María 

Alejandra Alvarez

Pharmacological
Properties of
Native Plants
from Argentina
Pharmacological Properties of Native Plants
from Argentina
María Alejandra Alvarez

Pharmacological Properties
of Native Plants from
Argentina
María Alejandra Alvarez
Department of Biotecnología Vegetal
CONICET/ CEBBAD
Universidad Maimónides
Ciudad de Buenos Aires, Argentina

ISBN 978-3-030-20197-5    ISBN 978-3-030-20198-2 (eBook)

https://doi.org/10.1007/978-3-030-20198-2

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Preface

There are a large number of native plants used by traditional medicine in Argentina.
The intention of this book is not to describe all the medicinal native species from
Argentina but those with pharmacological studies that validate their pharmacologi-
cal properties. To carry out this selection, the work by Barboza et al. (2009) was of
fundamental importance. The websites of the Darwinion Institute (www.darwin.
edu.ar) and Flora Argentina (www.floraargentina.edu.ar) have been consulted to
update the scientific names and corroborate the distribution data and botanical char-
acteristics found in other sources. The botanical terms used correspond to the glos-
sary compiled by P.P.J. Herman. Each chapter begins with a brief description of the
geographical, ecogeographical, or phytogeographical region in which the selected
species grow. It has not been the intention of this book to describe the medicinal
flora of each of these regions but to locate the selected species within the context of
one of the several regions where they could be found. For this reason, for each spe-
cies, all the provinces in which it is possible to find it are specified.
This work would not have been possible without the invaluable collaboration of Dr.
Chana Pilberg who gave me as a gift a large part of her botanical library; her friend-
ship is one of my greatest privileges. I must also thank my sister, Patricia G. Alvarez,
for generously giving me some of the photographs that illustrate this book. I also
thank all the colleagues who shared with me their publications and my colleagues
from the Pharmacy and Biochemistry School and CEBBAD at Maimónides University,
and Universidad de Buenos Aires for their constant support. Finally, I thank my family
and in particular my daughters for their patience and affectionate collaboration.

Buenos Aires, Argentina  María Alejandra Alvarez 

May 2019

References

Barboza GE, Cantero JJ, Núñez C, Pacciaroni A, Ariza Espinar L (2009) Medicinal plants: a gen-
eral review and a phytochemical and ethnopharmacological screening of the native Argentine
Flora. Kurtziana 34(12):7365

v
Book Introduction

The aim of this book is to offer information about native plants with pharmacologi-
cal properties from Argentina to students, researchers, and graduates interested in
the fields of Ethnobotany, Pharmacognosy, Phytochemistry, Pharmacy, and
Medicine. It describes some of the native species used in folk medicine whose phar-
macological activities have been experimentally tested. Chapter 1 summarizes the
characteristics of the geographical, ecological, and phytogeographical regions from
the country. Also, a brief reference about the legal regulatory framework of medici-
nal plants in Argentina is included. Chapter 2 reviews some aspects of ethnobotany
in Argentina. The following Chaps. 3, 4, 5, 6, 7, 8, and 9 describe native medicinal
plants in their botanical aspects, ethnomedicinal uses, chemical activity, and toxic-
ity and, when appropriate, in the establishment of in vitro cultures and their legal
status. Each of those chapters begins with a brief description of the region in which
the species grow. The following species are described: from Cuyo (Chap. 3), Aloysia
gratissima (Gillies & Hook. ex Hook.) Tronc., Lippia integrifolia (Griseb.) Hieron.,
Minthostachys mollis, Acantholippia seriphioides (A.  Gray) Moldenke, and
Achyrocline satureioides (Lam.); from the Pampa (Chap. 4), Erythrina crista-galli
L. var. crista-galli, Phytolacca dioica L., Salix humboldtiana Wild, Grindelia pul-
chella Dunal, Larrea cuneifolia Cav., Larrea divaricata Cav., and Larrea nitida
Cav; from Mesopotamia (Chap. 5), Cecropia pachystachya Trécul (Cecropiaceae),
Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul, Ilex para-
guariensis A.  St.-Hil var. paraguariensis, Tabebuia impetiginosa (Mart. ex DC.)
Standl., Allophylus edulis (A. St.-Hil., A. Juss., and Cambess.) Hieron. ex Niederl.,
Passiflora caerulea L., Blechnum occidentale L., and Maytenus ilicifolia Mart. ex
Reissek; from the Puna (Chap. 6), Acacia caven (Molina) Molina var. caven,
Chenopodium ambrosioides L., Anemia tomentosa (Savigny) Sw. var. anthriscifolia
(Schrad.) Mickel, Plantago australis Lam. subsp. Australis, and Alternanthera pun-
gens Kunth; from the Yungas (Chap. 7), Lepidium didymum L., Clinopodium gil-
liesii (Benth.) Kunze., Smilax campestris Griseb., and Smallanthus macroscyphus
(Baker ex Martius) A. Grau; and from Chaco and Espinal (Chap. 8), the medicinal
species Adiantum raddianum C. Presl., Aloysia citriodora Palau, Bauhinia forficata
subsp. pruinosa (Vogel) Fortunato & Wunderlin, Caesalpinia gilliesii var. gilliesii

vii
viii Book Introduction

(Wall. ex Hook.) D. Dietr., Pilocarpus pennatifolius Lemaire, and Jodina rhombifo-

lia (Hook. & Arn.) Reissek. Finally, from Patagonia Argentina, the species Adesmia
boronioides Hook f., Berberis microphylla G. Forest, Buddleja globosa Hope, and
Verbena litoralis Kunth var. litoralis are portrayed (Chap. 9).
Contents

1 Introduction: Native Plants of Argentina – A General Overview����������    1

1.1 Introduction����������������������������������������������������������������������������������������    1
1.2 República Argentina����������������������������������������������������������������������������    1
1.2.1 Geographical Regions������������������������������������������������������������    2
1.2.2 Eco-regions ����������������������������������������������������������������������������    3
1.2.3 Phytogeographical Regions����������������������������������������������������    6
1.3 Medicinal Plants����������������������������������������������������������������������������������   10
1.4 Native Medicinal Flora from Argentina����������������������������������������������   10
1.5 Regulatory Framework on Medicinal Plants in Argentina������������������   11
References����������������������������������������������������������������������������������������������������   12
2 Ethnobotany in Argentina������������������������������������������������������������������������   15
2.1 Introduction����������������������������������������������������������������������������������������   15
2.2 Ethnobotany����������������������������������������������������������������������������������������   15
2.3 Ethnobotany in Argentina ������������������������������������������������������������������   17
2.3.1 Ethnobotany in Indigenous and criollos Communities����������   19
2.3.2 Ethnobotany from Different Regions from Argentina������������   21
References����������������������������������������������������������������������������������������������������   26
3 Cuyo������������������������������������������������������������������������������������������������������������   33
3.1 Introduction����������������������������������������������������������������������������������������   33
3.1.1 The Cuyo Region��������������������������������������������������������������������   33
3.2 Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. gratissima (Verbenaceae)��������������������������������������������������   36
3.3 Lippia integrifolia (Griseb.) Hieron. (Verbenaceae) ��������������������������   39
3.4 Minthostachys mollis (Griseb.) Epling (Lamiaceae)��������������������������   41
3.5 Acantholippia seriphioides (A. Gray) Moldenke (Verbenaceae)��������   45
3.6 Achyrocline satureioides (Lam.) D.C. (Asteraceae) ��������������������������   47
References����������������������������������������������������������������������������������������������������   53

ix
x Contents

4 The Pampa: An Infinite Plain ������������������������������������������������������������������   63

4.1 Introduction: The Pampa��������������������������������������������������������������������   63
4.2 Erythrina crista-galli L. var. crista-galli (Fabaceae)��������������������������   65
4.3 Phytolacca dioica L. (Phytolaccaceae)����������������������������������������������   70
4.4 Salix humboldtiana Willd. var. humboldtiana (Salicaceae)����������������   73
4.5 Grindelia pulchella Dunal var. discoidea (Hook. & Arn.)
A. Bartoli & Tortosa (Asteraceae)������������������������������������������������������   75
4.6 Larrea spp. (Zygophyllaceae)������������������������������������������������������������   77
4.6.1 Larrea cuneifolia Cav.������������������������������������������������������������   78
4.6.2 Larrea divaricata Cav.������������������������������������������������������������   80
4.6.3 Larrea nitida Cav. ������������������������������������������������������������������   83
References����������������������������������������������������������������������������������������������������   84
5 Mesopotamia-Paranaense Forest��������������������������������������������������������������   91
5.1 Introduction: The Mesopotamia����������������������������������������������������������   91
5.2 The Paranaense Forest������������������������������������������������������������������������   92
5.3 The Jesuit Missions����������������������������������������������������������������������������   93
5.4 Cecropia pachystachya Trecul (Cecropiaceae)����������������������������������   95
5.5 Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.)
Altschul (Fabaceae)����������������������������������������������������������������������������   99
5.6 Ilex paraguariensis A. St.-Hil var. paraguariensis
(Aquifoliaceae) ����������������������������������������������������������������������������������  102
5.7 Tabebuia impetiginosa (Mart. ex DC.) Standl. (Bignoniaceae)����������  108
5.8 Allophylus edulis (A. St.-Hil., A. Juss. & Cambess.)
Hieron. ex Niederl. (Sapindaceae)������������������������������������������������������  111
5.9 Passiflora caerulea L. (Passifloraceae) ����������������������������������������������  114
5.10 Blechnum occidentale L. var. occidentale
(Blechnaceae, Pteridophyta) ��������������������������������������������������������������  118
5.11 Maytenus ilicifolia Mart. ex Reissek (Celesteraceae)������������������������  119
References����������������������������������������������������������������������������������������������������  125
6 Puna: A Surrealistic Landscape in the Argentinean Highlands������������  137
6.1 Introduction: The Puna�����������������������������������������������������������������������  137
6.2 Acacia caven (Molina) Molina var. caven (Fabaceae)������������������������  140
6.3 Chenopodium ambrosioides L. (Chenopodiaceae) ����������������������������  142
6.4 Anemia tomentosa (Savigny) Sw. var. anthriscifolia
(Schrad.) Mickel (Anemiaceae)����������������������������������������������������������  148
6.5 Plantago australis Lam. subsp. Australis (Plantaginaceae)����������������  150
6.6 Alternanthera pungens Kunth (Amaranthaceae)��������������������������������  154
References����������������������������������������������������������������������������������������������������  157
7 Yungas ��������������������������������������������������������������������������������������������������������  167
7.1 Introduction: Yungas, the Green Face of the Andes����������������������������  167
7.2 Lepidium didymum L. (Brassicaceae) ������������������������������������������������  169
7.3 Clinopodium gilliesii (Benth.) Kunze (Lamiaceae)����������������������������  171
7.4 Schinus molle L. Var. Molle (Anacardiaceae)������������������������������������  174
Contents xi

7.5 Smilax campestris Griseb. (Smilacaceae) ������������������������������������������  179

7.6 Smallanthus macroscyphus (Baker) A. Grau ex A.A. Sáenz
(Asteraceae)����������������������������������������������������������������������������������������  181
References����������������������������������������������������������������������������������������������������  184
8 Chaco and Espinal ������������������������������������������������������������������������������������  193
8.1 Introduction����������������������������������������������������������������������������������������  193
8.1.1 Chaco��������������������������������������������������������������������������������������  193
8.1.2 Espinal������������������������������������������������������������������������������������  195
8.2 Adiantum raddianum C. Presl (Pteridaceae)��������������������������������������  195
8.3 Aloysia citriodora Palau (Verbenaceae)����������������������������������������������  199
8.4 Bauhinia forficata subsp. pruinosa (Vogel) Fortunato
& Wunderlin (Fabaceae) ��������������������������������������������������������������������  203
8.5 Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.)
D. Dietr. (Fabaceae)����������������������������������������������������������������������������  208
8.6 Pilocarpus pennatifolius Lemaire (Rutaceae)������������������������������������  212
8.7 Jodina rhombifolia (Hook. & Arn.) Reissek (Cervantesiaceae) ��������  215
References����������������������������������������������������������������������������������������������������  218
9 Patagonia����������������������������������������������������������������������������������������������������  227
9.1 Introduction����������������������������������������������������������������������������������������  227
9.1.1 The Steppe and the Andine Forest������������������������������������������  227
9.2 Adesmia boronioides Hook. f. (Fabaceae)������������������������������������������  230
9.3 Berberis microphylla G. Forest (Berberidaceae)��������������������������������  232
9.4 Buddleja globosa Hope (Scrophulariaceae)����������������������������������������  236
9.4.1 Toxicity ����������������������������������������������������������������������������������  240
9.5 Verbena litoralis Kunth var. litoralis (Verbenaceae) ��������������������������  240
References����������������������������������������������������������������������������������������������������  242

Index������������������������������������������������������������������������������������������������������������������  249
About the Author

María  Alejandra  Alvarez  completed her studies at the Universidad de Buenos

Aires, Argentina. She graduated with degrees in Biochemistry and Pharmacy in
1981 and 1994, respectively, and obtained her MSc in Industrial Microbiology and
Biotechnology in 1986 and her PhD in Plant Biotechnology in 1993. She was lec-
turer in graduate and postgraduate courses of Botany, Pharmacobotany,
Biotechnology, Food Biotechnology, and Galenic Pharmacy at the Universidad de
Buenos Aires, Universidad Nacional de La Plata, and Universidad Maimónides.
She started working on the production of compounds of pharmaceutical interest
in in vitro plant cultures in 1986, being the author of numerous scientific articles.
Her research interests include chemotaxonomy, medicinal plants, production of sec-
ondary metabolites, and molecular farming. She was a researcher at the Universidad
de Buenos Aires and Centro de Ciencia y Tecnología Dr. César Milstein (CONICET/
Fundación Pablo Cassará). Currently, she is a member of the National Council of
Research and Technology (CONICET) in Argentina, professor of Pharmacobotany
and Pharmacognosy and director of the Plant Biotechnology Group at the
Universidad Maimónides, and professor of Biology at Colegio Divino Corazón. She
was the editor of the book Genetic Transformation (2011), Intech Open Access
ISBN: 978-­953-­307-364-4, and the author of the book Plant Biotechnology for
Health: From Secondary Metabolites to Molecular Farming (2014), Springer
International Publishing AG, ISBN 978-3-319-05770-5.

xiii
Abbreviations

2,4-D 2,4-dichlorophenoxyacetic acid

2iP 2 inositol phosphate N6-(2-isopentenyl) adenine
ALT Alanine aminotransferase
ANMAT National Administration of Drugs, Foods and Medical Technology
AST Aspartate aminotransferase
BAP 6-benzilaminopurine
DPPH (2,2-difenil-1,2-picrilhidrazil)
DW Dry weight
EOs Essential oils
FNA Argentina National Pharmacopeia
FW Fresh weight
IAA Indole-3-acetic acid
IBA Indole-3-butyric acid
Kin Kinetin
m.a.s.l. Meters above sea level
m.b.s.l. Meters below sea level
MBC Minimum bactericidal concentration
MIC Minimum inhibitory concentration
MPs Medicinal plants
MS Murashige and Skoog medium
MSG MS medium with Gamborg’s vitamins
MSRT MS medium with Khanna and Staba vitamins
NAA 1-Naphtalene acetic acid
PIC Picloram
ROS Reactive oxygen species
TDZ Thidiazuron

xv
Argentina Provinces

BAI Buenos Aires

CAT Catamarca
CHA Chaco
CHU Chubut
COR Córdoba
COS Corrientes
DFE Distrito Federal
ERI Entre Ríos
FOR Formosa
JUJ Jujuy
LPA La Pampa
LRI La Rioja
MEN Mendoza
MIS Misiones
NEU Neuquén
RNE Río Negro
SAL Salta
SCR Santa Cruz
SDE Santiago del Estero
SFE Santa Fe
SJU San Juan
SLU San Luis
TDF Tierra del Fuego
TUC Tucumán

xvii
List of Figures

Fig. 1.1 Phytogeographical regions from Argentina.

(Adapted from Cabrera (1971))��������������������������������������������������������    7
Fig. 3.1 The Cuyo region. Landscapes from Mendoza
(a), San Juan (b), and San Luis (c) provinces����������������������������������   35
Fig. 3.2 Some chemicals found in essential oils (EOs) from aerial
parts of Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. gratissima������������������������������������������������������������������������   38
Fig. 3.3 Some chemicals found in Lippia integrifolia (Griseb.) Hieron��������   40
Fig. 3.4 Some of the chemical compounds found
in Minthostachys verticillata (Griseb)����������������������������������������������   43
Fig. 3.5 Some of the chemical compounds found in the essential
oil from Acantholippia seriphioides (A. Gray) Moldenke����������������   46
Fig. 3.6 Some polyphenols and flavonoids detected
in Achyrocline satureioides (Lam.) DC��������������������������������������������   48
Fig. 3.7 Main components of the essential oils from Achyrocline
satureioides (Lam.) DC��������������������������������������������������������������������   49
Fig. 4.1 A landscape from the Pampa plain. PG Alvarez
personal collection����������������������������������������������������������������������������   64
Fig. 4.2 Plant communities from the Pampa eco-region.
PG Alvarez personal collection��������������������������������������������������������   65
Fig. 4.3 Erythrina crista-galli L. var. crista-galli (Fabaceae).
Branches with leaves and flowers (a), detail of the flowers
(b), fruits (c), flowers and fruits (d). MA Alvarez
personal collection����������������������������������������������������������������������������   66
Fig. 4.4 Some isoquinoline alkaloids from Erythrina crista-galli
L. var. crista-galli������������������������������������������������������������������������������   67
Fig. 4.5 Some compounds found in the Erythrina crista-galli
L. var. crista-galli essential oils��������������������������������������������������������   68
Fig. 4.6 Other chemical compounds found in leaves and flowers
of Erythrina crista-galli L. var. crista-galli��������������������������������������   68

xix
xx List of Figures

Fig. 4.7 Phytolacca dioica L., Phytolaccaceae. Tree (a), leaves

and fruits (b), pendulous flower clusters (c). MA Alvarez
personal collection����������������������������������������������������������������������������   71
Fig. 4.8 Ombuoside from Phytolacca dioica L. leaves����������������������������������   72
Fig. 4.9 Salix humboldtiana Willd. var. humboldtiana.
Tree (a) and leaves (b). MA Alvarez personal collection ����������������   74
Fig. 4.10 Chemical structure of the alkaloid salicine
from Salix humboldtiana Willd. var. humboldtiana��������������������������   75
Fig. 4.11 Chemical compounds from Grindelia pulchella Dunal var.
discoidea (Hook. & Arn.) A. Bartoli & Tortosa essential oils����������   76
Fig. 4.12 Grindelic acid and labdane from Grindelia pulchella Dunal var.
discoidea (Hook. & Arn.) A. Bartoli & Tortosa��������������������������������   77
Fig. 4.13 Some of the flavonoids found in Larrea divaricata Cav. leaves������   79
Fig. 4.14 Chemical compounds from Larrea divaricata Cav. ������������������������   81
Fig. 4.15 Chemical compounds from Larrea nitida Cav. aerial parts��������������   84
Fig. 5.1 The Iguazú Falls. Province of Misiones, Argentina.
MA Alvarez personal collection ������������������������������������������������������   92
Fig. 5.2 San Ignacio Miní Mission, Misiones, Argentina.
MA Alvarez personal collection ������������������������������������������������������   94
Fig. 5.3 Cecropia pachystachya Trecul (Cecropiaceae). Parque Nacional
Iguazú, Misiones, Argentina. MA Alvarez personal collection��������   96
Fig. 5.4 Some chemical compounds isolated from Cecropia pachystachya
Trecul������������������������������������������������������������������������������������������������   97
Fig. 5.5 Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.)
Altschul (Fabaceae). Parque Nacional Iguazú, Misiones,
Argentina. MA Alvarez personal collection�������������������������������������  100
Fig. 5.6 Chemical compounds from Anadenanthera colubrina (Vell.)
Brenan var. cebil (Griseb.) Altschul��������������������������������������������������  101
Fig. 5.7 Indole alkaloids from Anadenanthera colubrina (Vell.)
Brenan var. cebil (Griseb.) Altschul��������������������������������������������������  102
Fig. 5.8 Ilex paraguariensis A. St.-Hil var. paraguariensis
(Aquifoliaceae). San Ignacio Miní mission, Misiones,
Argentina (a and b), and in Buenos Aires Botanical Garden (c).
MA Alvarez personal collection ������������������������������������������������������  104
Fig. 5.9 Some chemical compounds found in Ilex paraguariensis
A. St.-Hil var. paraguariensis. (Filip et al. 1998)������������������������������  105
Fig. 5.10 Chemical structure of lapachol and other quinones found
in Tabebuia impetiginosa (Mart. ex DC.) Standl������������������������������  109
Fig. 5.11 Chemicals found on the dried inner bark of Tabebuia
impetiginosa (Mart. ex DC.) Standl��������������������������������������������������  110
Fig. 5.12 Chemical compounds found in Allophylus edulis
(A. St.-Hil., A. Juss. & Cambess.) Hieron. ex Niederl ��������������������  113
Fig. 5.13 Passiflora caerulea L. flower. MA Alvarez personal collection ������  115
List of Figures xxi

Fig. 5.14 Some of the chemical compounds determined

in Passiflora caerulea L. ������������������������������������������������������������������  117
Fig. 5.15 Chemicals found in Blechnum occidentale L. var. occidentale��������  120
Fig. 5.16 Maytenus ilicifolia Mart. ex Reissek, Celesteraceae.
Buenos Aires, Botanical Garden. MA A Personal collection ����������  121
Fig. 5.17 Some terpenoids from Maytenus ilicifolia
Mart. ex Reissek leaves��������������������������������������������������������������������  123
Fig. 5.18 Maytansinoid alkaloids from Maytenus ilicifolia
Mart. ex Reissek��������������������������������������������������������������������������������  124
Fig. 6.1 Puna, section in the province of Jujuy, Argentina.
Personal collection P G Alvarez�������������������������������������������������������  138
Fig. 6.2 Salinas grandes (a) and llama (Lama lama) in the province
of Jujuy (b), Argentina. Personal collection PG Alvarez������������������  139
Fig. 6.3 Acacia caven (Molina) Molina var. caven (Fabaceae)
tree (a), fruits and leaves (b), A. caven parasited
by Ligaria cuneifolia (c). MA Alvarez personal collection��������������  141
Fig. 6.4 Chemicals from Acacia caven (Molina) Molina var. caven��������������  142
Fig. 6.5 Chenopodium ambrosioides L. leaves and inflorescences.
MA Alvarez personal collection ������������������������������������������������������  143
Fig. 6.6 Chemicals found in Chenopodium ambrosioides
L. essential oil ����������������������������������������������������������������������������������  144
Fig. 6.7 Some triquinane sesquiterpenes found in Anemia
tomentosa (Savigny) Sw. var. anthriscifolia (Schrad.) Mickel����������  149
Fig. 6.8 Chemical compounds found in Anemia tomentosa
(Savigny) Sw. var. anthriscifolia (Schrad.) Mickel essential oils�����  150
Fig. 6.9 Chemical compounds from Plantago australis
Lam. subsp. Australis (Plantaginaceae)��������������������������������������������  152
Fig. 6.10 Chemical compounds found in Althernanthera pungens
Kunth (Amaranthaceae)��������������������������������������������������������������������  156
Fig. 7.1 Yungas, province of Jujuy, Argentina. (Personal collection
PG Alvarez)��������������������������������������������������������������������������������������  168
Fig. 7.2 Chemical compounds from Lepidium didymum L.��������������������������  171
Fig. 7.3 Chemical compounds from Clinopodium gilliesii
(Benth.) Kunze����������������������������������������������������������������������������������  173
Fig. 7.4 Chemical compounds from Schinus molle L. var. molle������������������  176
Fig. 7.5 Chemical compounds from Smilax campestris Griseb ��������������������  180
Fig. 7.6 Chemical compounds from Smallanthus macroscyphus
(Baker) A. Grau ex A.A. Sáenz��������������������������������������������������������  183
Fig. 8.1 Chaco (a) and Espinal (b) eco-regions from Argentina.
MA Alvarez personal collection ������������������������������������������������������  194
Fig. 8.2 Adiantum raddianum C. Presl (Pteridaceae). Buenos Aires
Botanical Garden. M A Alvarez personal collection������������������������  197
xxii List of Figures

Fig. 8.3 Chemical compounds found in Adiantum raddianum

C. Presl spores����������������������������������������������������������������������������������  197
Fig. 8.4 Some chemical compounds found in Adiantum
raddianum C. Presl ��������������������������������������������������������������������������  198
Fig. 8.5 Chemical compounds from Aloysia citriodora
Palau (Verbenaceae)��������������������������������������������������������������������������  200
Fig. 8.6 Bauhinia forficata subsp. pruinosa (Vogel) Fortunato & Wunderlin
(Fabaceae). Flower (a) personal collection M C Fernández
and leaves (b) MA Alvarez personal collection��������������������������������  204
Fig. 8.7 Chemical compounds of Bauhinia forficata subsp. pruinosa
(Vogel) Fortunato & Wunderlin (Fabaceae)��������������������������������������  205
Fig. 8.8 Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.) D. Dietr.
(Fabaceae). Buenos Aires Botanical Garden. MA Alvarez
personal collection����������������������������������������������������������������������������  209
Fig. 8.9 Some chemical compounds from Caesalpinia gilliesii var.
gilliesii (Wall. ex Hook.) D. Dietr. (Fabaceae) ��������������������������������  211
Fig. 8.10 Chemical compounds from Pilocarpus pennatifolius Lemaire
(Rutaceae) ����������������������������������������������������������������������������������������  213
Fig. 8.11 Jodina rhombifolia (Hook. & Arn.) Reissek
(Cervantesiaceae). Buenos Aires Botanical Garden.
MA Alvarez personal collection ������������������������������������������������������  215
Fig. 8.12 Chemical compounds found in Jodina rhombifolia
(Hook. & Arn.) Reissek, (Cervantesiaceae) aerial parts ������������������  217
Fig. 9.1 Patagonia Argentina. (a) Ushuaia, Vespignani glacier
viewpoint, (b) Patagonian steppe, (c) Perito Moreno Glacier,
(d) MA Alvarez personal collection��������������������������������������������������  228
Fig. 9.2 Chemical compounds from Adesmia boronioides Hook. f.��������������  231
Fig. 9.3 Berberis microphylla G. Forest (Berberidaceae) branches
with leaves and fruits. MA Alvarez personal collection ������������������  233
Fig. 9.4 Chemical compounds from Berberis microphylla G. Forest
(Berberidaceae) ��������������������������������������������������������������������������������  234
Fig. 9.5 Buddleja globosa Hope (Buddlejaceae). MA Alvarez
personal collection����������������������������������������������������������������������������  237
Fig. 9.6 Some chemical compound found in Buddleja globosa
Hope (Scrophulariaceae)������������������������������������������������������������������  238
Fig. 9.7 Chemical compounds from Verbena litoralis
Kunth var. litoralis����������������������������������������������������������������������������  241
List of Tables

Table 2.1 Ethnobotanical studies related to ethnic groups

from Argentina (indigenous and criollos)����������������������������������������  20
Table 2.2 Ethnobotanical research in different regions of Argentina ��������������  22

xxiii
Chapter 1
Introduction: Native Plants of Argentina –
A General Overview

1.1  Introduction

Plants are the perfect machinery that transforms light into chemical energy and
produce also oxygen, a vital element for most of the organisms on Earth. They are
a source of food, fodder fuel, cellulose, paper, timber, textile fibers, essences and
flavors, and medicines. They are also natural pesticides, are helpful to mitigate envi-
ronmental pollution, and have a vital role to fight climate change. Plants are also
central to scientific research on basic aspects of plant life and on challenges related
to agriculture, health, and environment. Besides, plants have been part of ritual,
religious, and festive practices.

1.2  República Argentina

Argentina is located in the Southern tip of South America. It shares borders with
Bolivia and Paraguay to the North; Chile to the West; Brazil, Uruguay, and the
South Atlantic Ocean to the East; and Chile and the South Atlantic Ocean to the
South. Their endpoints are North, 21° 46’ S, 66° 13’ W.; South, 55° 03’ S, 66° 31’
W; East, 26° 15’ S, 53° 38’ W; and West, 50° 01’S, 73° 34’ W.  The Antarctica
Argentina is located between the meridians 25° W and 74° W and the parallels 60°
S and 90° S (South Pole). The country has a total surface area of 3.761.274 km2,
2.791.810 km2 in the American continent and 969.464 km2 in the Antarctica. The
population is around 40.117.096 inhabitants with a population density of 10.7
inhabitants/km2 (Instituto Geográfico Nacional República Argentina). The country
is composed of 23 provinces and the Autonomous City of Buenos Aires which is the
capital city of the country. It is also divided in seven geographical regions: Northwest,
Mesopotamia, the Gran Chaco, the Sierras Pampeanas, Cuyo, Región Pampeana,
and Patagonia. The country has various ecosystems with a substantial biodiversity
due to its large size, geographic variety, and diverse climate types.

© Springer Nature Switzerland AG 2019 1

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_1
2 1  Introduction: Native Plants of Argentina – A General Overview

1.2.1  Geographical Regions

The geographical regions from Argentina are the Northwest region, Mesopotamia,
the Gran Chaco, Cuyo, the Pampas, Patagonia, and South Atlantic Islands and
Antarctica.
The Northwest region includes the provinces of Catamarca (CAT), Jujuy (JUJ),
La Rioja (LRI), Salta (SAL), Santiago del Estero (SDE), and Tucumán (TUC). The
climate is varied, with rainfall diminishing from North to South and from East to
West. It comprises two subregions, the Puna and the Yungas. The Puna, to the high
Andean west, is dry and with a great temperature oscillation, mostly cold, frequently
decreasing below freezing point at night. The Yungas, to the East, is tropical, very
hot, and humid.
Mesopotamia includes the provinces of Misiones (MIS), Entre Ríos (ERI), and
Corrientes (COS). The climate is mostly subtropical, with a hot and very humid
tropical climate to the North, gradually becoming temperate and semi-humid to the
South.
The Gran Chaco includes the provinces of Chaco (CHA), Formosa (FOR), and
Santiago del Estero (SDE). The region has a very hot subtropical to tropical climate,
characterized by humid summers and mild dried winters. It has heavy seasonal rain-
falls and periodic droughts.
Cuyo includes areas from the provinces of San Juan (SJU), San Luis (SLU),
and Mendoza (MEN). It has an arid or semiarid climate with an average annual
precipitation of about 100 to 500 millimeters (4 to 20 in). The diurnal temperature
range is very large with extremely hot temperatures during the day followed by
cold nights.
The Región Pampeana could be divided into Llanura pampeana (Pampa’s
plains) and the Sierras Pampeanas (Pampa’s hills). The Llanura pampeana covers
the provinces of Buenos Aires (BAI), La Pampa (LPA), Córdoba (COR), Santa Fe
(SFE), and Entre Ríos (ERI). The climate is temperate, with hot, stormy summers,
and cool winters. The Sierras Pampeanas comprises sectors from the provinces of
San Luis (SLU), San Juan (SJU), Córdoba (COR), La Rioja (LRI), Catamarca
(CAT), Santiago del Estero (SDE), and Tucumán (TUC). They have a temperate
and semiarid climate, with warm summers and cool winters. The Northern-East
slope is covered by rainforest, generating high humidity. The area inside COR and
SLU has a Mediterranean-type climate, with intense summer rainstorms and
snowy winters. The eastern slopes that receive the moist winds from the Atlantic
Ocean have more rainfalls.
Patagonia includes the provinces of Neuquén (NEU), Río Negro (RNE), Chubut
(CHU), Santa Cruz (SCR), and Tierra del Fuego (TDF). The region is very windy,
with mild summers, cold to very cold winters, and heavy snowfall and frost, espe-
cially in mountainous zones. Precipitation steeply diminishes from west to east.
South Atlantic Islands and Antártida Argentina. Argentina is a founding signa-
tory member of the Antarctic Treaty and is also a permanent consulting member.
There are 13 Argentinean bases (6 permanent and 7 seasonal) in Antarctica with
1.2 República Argentina 3

activities coordinated by the Instituto Antártico Argentino and the Dirección

Nacional del Antártico. There the temperatures range from 0 °C in the summer to
−60 °C to −80 °C in winter.

1.2.2  Eco-regions

An eco-region is defined as a relatively large geographical area characterized by a

unique geology, weather, soil, hydrology, and plant and animal species. Argentina is
divided into eighteen eco-regions, fifteen of them are continental, two marine, and
one Antarctic. The eco-regions are Altos Andes (High Andean Complex), Puna
(Puna grassland), Monte de sierras y bolsones (Argentine Northwest Monte and
Thistle of the Prepuna), Selva de Yungas (Yungas), Chaco seco (Arid Chaco), Chaco
húmedo (Humid Chaco), Bosques patagónicos (Patagonia forests), pastizales y
matorrales (grasslands and shrublands), Delta e Islas del Paraná (Paraná flooded
savanna), Espinal (Espinal), Estepa patagónica (Patagonian steppe), Esteros del
Iberá (Iberá Wetlands), Monte de llanuras y mesetas (Argentina Low Monte),
Pampas (Pampa), Selva Paranaense (Alto Paraná Atlantic forests), Islas del
Atlántico Sur (South Atlantic islands), Mar Argentino (Argentina sea), and Antártida
(Antarctic Peninsula) (Burkart et al. 1999; Diminich et al. 2018).
The High Andean Complex or Altos Andes is the high mountain sector in the
West of Argentina, from the border with Bolivia to the North of the province of
NEU, across the provinces of JUJ, SAL, TUC, CAT, LRI, SJU, and MEN. To the
North it includes the summits and slopes of the Eastern Andes mountain range.
Towards the South it includes some sections of the Andes foothills (Matteucci
2012). The climate is cold with everlasting snow. The rains are around 100 to
200 mm. Vegetation is grassy or shrubby, highly adapted to the extreme weather
and altitude.
The Puna grassland or Puna extends from the Northwestern border of the coun-
try (province of Jujuy) to the North of the province of SJU. It has a cold desert cli-
mate characterized by cushion bog vegetation. Its plateau is crossed by mountains
with numerous volcanoes.
Argentine Northwest Monte and Thistle of the Prepuna extends from JUJ to the
North of MEN. It has extensive and pronounced slopes that connect with the Puna
to the North and with the High Andes to the South. At the foothills are the Quebrada
de Humahuaca, the Quebrada del Toro, the Calchaquí Valleys, and the rivers Jáchal
and Bermejo. The climate is subtropical dry to the North; in their valleys and can-
yons, the rains concentrated in the summer season, while in the South, rains are
distributed throughout the year. The radiation is intense, and the cloudiness is low.
Temperature changes are very marked, both during the day and between seasons.
Soil is predominantly sandy and poor in organic matter.
The Yungas are located to the north of the country with a discontinuous distribu-
tion in the provinces of SAL, JUJ, TUC, and CAT. The climate is warm and humid
4 1  Introduction: Native Plants of Argentina – A General Overview

to sub-humid. The conditions of temperature and humidity are variable according to

altitude and latitude. The region has a high biodiversity with evergreen forests.
Arid Chaco extends over the western half of FOR and CHA; the eastern half of
SAL; almost all of SDE, North of SFE, and COR; and sectors of CAT, LRI, and
SLU. The climate is continental and subtropical, with areas that attain the maximal
absolute temperature of the country. The type of vegetation is the xerophilous forest,
whose trees become lower and narrower towards the southwest. There are abundant
mountain forests, savannas, and grasslands.
Humid Chaco occupies the eastern half of FOR and CHA and the extreme north-
west of COS and North of SFE.  The climate is subtropical warm. The average
annual temperature decreases from North to South and from 23 °C in the border
with Paraguay to about 18 °C in the center of the Province of SFE. The vegetation
has greater diversity than in the Arid Chaco.
Patagonia forest is the driest part of the Andean-Patagonic forest; it extends from
NEU to TDF. The landscape is mountainous, with abrupt and steep reliefs, snowy
peaks, glacial valleys, volcanic geo-forms, numerous water courses, and lakes that
drain through different rivers to the slopes of the Atlantic or the Pacific oceans. The
climate is temperate to cold and humid, with heavy snowfall, rains, strong winds,
and frost most of the year. The dominant plant formation is the humid, semidry
temperate forest.
Grassland and shrublands are located in South MIS and Northeast COS being
mostly pastures and grasslands corresponding to the phytogeographical Paraná
flooded savanna. The dominating flora is herbaceous.
Paraná flooded savanna borders the southern Paraná River, from CHA to BAI. It
runs through the Humid Chaco and the Humid Pampas eco-regions, ending up in the
Río de la Plata estuary. It has a landscape of low and floodable islands, delimited by
the lateral arms and main channels of rivers (e.g., Paraná Guazú, Paraná de las
Palmas, Paraná Miní, Barca Grande, Luján) and extensive coastal waters. From the
original habitat, only patches including strips of forest and shrubs, scrub and pas-
ture, and hydrophilic and aquatic flora still remain.
Espinal is located in central Argentina involving the south of the Province of
COS, northern half of ERI, a central belt of SFE and COR, central and southern
SLU, and half East of LPA and south of BAI. The predominant landscape is flat
plain to gently undulating, occupied by low forests, savannahs, and grasslands,
nowadays largely adapted to agriculture.
The Patagonian steppe is around 730.000 Km2 extended from southwest MEN,
center of NEU, southwest of RNE, and most of CHU and SCR. The soils are rocky
and sandy, lacking rich organic materials. The weather is temperate-cold, with snow
in winter and frosts almost at any time of the year. Precipitations are scarce, ranging
from 100 to 300 mm per year, and are concentrated on the coldest months, from
April to September. In spring and summer, strong West winds blow almost con-
stantly. There are low shrubs with small leaves and thorns and grasslands.
Iberá Wetlands are a deposit of stagnant water that occupies the northern center
of the province of COS. It is one of the most important fresh water reservoirs of the
continent. The wetlands, with a deep no higher than 3 m, are covered by aquatic
1.2 República Argentina 5

plants (aguapés, poppies, and sequins of water) forming floating islands with up to
2 m of thickness. It is protected in the Iberá Provincial Reserve.
The Argentina Low Monte is the driest region of the country located in North-­
central Argentina covering the eastern foothills of the Andes. It extends from the
Andean mountain range in MEN, along NEU and LPA, to the coast of the Atlantic
Ocean of RNE and northeast of CHU. It shares with the eco-region of the Monte de
Sierras and Bolsones the most arid characteristics of Argentina but differs on its
prevailing landscape of plains and extensive plateaus. The climate is temperate-arid
with scarce precipitations. Average annual temperatures are on the order of
10–14 °C. The vegetation is poorer than in the Mount of Sierras and Bolsones.
The Pampa represents around 60% of the grasslands of the country; it is extended
from BAI (except the southern section), northeast of LPA, and south of COR, SFE,
and ERI.  The rains are distributed during the year. Average annual temperatures
range from 15 °C in the South to about 18 ° in the North.
The Alto Paraná Atlantic forests extend from the center and North of the prov-
ince of MIS through ERI as a streamside zone of narrow hills by the rivers from the
basin of the Río de la Plata. The weather is warm and humid, with more intense
rains in the summer. The average annual temperature is on the order of 20 °C. The
red soils, characteristic of the eco-region, are a consequence of the transformation
of the basaltic material under warm and humid conditions. It has 2000 vascular
plants, and the Argentinean sector exhibits the highest biological diversity of all the
country eco-regions. The dominant vegetation is the subtropical forest.
In the South Atlantic islands, due to the extreme climatic conditions and the
rocky nature of the terrain, the soils are poorly developed. There are no natural for-
ests, and the dominant vegetation is steppe grasses with tundra bushes. This eco-­
region is remarkably poor in phanerogams, several of which are endemic. Among
the most prominent plant communities are the Tussock grass pastures, which occupy
the coastal areas of Malvinas and South Georgia Islands, with different species of
grasses and mosses, among other plants.
Mar Argentino includes the Argentina Continental Platform which can be subdi-
vided into a coastal subregion, represented by the strip of coast up to 40 m depth and
the subregion of the external platform, which extends from the depths of 40 m to
200 m. On the coasts of CHU, SCR, and TDF, marine vegetation develops with a
huge diversity of green, red, and brown algae on the seabed.
The Antarctic Peninsula has an extremely cold climate, with average tempera-
tures below zero and with snow and ice all year round. Two types of climate are
distinguished: the glacial one, dominant in the sectors covered by ice, and the insu-
lar one that includes the Antarctic Peninsula and adjacent islands. Winds are very
strong; temperature is always very low, reaching – 42 °C on polar night and 0 °C in
the hottest months. The flora is very scarce and limited to sectors near the sea, which
lose the snow cover during the brief Antarctica summer. There are lichens or moss
cushions. The only native vascular plant species are Colobanthus quitensis (with
tiny white flowers) and Deschampsia antarctica (Antarctic grass) usually among
moss communities. A nitrophilous alga (Prasiola crispa) grows associated with
penguin breeding ground. There are two introduced species (Poa spp.), which grow
near to the permanent bases.
6 1  Introduction: Native Plants of Argentina – A General Overview

1.2.3  Phytogeographical Regions

Phytogeography can be defined as the geographical distribution of plants according

to the temperature, humidity, light intensity, day length, soil composition, and bio-
logical communities, among other characteristics, of each region. With the excep-
tion of some species that grow in very hostile environments, plants usually grow
with other individuals, from the same or other species, establishing plant communi-
ties or associations with all their members having similar ecological needs. There
have been several attempts to organize the Argentine vegetation in phytogeographi-
cal regions, as is shown in the works of Lorentz (1876), Holmberg (1898), Hauman
(1920, 1931), Frenguelli (1941), Castellanos and Pérez Moreau (1941, 1945),
Cabrera (1953, 1958, 1971), and Cabrera and Willink (1980).
Cabrera (1971) divided Argentina into three phytogeographical regions:
Neotropical, Antarctic, and Oceanic. Each region is categorized in domains, which
are divided in provinces that in turn are subdivided into districts. For each domain
Cabrera specifies its geographical location and for each province the location and
characteristic vegetation. The Neotropical region was divided in the Amazonian
(subdivided in the Yungas and Paranaense provinces), Chaqueño (subdivided in the
Chaqueña, Espinal, Prepuna, Monte, and Pampeana provinces), and Andino
Patagónico (subdivided in Altoandina, Puneña, and Patagónica provinces) domain.
The Antarctic region was subdivided into the sub-Antarctic (subdivided in the sub-­
Antarctic and insular provinces) and the Antarctic (Antarctic province) domain and
the Oceanic region without subdivisions (Cabrera 1971) (Fig. 1.1).

1.2.3.1  Neotropical Region

Amazonian domain: Despite its small area, the Amazonian domain is the largest
floristic wealth in Argentina.
Yungas province: it has many genus and species in common with the Paranaense
Province, such as Nectandra, Cedrela, Blepharocalyx, Jacaranda sp., etc. However,
unlike the Paranaense province, it has few bamboos and no palm trees. It presents
exclusive genera such as Tipuana, Juglans, Phoebe, and Myroxylon.
Paranaense province: more than 200 arboreal species have been counted in the
Misiones Forest. In the district of the Mixed Jungle are arboreal communities of
laurel forest (Nectandra saligna), Guatambú (Balfourodendron riedelianum),
palo rosa (Aspidosperma polyneuron), and palmito (Euterpe edulis). Also, there
are arboreal species such as anchico colorado (Parapiptadenia rigida), cancha-
rana (Cabialea oblongifolia), and cocó (Allophylus edulis) among others. The
pindó (Syagrus romanzoffiana) palm are scattered and distributed into the forest.
In the northeastern sector of the MIS jungle, there are Paraná pine (Araucaria
angustifolia) communities.
1.2 República Argentina 7

Fig. 1.1  Phytogeographical regions from Argentina. (Adapted from Cabrera (1971))
8 1  Introduction: Native Plants of Argentina – A General Overview

Chaqueño domain: it has polymorphic vegetation with deciduous xerophilous

forests, steppe shrubs, palms, savannas, meadows, steppes, grasslands, etc. The
weather is varied, with predominance of the continental type with moderate to
scarce rainfall, moderate winters, and warm summers.
Chaqueña Province: in this province the predominant species are from Schinopsis,
Aspidosperma, and Prosopis genus.
Province of Espinal: this province has connections to Chaqueña province, but in
Espinal species from the genus Prosopis predominate.
Prepuneña province: it has few tree species, with prevalence of cacti,
Zygophyllaceae spp., and shrubs.
Monte province: it mainly has trees from the genus Prosopis, dwarf arboreal spe-
cies, and prevalence of Zygophyllaceae shrubs (Larrea spp.).
Pampeana Province: here predominate xerophilous grasses and species from the
genus Stipa, Piptochaetium, Andropogon, Leonurus, etc.
Andino Patagónico domain: its climate is cold and dry, with frost almost all year
round and snow in the winter months.
Altoandino province: it has a rocky, stony, and sandy soil. It has shrub and grass
steppes with supremacy of xerophilous grasses and creeping or cushioned dicotyle-
dons. The endemic genera are numerous (Compositae, Cruciferous, Verbenaceae,
Poaceae, Papilionaceae, and Solanaceae).
Puneña province: it has a vegetation of steppe, grasslands, meadows, and bushes.
There are some shrubby communities such as Adesmia sp., Azorella yareta (yareta),
Stipa sp., Festuca sp., Fabiana densa, Baccharis boliviensis, and Opuntia sp.,
among others.
Patagonia province: depending on the characteristics of soil and water availabil-
ity, there are shrubby, halophytic, or grass steppe and, near small streams, fertile
valleys (vegas). Some typical species are Festuca pallescens, F. gracillima, Mulinum
spinosum, Trevoa patagonica, Chuquiraga avellanedae, Nassauvia glomerulosa,
and Junellia tridens.

1.2.3.2  Antarctic Region

The only native species from the region are Deschampsia antarctica and Colobanthus
quitensis. Also, there are several species of lichens, mosses, and algae.
Sub-Antarctic domain.
Sub-Antarctic province: The dominant vegetation is the deciduous and evergreen
forest. Other types of vegetation are scrubs, meadows, tundra, etc.
Insular province: the Malvinas Islands have a poor phanerogam flora that is still
poorer in the South Georgia. The dominant species is the tussock grass (Poa flabel-
lata). There is no community of shrubs.
Antarctic domain: Antarctic province—the seaweed Prasiola crispa forms green
wavy, snow-free emerald mats. There are also mosses and lichens grass.
1.2 República Argentina 9

1.2.3.3  Oceanic Region

It is the most extensive region that covers all the oceans. In Argentina, it extends
through the epicontinental sea, from the mouth of the Río de la Plata to the
Antarctica.
The Cabrera classification of the phytogeographical regions is still the most pop-
ular. However, there are other classifications that should be considered.
For Takhtajan (1986), the terrestrial flora of Argentina could be distributed in
three phytogeographical regions, two within the Neotropical (Brazilian and
Andean) and one in the Holantarctic (Patagonian-Chilean) kingdoms. According
to Takhtajan, the region and kingdom categories are equivalent to the domain and
region categories defined by Cabrera (Matteucci et al. 2016). In a detailed work,
for each of the phytogeographical regions defined by Cabrera, Apodaca et  al.
(2015) established the eco-regions involved, its dominant vegetation, area (in km2),
general weather, annual rain (expressed in mm), average annual temperature
(expressed in °C), and species, genus, and/or families from its endemic and char-
acteristic plant species. They also included a concise description of international,
national, and provincial protected areas. They recognized three main biomes in
Argentina: steppe (that include seven of the phytogeographic provinces), wood-
land (Chaco, Espinal, sub-­Antarctic), and forest. In the steppe predominate the
Gramineae; according to the accompanying species, the steppe could be shrubby,
herbaceous, etc. In the woodlands prevail trees, and in the forest prevail tall trees
along with copious lianas and epiphytes and stratified vegetation (Paranaense,
Yungas) (Apodaca et al. 2015).
Ribichich (2002) disclosed weaknesses and discrepancies in the work of
Cabrera (1971) and Cabrera and Willink (1980), particularly related to the impre-
cise use of the terms endemic and endemism, the irregular use of the taxonomic
categories to classify phytogeographical areas in different hierarchical levels, the
borderlines of the Argentinean provinces and geographical regions, and the ambi-
guities or incoherencies related to the characterization of the phytogeographical
territories. Besides, Ribichich compares the phytogeographical systems from
Cabrera and Takhtajan.
Recently, Oyarzábal et  al. (2018) published a physiognomic-floristic map of
spontaneous vegetation of Argentina based in vegetation surveys published in the
last decades. For each phytogeographic province identified by Cabrera (1976),
they reviewed the charts and available descriptions and mapped their internal het-
erogeneity. However, they made two modifications on Cabrera classification; on
the one hand, they incorporated the documented ecotone Monte-Patagonia (León
et al. 1998; Cingolani et al. 2000). On the other hand, they established slightly
different phytogeographic limits (Cabrera 1976; León et al. 1998). The resulting
map had 50 vegetation units as subdivisions of the phytogeographic ecotone and
provinces described by Cabrera and a brief physiognomic-floristic description of
each ecotone, province, and vegetation unit. A nomenclature of each vegetation
unit based on dominant type of spontaneous vegetation and characteristic species
was also proposed.
10 1  Introduction: Native Plants of Argentina – A General Overview

1.3  Medicinal Plants

The use of medicinal plants by early civilizations has been recorded for as old as
5000 years ago as is shown in Sumerian clay slabs containing the recipes for drug
preparations (Sumner 2000), the clay tablets from ancient Mesopotamia dated back
to around 2600 BC (Scurlock 2014), the registers from the Chinese emperor Shen
Nung ca. 2500 BC (Zhu 1998), the Code of Hammurabi from 1800 BC (Sigerist
et  al. 1955), the Egyptian Ebers Papyrus dated 1500  B.C. (Ebers 1889), and the
Sanskrit text Charaka Samhita on Ayurveda medicine dated 700 B.C. (Van Loon
2003). In occident, Hippocrates (460–377 BC) recorded and categorized by their
physiological action over 200 medicinal plants in the Book of [Medical] Aphorisms
(Castiglioni 1985), Dioscorides wrote the De Materia Medica (77 AD) with over
900 drugs mostly of plant origin (Riddle 1985), and Galen in the Treatise of Simple
Medicines emphasized on the properties of plants, animals, and minerals as simple
medicines (130–200 AD) (Petit 2017; Malapani 2016). In Egypt, the Maimonides
Medical Writings (1100  AD) compiled medical theory and herbal information,
especially in the 7th volume Treatise on Poisons and Their Antidotes (Rosner 2002).
In America, the Aztec and Maya Indian cultures of Mexico and Central America
also used natural substances as medicines. As is referred by Evans, postclassic
Mesoamericans developed a large and effective pharmacopoeia, formulae for medi-
cines concocted from animals, minerals, and especially plants (Evans 2004). The
ancient Aztec healers exploited at least 132 medicinal herbs to treat ailments rang-
ing from pimples and nosebleeds to gout and epilepsy and respiratory and gastroin-
testinal infections (Berdan 2005). After the arrival of Columbus, the Spanish Crown
dominated and exploited the medicinal plants market from the Americas up to the
eighteenth century (Miller 2017). Nowadays, each country has a particular approach
to medicinal plants; some countries have specific regulations and national pharma-
copeia that included plants or their derivatives, while others do not have regulations
about the use of traditional medicines of plant origin.
The World Health Organization (WHO) confirmed that traditional medicine
(TM) is still used by a high proportion of the world’s population. It has been esti-
mated that more than 80% of the people in developing countries depend on herbal
medicine for basic healthcare needs. In Western countries, where chemistry is the
backbone of the pharmaceutical industry, about 25% of the bioactive molecules are
of plant origin. The WHO has released the WHO guidelines on good agricultural
and collection practices (GACP) for medicinal plants as a means to recommend
practices to ensure good quality, safe herbal medicines, and ecologically sound cul-
tivation practices for future generations (WHO 2003).

1.4  Native Medicinal Flora from Argentina

Traditional medicine is a set of empirical practices from a group, which are trans-
mitted from generation to generation with the intention of solving health problems
from a specific population. The recognized native medicinal flora of Argentina
1.5 Regulatory Framework on Medicinal Plants in Argentina 11

comprises around 1529 taxa grouped into 175 families and 688 genera of vascular
plants (pteridophyte, gymnosperm, and angiosperm). Barboza et  al. (2009) have
made an exhaustive analysis of medicinal plants in Argentina. They have found that
the aerial parts (leaf, stem bark, etc.) and leaves are the most widely parts used in
medicinal preparations. Stems, underground organs, and flowers are used in a lesser
extent. Also, they found a high amount of references without specifications on the
parts used in folk medicine. More information related to medicinal Dicotyledoneae
than about Monocotyledoneae and Pteridophyta (58.92%) were found, being
Asteraceae, Fabaceae, and Solanaceae the three best representative families, fol-
lowed by Verbenaceae, Lamiaceae, Myrtaceae, and Apiaceae. From the available
data, the anti-inflammatory, antioxidant, and antimicrobial activities were the most
frequently cited; however, around 58% of the medicinal plants were not yet studied
on their pharmacological properties. Finally, they found that most of the available
information is referred to species from the provinces of SAL, JUJ, TUC, COS, MIS,
ERI, and CHA. Less cited are species from COR, BAI, and CAT. SDE and CHA are
the provinces with more medicinal plants and TDF the one with the lesser (Barboza
et al. 2009). In the following chapters, some of those species having pharmacologi-
cal studies that support their popular use will be described.

1.5  R
 egulatory Framework on Medicinal Plants
in Argentina

Since 1999 Argentina has a legislation that normalizes and regulates the use of
medicinal plants. The National Administration of Drug, Food and Medical
Technology (ANMAT) is the body that aims to strengthen the regulatory, inspec-
tion, and surveillance capacities about health products, including those referring
to medicinal plants and medicines of plant origin, at national and provincial levels.
The law N° 16,463, Resolution 1817/2013, from the Ministry of Health, approved
a glossary of terms related to phytomedicines, established the regime for register-
ing herbal medicines, defined a simplified regime for registering traditional herbal
medicines, and established the requirements that these medicines must fulfill to be
registered, their methods of elaboration, and the specifications for labeling and
advertising. The Argentine National Pharmacopoeia (FNA) is a national law that
contains the information about plants and their products that are authorized as
medicinal in the country. The last printed versions are the FNA VI edition (FNA
VI) and the last revision, Volume I of the FNA VII. Volumes II, III, and IV of FNA
VII were approved in November 2013 and published in the Official Gazette. The
FNA VIII edition has been sent to the National Congress for being sanctioned.
However, the FNA VI edition is still consulted. The FNA VII established that a
Vegetable Drug is a plant or their parts, entire, ground or powdered (flowers,
fruits, seeds, tubers, bark, etc.) fresh or dried, as well as juices, gums, latex, essen-
tial or fixed oils and others similar components, which are used pure or mixed in
the elaboration of medicines (Argentina National Pharmacopeia 2013).
12 1  Introduction: Native Plants of Argentina – A General Overview

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definición y sus principales áreas protegidas. In: El deterioro del suelo y del ambiente en la
Argentina, vol 1. Editorial Dunken, Argentina, pp 79–101
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pfds/Farmacopea_Argentina_2013_Ed.7.pdf
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Berdan FF (2005) The Aztecs of Central Mexico: an Imperial Society. Thomson Wadsworth,
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Burkart R, Bárbaro NO, Ro S, Gómez DA (1999) Eco-regiones de la Argentina. Administración
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Cabrera AL (1953) Esquema Fitogeográfico de la República Argentina. Rev Mus La Plata (N.S.)
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Cabrera AL (1958) Fitogeografía. In: de Aparicio F, Difrieri HA (eds) La Argentina: Suma de
Geografía, vol 3, pp 101–207
Cabrera AL (1971) Fitogeografía de la República Argentina. Boletín de la Sociedad Argentina de
Botánica 14:1–42
Cabrera AL (1976) Regiones fitogeográficas argentinas. In: Kugler WF (ed) Enciclopedia
Argentina de Agricultura y Jardinería. 2nd edn. Acme, Buenos Aires, Argentina. 2(1):1–85
Cabrera AL, Willink A (1980) Biogeografía de América Latina. 2nd edition Serie de Biología.
Secretaría General de la Organización de los Estados Americanos. EEUU, Washington, D.C.,
p 120
Castellanos A, Pérez Moreau RA (1941) Contribución a la Bibliografía Botánica Argentina.
Lilloa 7
Castellanos A, Pérez Moreau RA (1945) Los tipos de vegetación de la República Argentina.
Monogr Inst Estud Geogr Univ Nac Tucumán 4:1–154
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Cingolani AM, Bran D, López C, Ayesa J (2000) Comunidades vegetales y ambiente en el ecotono
boreal entre los distritos patagónicos Central y Occidental (Río Negro, Argentina). Ecol Austral
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Diminich MC, Franco RD, Fernández EM, Zorzi F (2018) Capítulo 5 Ecosistemas. In: Biología 3.
Flujo de la información genética, los procesos evolutivos, de los individuos a los ecositemas,
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Aires, pp 94–109
Ebers G (1889) The Papyrus Ebers. A translation. https://archive.org/details/
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Evans ST (2004) Ancient Mexico and Central America. Thames and Hudson, New York
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Hauman L (1920) Ganadería y Geobotánica en la Argentina. Rev Centr Estud Agron y Vet Bs As
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Belgique 64:20–79
Holmberg E L (1898) La Flora de la República Argentina, en Segundo Censo de la República
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la Patagonia extra andina. Ecol Austral 8:123–141
Lorentz PG (1876) Die Vegetationsverhältnisse der Argentinischen Republik. In: Richard Napp:
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Malapani A (2016) Galen on pharmacology: his scientifi c way of thinking and contribution to the
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témicos argentinos (Jorge Morello, Silvia D. Matteucci, Andrea F. Rodríguez, Mariana Silva).
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ficas, ecorregiones o paisajes? Fronteras 14:1–16
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Academic Publishers, Amsterdam
Chapter 2
Ethnobotany in Argentina

2.1  Introduction

Ethnobotany is the scientific approach to the relationships between people and

plants into the frame of their sociocultural context. In this chapter a review of some
aspects from ethnobotany in Argentina from the first references made by the
members of the Society of Jesus (Compañía de Jesús) up to this day is introduced.
Most of the studies already made were focused from a regional or ethnic point of
view, although some combine both aspects.

2.2  Ethnobotany

In ancient times, naturalists, travelers, and missionaries collected information

about habits and practices of societies in distant and new discovered communities
or tribes. That information was often preserved in monasteries, abbeys or, in the
New World, in catholic reductions called missions. The written documentations on
the therapeutic uses of plants in the ancient Mediterranean world currently known
are the Hippocratic Collection, Historia Plantarum by Theophrastus, and De
Materia Medica by Dioscorides. The last one, often dated to ca. 70 AD, has been
the basis of most of the literature on materia medica in the Mediterranean and
Western tradition up to the surge of modern pharmacy. The study of useful plants
is probably one of the oldest sciences in the world. Moreover, people that gathered
the information about plant properties were always respected in their communities
for having a knowledge that was hidden to the majority. Even before the writing
was developed, the man left records of the many uses, including medicinal and
religious, of plants. Traditionally, ethnobotanists were looking into documents and
archeological sites for connections between ancient civilizations and settlements
with plants. Those data revealed the migration habits, commercial routes,

© Springer Nature Switzerland AG 2019 15

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_2
16 2  Ethnobotany in Argentina

agricultural practices, and the origin and dispersion of those plant species.
Similarly, ethnobotanists study the relationship between people and plant species
of today compiling the knowledge safeguarded by indigenous and rural people
(Álvarez 2014). Nonetheless, ethnobotany is not only restricted to the interactions
between traditional cultures and plants. In a more broad and modern approach,
ethnobotany comprises the study of both traditional and nontraditional populations
and their connections with their botanical environment. Then, ethnobotany is not
confined to the use of plants by ancient communities but also to the relationship
that modern societies have with them. Recently, some authors are referring to the
ecological dimension of ethnobotany as is the case of the relation with plants
among immigrants (Kujawska and Pieroni 2015; Scarpa et al. 2016), in urban envi-
ronments (Hurrell 2014), gardens (Kimber 2004; Calvet-Mir et al. 2014), and in
fairs and markets (Ladio et al. 2013; Albuquerque et al. 2017). Some new questions
are: How does the botanical background of a group change with migration or the
arrival of immigrants? When the communities age and young people move, are the
knowledge moving with them or it remains nested in the old community? How do
traditional communities manage horticultural and agricultural techniques without
harming the ecosystem? How was the environment of the communities modified
by human interaction along time? What represent plants to people? (Korstanje
2016; Albuquerque et al. 2017). Regularly, in traditional communities plants are
more than food and health; they are also part of their religion, magic, and rites. In
brief, they are part of their culture and everyday life. However, it is usual to find
reports describing the medical use of plants avoiding the underlying significance of
that use. In that case, the current consensus is to call that type of studies as medical
ethnobotany (Pake 1987). Consequently, with a broad-minded analysis, ethnobot-
any focus is turning to the study of the dynamic interrelationship between human
societies and plant communities within particular sociocultural and environmental
contexts (Rodríguez et al. 2006; Bonomo et al. 2011).
From the scientific point of view, ethnobotanical research evolved from invento-
ries of useful plants to detailed quantitative studies supported by solid methods and
analysis. Those methods should aim at collecting high-quality, representative data
that have statistical strength. First of all, ethnobotany should accurately identify
plant species. In doing that, the researcher will allow for the recovery of all the
information linked to that species over the years. Not only the scientific name of the
species must be properly recorded, but also herbarium specimens must be collected
and appropriately deposited. Misidentifications will produce strong inaccuracies on
current and future research (Blanché et al. 1996; Arenas and Kamienkowski 2014).
Second, information must be analyzed; cultural domain analysis, ethnographic
methods, participatory mapping and ground-truthing, plot, transects and tree trials,
etc. are approaches that can be used to manage and analyze the collected informa-
tion (Caruso et al. 2015).
The great importance of traditional knowledge is recognized in international ini-
tiatives as the Ecosystem Assessment (Reid et  al. 2006), the Economics of
Ecosystems and Biodiversity (Brondizio et  al. 2010), or the Intergovernmental
Platform Scientific and Technological Advice on Biodiversity and Ecosystem
Services (IPBES) (de Santanaya 2014). Certainly, its inclusion in the United Nations
2.3 Ethnobotany in Argentina 17

Convention on Biological Diversity (1992) was a landmark in its recognition and

appreciation. Therefore, it has been recommended to all states to deepen scientific
research, especially with regard to biological inventories and methods of extraction,
identification, and standardization of substances from plants (OMS 1993). On the
other hand, the ethical dimension of research must be considered; the Code of Ethics
from the International Society of Ethnobiology proposes that “all ethnobiological
research should be collaborative; it establishes a community-centered approach,
and calls for all research to strengthen community goals” (http://ethnobiology.net/
code-of-ethics/).

2.3  Ethnobotany in Argentina

The oldest reference about medicinal plants from Latin America was a Royal decree
dated on the year 1570 signed by King Felipe II from Spain. There, he established
that “... all herbs, trees, plants, or seeds of any medicinal value that can be found in
those places will be sent to this Kingdom.” Hence, the first collection on medicinal
plants from the New World was published in 1597 under the title Introduction and
memories of the description of the Indians that His Majesty ordered to do for the
good government and ennoblement of them (Trillo et al. 2011).
At least empirically, ethnobotany in Argentina dates from the seventeenth cen-
tury, when the Jesuits settled the reductions or missions at the northeast of the coun-
try in the Mesopotamia and also in the neighboring regions from Brazil, Bolivia,
and Paraguay. Besides their religious, social, and economical purposes, the Jesuits
also gathered information about native traditions, language, and customs which also
included the characteristics of the native plants and their medicinal uses. However,
there is scarce documentation about their findings. Some of them are the publica-
tions of three missionaries from the Society of Jesus, Pedro Lozano, Florian Paucke,
and Pedro Montenegro. Lozano was designated historian from the Society (histo-
riograpus provinciae) from 1730; hence, as an endorsed historian, he offered an
official chronicle following the constraints of the order. He wrote the Descripción
Chorográfica del Gran Chaco Gualamba and other works of historical value, La
historia de la conquista, La historia de la Compañía, and La historia de las revolu-
ciones del Paraguay. The book Descripción Chorográfica del Gran Chaco
Gualamba was published in 1733  in Córdoba, Spain, and reedited in 1941  in
Tucumán, Argentina (Lozano 1941). It was accompanied by a map of Gran Chaco
by father Antonio Machoni that could also be found in the reedition. Florián Paucke
was a missionary in the mocovíes reductions of San Francisco Javier and of San
Pedro, from 1749 up to the eviction of the Society in 1767. His work Hacia allá y
para acá: Una estada entre los indios Mocobíes 1749–1767 was written between
1778 and 1779, but first published in 1944. His style differs from Lozano in the
close approach to the indigenous and locations he made; consequently, his work is
considered by some authors comparable to that of an ethnographer. The final
­version of his work, which included watercolor illustrations, was written from exile
with the subtitle There we went pleasant and happy, here we returned bitter and
18 2  Ethnobotany in Argentina

saddened (Paucke s/f 2010; Rosso and Cargnel 2012; Rosso and Medrano 2013).
Brother Pedro de Montenegro, a physician that entered the Society and had estab-
lished in the Paraguay province, wrote the Materia Médica Misionera (1710). It was
described by Lozano as “an incredible surgeon (chirurgus), nurse (infirmarius), and
herbalist.” His Materia Medica contains a rich description of native plant species,
their therapeutic properties, and mode of administration (Montenegro 1945).
Materia Médica Misionera is a codex of 458 illustrated pages with 136 figures,
drawn by pen and ink. On the cover there is a picture of Our Lady of Sorrows,
patron saint of the sick, and at the bottom the date “Year of 1710.” The first three
sections refer to the botanical nomenclature, to the properties of the plants, the time
of collect, and how to preserve them, their healing virtues, and how to take advan-
tage of them. The fourth part is a medical study of diseases that are cured by herbs,
roots, and barks (Perkins de Piacentino 2007). Hence, the Plantae Diaphoricae
Florae Argentinae (Hieronymus 1882) is considered as the first compilation of
medicinal, poisonous, or useful plants that grew in Argentina (native or exotic). A
remarkable characteristic of that book is that the author only included species whose
scientific names were known in those days, producing a valuable scientific enu-
meration of species, their characteristics, and uses. Other scientists also made a
great contribution to plant knowledge related to the primitive inhabitants of the
country. Aimé Bonpland, a French explorer, physician, and botanist, traveled with
Alexander von Humboldt to Spain, Venezuela, Colombia, Ecuador, Cuba, México,
and the United States (1799–1804). They produced an herbarium with approxi-
mately 6000 samples, 4500 of them new (described in Nova genera et species plan-
tarum, from Humboldt, Bonpland, and Kunth) that are deposited in Paris. Later on,
he traveled to South America, arriving to Argentina in 1817. His archive, with more
than 2050 documents (scientific, epistolary, historical) is in the Museo de
Farmacobotánica, Juan A. Domínguez, in the Faculty of Pharmacy and Biochemistry,
University of Buenos Aires. Among his manuscripts are the Botanical Journals
where he described 2884 plants that he collected in the Río de la Plata region.
Juan A. Domínguez was a pharmacist committed to the study of phytochemistry
and botany. He was the author of Contribuciones a la Materia Médica Argentina
(1928) and a series of publications on medicinal plants, particularly those used by
indigenous from Argentina. His herbarium is part of the Museo of Farmacobotánica
Juan A. Domínguez collection (Amorín 1990; Giberti 2008; Krapovickas 2008).
Raúl N. Martínez Crovetto was the first Argentinean researcher that introduced
the expression ethnobotany in his articles. His work was focused on both indige-
nous and rural people from different locations in the country, mainly from the
northeast region. His approach was qualitative, meticulously descriptive and
interpretive (Pirondo and Keller 2012). Among its prolific series of publications
are the Estudios etnobotánicos I-V, the last one published post-mortem by his col-
laborators in Bonplandia, the journal he founded. The library of the Instituto de
Botánica del Nordeste preserves numerous documents belonging to him (Pirondo
and Keller 2012).
Domingo Parodi, Eduardo Matoso, Nicolás Rojas Acosta, Lorenzo R. Parodi,
and Arturo Ragonese were also remarkable contributors to the ethnobotanical stud-
ies in Argentina.
2.3 Ethnobotany in Argentina 19

Currently, there are profuse investigation covering aspects such as the relation-
ship between people and plants in connection with their ethnical origin, history, and
cultural landscape (Arenas 2009; Arteaga 2010; Capparelli et al. 2011; Rosso 2013;
Kujawska 2016; Scarpa and Rosso 2018). I will make here a brief review of some
of the available literature in order to bring a general glance about the work already
made. For more information there is a great amount of material that could be found
in the articles cited in this chapter.
I consider the work of Barboza et al. (2009) of a paramount relevance, since they
have performed an exhaustive and detailed review of native medicinal plants
including bibliometric, phytochemical, and etnopharmacological screenings. In that
publication are described around 1520 taxa. Among the species listed, only about
half of them have chemical and pharmacological studies carried out. Consequently,
there is still a lot of work to be done in order to substantiate the accuracy of their
traditional use. From the article, it is clear that the uppermost diversity of plants
with medicinal properties is located at the Northwest, Mesopotamia, and Gran
Chaco regions, being SAL and JUJ the provinces with the highest number of
medicinal taxa. In consequence, reports are more numerous from the central and
northern part of the country than from the south.
There are records compiling plant catalogues, references about plant handling,
methods of collection, uses, part used, preparations, route of administration,
conservation, and the social-cultural context of the consumers.

2.3.1  Ethnobotany in Indigenous and criollos Communities

There are a large number of publications specifically related to the ethnobotany of

indigenous communities (Arenas 1995). I will refer here only to some of them; a list
of ethnobotanical research related to communities from Argentina (indigenous and
criollos) is displayed in Table 2.1. In 1967 Martínez Crovetto gathered information
about the ethnic group mocovi, or moqoit, from the southwest of the province of
CHA. Those records were included in several articles describing the medicinal plants
of mocovis, their scientific name, specific use, part of the plant used, elaboration, and
administration (Martínez Crovetto 2014; Scarpa and Rosso 2014). A comparison
was later made between the contemporary alimentary ethnobotany and the one of
mocovis during the eighteenth and twentieth centuries (Rosso and Scarpa 2017)
founding the highest similarities with the information recorded in the twentieth cen-
tury. Bromelia hieronymi, Aechmea distichantha, Araujia odorata, and Sarcomphalus
mistol were the more consumed species, preferably as raw fruit. In those communi-
ties, the edible use of the fresh fruits of Scutia buxifolia, the flowers of Glandularia
peruviana, and the use as salt of cfr. Sporobolus spartinus ashes were there for the
first time recorded. In the case of chorote, hunter-gatherer and fishermen who live in
the Northwest of Argentina, a description was made about the edible wild plants they
collected as well as a detailed description of the used parts, their elaboration, mode
of conservation, and storage. Some of the species were also consumed by other
groups of the same linguistic family (Mataco-Macá), and only a few ones were
20 2  Ethnobotany in Argentina

Table 2.1  Ethnobotanical studies related to ethnic groups from Argentina (indigenous and
criollos)
Communities Research subject References
Aónikenk, Selk’nam, Ethnobotany mushroom Díaz (2010)
Kawésqar, Yagan, and
Haush
Chorote Alimentary ethnobotany Arenas and Scarpa
(2007), Scarpa and
Pacor (2017)
Medical ethnobotany Scarpa (2009)
Ethnobotany and religion Scarpa and Pacor
(2015)
Criollos Ethnobotany Scarpa et al. (2016)
Medical ethnobotany Scarpa and Rosso
(2019)
Huarpes Medical ethnobotany Montani (2013)
Mapuches Alimentary and medical ethnobotany Molares and Ladio
(2015)
Mataco-ihoko’tax and Alimentary ethnobotany Arenas (2000)
toba-pilagá
Mbya Medical ethnobotany Remorini and Sy
(2002)
Mocovi Ethnobotany Martínez Crovetto
(2014), Scarpa and
Rosso (2014)
Alimentary ethnobotany Rosso and Scarpa
(2017)
Historical ethnobotany Rosso (2013)
Pilagá Medical ethnobotany Filipov (1994)
Quilmes Medical ethnobotany Ceballos and Perea
(2014)
Ranqueles Ethnobotany Steibel (1997)
Selknam Archeobotany Caruso et al. (2008)
Toba-pilagá Ethnophytogeography Scarpa and Arenas
(2004)
Toba Sorcery and funeral practices Arenas (2013)
Wichi Ethnobotany, phytonomy of herbs and Suárez (2011), Suárez
climbing plants of the semiarid Chaco, Salta (2014)
Province

exclusive to them (Arenas and Scarpa 2007). As the knowledge about the chorote
ethnobotany is scarce, that work is a contribution of great relevance; however, it was
only referred to chorotes living in Argentina, not in Paraguay. Scarpa and Pacor
(2015) have made an analysis about the religious symbolism of Anisocapparis spe-
ciosa (Capparaceae) (axis of the world and inexhaustible regeneration) for chorotes
and its medicinal use. It was interesting to found that chorotes and criollos from the
same region shared approximately half of their ethnomedicine. Maybe the
2.3 Ethnobotany in Argentina 21

incorporation by the chorotes of some species used by criollos is a result of the

restrictiveness of their ethnomedicine (Scarpa 2009). On the other hand, it was
reported a cultural change in chorote society as a consequence of the growing stigma
and prejudices involved in the collection of edible plants (Scarpa and Pacor 2017).
As for the wichis from semiarid CHA, the phytonymy of herbs and climbing
plants was made (Suárez 2011). The vernacular name of 90 species was recorded
along with their translation to Spanish and also a semantic and linguistic analysis in
order to establish nomenclature patterns.
A portrayal of the Mapuche cosmovision, the participation of shamans and other
mediators, and the description of specific healing rites are portrayed in the article of
Albornoz et al. (2004). A record of their medicinal species, with their scientific and
vernacular name and their uses, was included. A similar analysis was made for the
mbya-guaraní communities from Misiones (Remorini and Sy 2002).
An inventory of the species of ethnobotanical interest for the Aónikenk, Selk’nam,
Kawésqar, Yagan, and Haush communities from West TDF-Patagonia with their
uses and medicinal properties was made based on bibliographic information (Díaz
2010). A number of 68 taxons were registered, mainly used as food, medicine (anal-
gesic, digestive, disinfectant, diuretic, etc.), and to a lesser extent, as fuel or as part of
rituals. The different ways of plant use among the Selknam from Tierra del Fuego
were discovered from carbonized and non-carbonized wood remains recovered at
Ewan site. Woody species in that hunter-gatherer group were particularly used for
construction and as fuel. Seeds of Empetrum rubrum were found in one of the arche-
ological sites (Caruso et al. 2008). A record of the species of ethnomedical interest
was also made for the pilagá from Central CHA (Filipov 1994), ranqueles from LPA
(Steibel 1997), and quilmes from TUC (Ceballos and Perea 2014).
Also, a historical analysis from non-medicinal plant species used by criollo
groups was made from the information presented by the Argentinian Government at
the Universal Exhibition of Paris of 1889. The author identified the most employed
plant families of that time and their uses (Scarpa 2017). The more reported uses
were, from most cited to less cited, as dyes, utensils, furniture, food, transport,
tanning, home building, other constructions, and fuel.
The medical ethnobotany from Criollos from the Humid Chaco region was ana-
lyzed considering the data from the 1921 National Folklore Survey. The remedies
for the digestive system, respiratory system, and vulneraries were the most repre-
sented (Scarpa and Rosso 2019).

2.3.2  Ethnobotany from Different Regions from Argentina

The ethnobotany from different regions from Argentina was also profusely analyzed
(Table 2.2). There are numerous catalogues of medicinal species from the province
of BAI with its corresponding taxonomic classification, botanical characteristics,
uses, and medicinal preparations (D’Alfonso et  al. 2011; Hurrell et  al. 2015).
Besides, agricultural practices in the conservation and selection of vegetables
22 2  Ethnobotany in Argentina

Table 2.2  Ethnobotanical research in different regions of Argentina

Region Research subject References
Buenos Aires
Azul Medicinal ethnobotany Vercelli et al. (2013)
Tandilia system Botanical knowledge D’Alfonso et al. (2011), Hilgert et al.
(2010)
Metropolitan area Urban ethnobotany Hurrell et al. (2015), Hurrell et al.
(2013), Puentes and Hurrel (2015)
Buenos Aires-La Plata Urban ethnobotany Hurrell et al. (2016), Puentes and Hurrel
conurbation (2015)
La Plata Ethnobotany Doumecq and Arenas (2018), Puentes
and Hurrel (2015)
Isla Santiago Medical ethnobotany Hernández et al. (2013)
Parque Costero del Alimentary and medical Echenique et al. (2018)
Sur ethnobotany
Laguna de Mar Medical ethnobotany Arias Toledo and Trillo (2014)
Chiquita
Hurlingham Urban ethnobotany, urban Bach et al. (2014)
phytotherapy
Paraná Delta Archeobotany Bonomo et al. (2011)
Misiones Medical ethnobotany Stampella et al. (2018), Zamudio et al.
(2010), Kujawska and Pieroni (2015),
Kujawska (2016)
Ethnopharmacology Amat and Vajía (1991)
Formosa Historical ethnobotany Anconatani and Scarpa (2015), Scarpa
and Anconatani (2017)
Chaco Medical ethnobotany Arenas (1997), Scarpa (2004), Marinoff
et al. (2006)
Corrientes Urban ethnobotany Pirondo et al. (2011)
La Pampa Alimentary ethnobotany Muiño (2012)
from peasants
Sierras Pampeanas Archeobotany Salazar (2015)
Medical ethnobotany from Novoa et al. (1998)
monocotyledonous
Patagonia Urban ethnobotany Ladio et al. (2013), Molares and Rovero
(2016)
Yungas Alimentary and medical Hilgert (1999), Hilgert and Gil (2006)
ethnobotany
Salta Medical ethnobotany Martínez et al. (2004), Hilgert (2001)
Upper Bermejo Basin Plants involved in the Hilgert (2000)
manufacture of yista
North West Region Archeobotany research Oliszewski (2005)
(NOA) (0–600 a.C.)
Jujuy Medical ethnobotany Acosta et al. (2015), Romeo (2015)
Urban ethnobotany Acosta et al. (2017)
Tilcara Ethnobotany Barbarich and Suárez (2018)
(continued)
2.3 Ethnobotany in Argentina 23

Table 2.2 (continued)
Region Research subject References
Los Juríes, Santiago Medical ethnobotany Riat and Pochettino (2015)
del Estero
Puna Archeobotany Rodríguez et al. (2006)
Córdoba Medical ethnobotany Martínez (2005), Arias Toledo et al.
(2007), Martínez (2008a, 2008b), Arias
Toledo (2009)
Veterinary ethnobotany Martínez and Luján (2011)
Cultural landscape Trillo (2016)
Diachronic ethnobotany Fernández and Martínez (2019)
Ethnobotany Arias Toledo et al. (2009)
Sierra de Medical ethnobotany Goleniowski et al. (2006)
Comechingones
Río Cuarto Medical and urban Madaleno and Montero (2012)
ethnobotany
Chaco forest Alimentary and Arias Toledo et al. (2007)
ethnomedicine
Pampa-Cuyo Medical ethnobotany Muiño (2011)
Andean Argentina Archeobotany and Caparelli et al. (2005)
ethnohistory
Rio Negro, Patagonia Medical ethnobotany Ochoa et al. (2010)
Ethnobotany Lozada et al. (2006)
Tierra del Fuego Archeobotany Caruso et al. (2008)

(Ahumada et al. 2010), traditional botanical knowledge in urban zones (Pochettino

et al. 2008), marketing of medicinal plants (Arenas et al. 2011; Hurrell et al. 2013),
and the use of forest species as food and medicine, (Sharry et al. 2011) among other
subjects. Were also studied. The current sociocultural changes in the metropolis of
Buenos Aires and the province are reflected in research related to more recent
immigration. Such is the case of two articles on urban ethnobotany. One of them
revealed an increase in  local biocultural diversity by analyzing the presence of
medicinal plant products (e.g., Stevia rebaudiana, Achyrocline alata, Cuphea
lysimachioides, Euphorbia serpens, etc.) brought by immigrants from Paraguay
(which represents 30.5% of the immigrants in Argentina) to the metropolitan area of
BAI (Hurrell et al. 2016). The others refer to the traditional Chinese phytotherapy
insertion in ciudad de Buenos Aires. For each species their scientific and vernacular
names in Spanish, Chinese, and Latin are included along with their distribution, use
linked with tradition, biological activity, and effects (Hurrel and Puentes 2017).
The province of COR is proliferous in investigations related to its flora. The
changes in the knowledge and management of medicinal plants were evidenced by
various articles. A study made in a rural community revealed that people used and
cared more medicinal plants than wild edible plants, with a marked influence by the
sociocultural context (Arias Toledo et al. 2007). From the individuals studied, those
24 2  Ethnobotany in Argentina

with better awareness were people that had acquired that knowledge through formal
education and people, old or native residents, that had an empirical knowledge
about nature. A third and extensive group was composed by people without any
familiarity with medicinal plants. The same indifference was found related to the
use of native plants as food, which was attributed to the prevalence of meat as the
main ingredient of the diet for Argentinean people, neglecting the ingestion of veg-
etables, and to the extended association of the use of wild plants as food with pov-
erty (Pelto et al. 1989; Aguirre 2005). The knowledge and use of medical plants by
residents living near natural protected areas in Sierras de COR, which has a higher
variety of species, did not show differences with people living in remote places.
Both groups used the same species in cases of digestive, respiratory, or circulatory
problems (Furlan et  al. 2011). In the chaqueña phytogeographical region from
COR, the familiarity about medicinal plants was higher in older residents than in
young people, which is probably related to the modifications in lifestyle, the access
to medical assistance, and a growing detaching of rural practices. On the contrary,
no differences were found related to gender (Arias Toledo et al. 2009).
In MIS, which also has a great amount of literature referred to its medicinal
plants, studies related to the behavior of natives, criollos, and descendant of
immigrants coming in the first moiety of the twentieth century were made. There
were found patterns of medicinal pluralism among peasant descendants from Polish
immigrants in MIS (in the settlements Wanda and Gobernador Lanusse). Their
determination to retain their traditional healing practices was mixed with the
apprehension of knowledge from natives and criollos through social cohabitation
and mixed marriages. The study revealed that they used medicinal plants (22 species
known from the home country and other native species), restored their health at
home, and required the assistance of healers as curanderos, hueseros, and naturis-
tas. Medicalization was restrained to childbirth and fractures or when life was
threatened (Kujawska 2016). Polish and criollo community from MIS diverged on
the use of wild honeys produced by insects of the order Hymenoptera (bees, wasps,
and bumblebees) as food and medicine. All the investigated honeys were used by
both communities as food, but only a few of them are considered also as a medicine
(honey from bee, yateí, mirí, carabozá). The ethnospecies borá, mandasaia, and
iratin were exclusively mentioned as medicinal by the criollo population. Home
remedies based on honey are most frequently used to treat illnesses related to the
respiratory system in both populations. Polish and criollos prepared home remedies
in a similar way but showed differences in the frequency and mode of application
(Zamudio et al. 2010). As for the relevance regarding the cultural context, it was
seen that the main Citrus spp. from the region varies according to the cultural
context; for criollos the more relevant species are limón mandarina and naranja
dulce, whereas for residents of the Yabotí biosphere is lima ácida (Stampella et al.
2018). The medicinal use of Citrus among the criollos revealed that the juice, tea, or
decoctions made with non-processed endocarp and leaves (7 ethnospecies, 5 taxa)
alone or combined with other plants were used to treat flu, fever, hypertension,
cough, and digestive or nerve disorders.
2.3 Ethnobotany in Argentina 25

An alimentary ethnobotanical analysis from peasant descendants from immi-

grant settlers in LPA registered 62 taxons (4.7% of the wild species, 31% native
taxons) that were used to prepare meals, sweets, and beverages. The knowledge
about those plants was higher in older people (Arias Toledo et  al. 2009; Muiño
2012).
In the Yungas the edible plants from the Quebrada de Humahuaca were recorded
including their scientific and vernacular name, parts of the plants employed, methods
of collection and application, cooking, commercial relationships with neighboring
settlements, and their geographical localization (Hilgert 1999). In the Northwest
(Korstanje 2016), in peri-urban neighborhoods from the city of SAL, the habit of
chewing leaves of Erythroxylon coca (coqueo) was extended to a 54% of the
population which is related to ancestral cultural customs not only restricted to the
descendants of the original natives. Other medicinal plants that were reported to be
commonly used were quimpe (Lepidium didymum), tusca (Acacia aroma), paico
(Chenopodium ambrosioides), molle (Schinus areira), ruda (Ruta chalepensis), and
hediondilla (Cestrum parqui), being taken in cases of gastrointestinal, respiratory,
and kidney alterations in different preparations. In opposition with the finding of
Arias Toledo et al. (2009), no differences in knowledge about medicinal plants were
found related to age (Oliva 2016). In the Calchaquí Valley (Salta), traditional healers
managed a phytotherapy of 42 species specific for regional diseases related to the
environment and the rural activities of the population, such as osteoarticular and
cardiocirculatory problems. The article of Martínez et  al. (2004) brings a deep
description of species, parts of the plants used, method of preparation, medical uses,
and route of administration. In Patagonia, the commercialization of weeds as food
and medicinal plants in urban fairs from Bariloche was analyzed, paying particular
attention to the cosmovision, attitudes, and behavior of the horticulturalists. There,
as was referred for the province of COR (Ladio et al. 2013), the use of edible wild
plants was initially associated to poverty, a perception that slightly changed after an
instructive workshop. Another study made in CHU, in a region inhabited
predominantly by descendants of the Mapuche-Tehuelche indigenous people,
revealed that the most versatile species (Acantholippia seriphioides, Valeriana spp.,
Mentha, and Artemisia absinthium) were the most relevant in folk medicine since
they can treat a variety of medical conditions (Richeri et al. 2013).
There are also ethnomedical studies related to veterinary medicinal plants. In
CAT, it was found 43 species from 30 botanical families with 62 medicinal uses
such as healing, digestive, antiparasitic and oxytocic. Also, plants were involved in
cultural practices of religious-ritualistic nature to heal animals (Martínez and
Jiménez-Escobar 2017). Similar studies were made in Sierras de COR with the
additional comparative analysis with the use of those plants in human medicine. A
total of 127 medicinal uses were registered, corresponding to 70 species of plants
belonging to 39 botanic families (Martínez and Luján 2011).
Ethnobotanical studies focused on a particular species or biological activity were
also profused. Some examples are the ethnobotanical studies about cardón
(Trichocereus atacamensis) from Tilcara in JUJ (Barbarich and Suárez 2018),
26 2  Ethnobotany in Argentina

Hypericum robsonii spec. nova sect. Trigynobrathys (Hypericaceae) from MIS

(Keller and Crockett 2015), ligas (Loranthaceae), and Synandrospadix vermitoxicus
(Scarpa and Montani 2011; Arenas 2016). As for the ethnobotanical studies focused
on the biological activities, there are reports about antimicrobial and antifungal
activities, among others (Demo et al. 2008; Rondina et al. 2010).

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Chapter 3
Cuyo

3.1  Introduction

3.1.1  The Cuyo Region

Cuyo is a geographical region located in the western center of Argentina; it covers

the provinces of Mendoza (MEN) and San Juan (SJU) and the occidental side of
San Luis (SLU). It is flanked to the South by the Patagonia, to the East by the
Sierras Pampeanas and the Pampeanas Plain, and to the West by Chile. The geo-
graphical characteristics of the region are diverse as there are cordilleras (high
mountain ranges), mountains, and valleys with sparse vegetation and a desert cli-
mate. To the West of Cuyo is located the Andes Mountain range (Cordillera de los
Andes) including the Aconcagua, the highest mountain of America (6959 meters
high). Parallel to that are the Cordillera Frontal that has several independent cords
and the Andean foothills of MEN and SJU. By the East, there are vast plains, the
Travesías, which ended in the Pampa region. There are also several isolated moun-
tain ranges belonging to the Sierras Pampeanas; in the lower zones of the plains,
there are salt flats (e.g., Salinas del Diamante, a national protected area). In the
South of MEN are plateaus and volcanoes of the La Payunia Provincial Reserve. In
Cuyo prevails an arid climate, cold in winter and hot in summer; hence, it could
have both snowstorms and frost or heat waves according to the season. In the Andes
as in the desert territory, the weather is extremely hot during the day and very cold
at night. The Zonda (foehn wind) is a northerly dry, hot, and suffocating mountain
wind typical from this region, which appears usually between May and November
but especially in winter. Cuyo is an area of seismic risk, particularly in the provinces
of SJU and MEN. Also, there are chances of tornadoes, thunderstorms, landslide,
alluvium, and wildfires. The wine industry is the prevailing commercial activity,
being this region one of the main wine producers from South America. Dry weather
and cold winters favor the development of the vines; despite the heat during sum-
mer, the height where the crops are located favors the quality of the internationally

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34 3 Cuyo

valued wines. The most common varieties are Malbec, Bonarda, Cabernet
Sauvignon, Pedro Giménez, Syrah, Merlot, Tempranillo, and Chardonnay. The agri-
cultural production includes olives, plums, damasks, peach, nuts, corn, sorghum,
soy, oats, and horticultural and ornamental species. Also, there is production of
honey, mineral water, fruit juices, beer, cider, and by-products of the wine industry
(e.g., vinegar). Livestock production includes cattle, sheep, caprine, poultry, chin-
chilla, and ñandúes. As for mining, there is exploitation of salt, onyx, tungsten,
wolfram, uranium, gold, silver, dolomites, bentonites, quartz, marble, and feldspar,
among others (mainly in the provinces of SLU and SJU). Also, there is one of the
main oils and gas basins of the country (Luján de Cuyo, Tupungato, Rivadavia,
Malargüe, Maipú, and Junín departments).
The fauna is diverse; there are pumas, foxes, wild cats, guanacos (Lama guani-
coe), red lizards, ferrets, maras or Patagonia hare (Dolichotis patagonum), and
cuises (Southern mountain cavy). As for birds, there are ñandúes (Rhea), spot-
winged falconet (Spiziapteryx circumcincta), elegant-crested tinamou (Eudromia
elegans), and burrowing parrot (Cyanoliseus patagonus), among others. In some
lakes (e.g., Lancanelo lake), there are flamingos, plovers, ducks, rails (Rallidae
spp.), and swans.
The vegetation is characteristic of an arid region, with predominance of xerophi-
lous plants. Among the flora predominate those from the Monte phytogeographical
region with Prosopis alba and P. nigra (algarrobos); a variety of shrubs such as
Condalia microphylla (piquillín), Larrea nitida (jarilla), Opuntia quimilo (quimil),
Celtis tala (tala), and Geoffroea decorticans (chañar); as well as cactus (Fig. 3.1).
Some of the medicinal plant species reported in Cuyo are Acacia caven (Molina)
Molina (Fabaceae), Acantholippia seriphioides (A. Gray) Moldenke (Verbenaceae),
Achyrocline  satureioides (Lam.) D.C. (Asteraceae), Acmella decumbens (Sm.)
R.K. Jansen (Asteraceae), Acaena magellanica (Lam.) Vahl (Rosaceae), Adiantum
raddianum Presl. (Adiantaceae), Aloysia gratissima (Gillies at Hook, ex Hook)
Tronc. (Verbenaceae), Amaranthus muricatus (Moq.) Hieron. (Amaranthaceae),
Ambrosia tenuifolia Spreng. (Asteraceae), Anemia tomentosa (Savigny) Sw. var.
australis Mickel (Anemiaceae), Apium sellowianum Wolff (Apiaceae), Argemone
subfusiformis G.B.  Ownbey (Papaveraceae), Aristolochia argentina Griseb.
(Aristolochiaceae), Asclepias campestris Decne (Asclepiadaceae), Aspidosperma
quebracho-blanco Schldl. (Apocynaceae), Atriplex crenatifolia Chod. & Wilczk
(Chenopodiaceae), Bacharis articulata (Lam.) Pes. (Asteraceae), Bauhinia forfi-
cata Link (Fabaceae), Bidens pilosa L. (Asteraceae), Buddleja  araucana Phil.
(Buddlejaceae), Buddleja mendozensis Benth (Buddlejaceae), Caesalpinia gilliesii
(Wal. Ex Hook.) D. Dietr. (Fabaceae), Capparis atamisquea Kuntze (Capparaceae),
Capsicum chacoense A.T. Hunz. var. chacoense (Solanaceae), Cardionema ramo-
sissimun (Weinm.) Nelson & Macbr. (Caryophyllaceae), Celtis tala Planchon
(Ulmaceae), Cestrum parqui L’Her. (Solanaceae), Chaptalia nutans (L.) Pol.
(Asteraceae), Chenopodium ambrosioides L. (Chenopodiaceae), Colletia spinosis-
sima Gmelin (Rhamnaceae), Conyza bonariensis (L.) Cronquist. (Asteraceae),
Coronopus didymus (L.) Smith (Brassicaceae), Cortaderia  selloana (Schultes)
Asch. & Graeb. (Poaceae), Cucurbitella asperata (Gill.) Walpers (Cucurbitaceae),
Cuphea glutinosa Cham. & Schldl. (Lythraceae), Cuscuta indecora Choisy
3.1 Introduction 35

Fig. 3.1  The Cuyo region. Landscapes from Mendoza (a), San Juan (b), and San Luis (c)
provinces

(Cuscutaceae), Fabiana imbricata Ruiz et Pavon (Solanaceae), Gentianella acha-

lensis (Hieron. Ex Gilg.) Fabris (Gentianaceae), Geoffroea decorticans (Gill. Ex
Hook. et Arn.) Burkart (Fabaceae), Gomphrena pulchella Mart. (Amaranthaceae),
Grindelia pulchella Dunal. (Asteraceae), Hedeoma multiflorum Benth. (Lamiaceae),
Heliotropium amplexicaule Vahl. (Boraginaceae), Hypericum connatum Lam.
(Hypericaceae), Ipomoea purpura (L.) Roth (Convolvulaceae), Jodina rhombifolia
(Hook & Arn.) Reissek (Cervantesiacaceae), Larrea cuneifolia Cav.
(Zygophyllaceae), Larrea divaricata Cav. (Zygophyllaceae), Lepidium bonariense
L. (Brassicaceae), Ligaria cuneifolia (Ruiz et Pav.) Tiegh. (Loranthaceae), Lippia
integrifolia (Griseb.) Hieron. (Verbenaceae), Lippia turbinata Griseb. F.
(Verbenaceae), Lithraea molleoides (Vell.) Engl. (Anacardiaceae), Margyricarpus
pinnatus (Lam.) Kuntze (Rosaceae), Minthostachys mollis (Griseb.) Epling
(Lamiaceae), Mirabilis  jalapa L. (Nyctaginaceae), Passiflora  caerulea L.
(Passifloraceae), Perezia multiflora (Humb. & Bonpl.) Less spp. multiflora
(Asteraceae), Petunia axillaris (Lam.) Britton subsp. Parodii (Steere) Cabrera
(Solanaceae), Physalis mendocina Philippi (Solanaceae), Physalis viscosa L.
(Solanaceae), Plantago tomentosa Lam. ssp. Tomentosa (Plantaginaceae), Pluchea
36 3 Cuyo

sagittalis (Lam.) Cabrera (Asteraceae), Prosopanche bonacinae Speg.

(Hydnoraceae), Prosopis alba Griseb. (Fabaceae), Salix humboldtiana Wild.
(Salicaceae), Salpichroa origanifolia (Lam.) Thell. (Solanaceae), Satureja parvifo-
lia (Phil.) Epling (Lamiaceae), Schinus areira L (Anacardiaceae), Schinus
­fasciculatus (Griseb.) I.M. Johnst (Anacardiaceae), Senna corymbosa (Lam.) Irwin
& Barneby (Fabeaceae), Sida rhombifolia L. (Malvaceae), Sisyrinchium macrocar-
pum Hieron. (Iridaceae), Solanum eleagnifolium Cav. (Solanaceae), Solanum sisym-
brifolium Lam. (Solanaceae), Solanum argentinum Bitter et Lillo (Solanaceae),
Solidago chilensis Meyen (Asteraceae), Tagetes argentina Cav. (Asteraceae),
Tagetes filifolia Lag. (Asteraceae), Tagetes mendocina Phil. (Asteraceae), Tagetes
minuta L. (Asteraceae), Talinum paniculatum (Jacq.) Gaertn. (Portulacaceae),
Tessaria absinthioides (Hook. & Arn.) DC. (Asteraceae), Teucrium grisebachii
Hieron. (Lamiaceae), Tillandsia myosura Gris. & Baker (Bromeliaceae), Tillandsia
pedicellata (Mez) Castell. (Bromeliaceae), Trixis cacalioides (Kunth) D.  Don
(Asteraceae), Typha dominguensis Persoon (Typhaceae), Urtica dioica L. var. mol-
lis (Steudel) Wedd. (Urticaceae), Usnea densirostra Taylor (Usneaceae), Valeriana
ferax (Griseb.) Höck (Valerianaceae), Verbesina encelioides (Cav.) Benth & Hook.
Ex Gray (Asteraceae), Wedelia glauca (Ortega) Eifert (Asteraceae), Xanthium spi-
nosum L. (Asteraceae), and Zuccagnia punctata Cav. (Fabaceae), among other
species (Bustos et  al. 1996; Del Vitto et  al. 1997; Barboza et  al. 2009; Martínez
Carretero 2013; Agüero 2017).

3.2  A
 loysia gratissima (Gillies & Hook. ex Hook.) Tronc. var.
gratissima (Verbenaceae)

Synonymy  Aloysia lycioides Cham., Lippia lycioides (Cham.) Steud., Lippia

ligustrina (Lag.) Kuntze var. paraguariensis Briq., Verbena gratissima Gillies &
Hook. Ex Hook.

Vernacular name  Arrayán, arrayán del campo, azahar del campo, cedrón, cedrón
del monte, azar del monte, favorita, palo, muña del monte, palo amarillo, usillo,
niño rupá.

General information  A. gratissima grows in xerophilous environments and dry

forests from BAI, CAT, COR, COS, ERI, FOR, LPA, MEN, MIS, SDE, SFE, SJU,
SLU, and TUC. It is also found in Paraguay and Uruguay. It is an aromatic species
with a lemony scent. Also, it is a melliferous and ornamental species ideal for urban
gardens in temperate or temperate-warm climate (Haene 2007; Cané 2016).

Botanical characteristics  A. gratissima is an aromatic suffriticose or shrubby

plant, highly branched, 1–3 m in height, and 4–5 m in diameter. Stems are four-­
angled when young and rounded in maturity. The leaves are simple, deciduous,
oblong-obovate, 5–35 mm in length, entire or 3- to 5-lobed by the base, opposite
3.2  Aloysia gratissima (Gillies & Hook. ex Hook.) Tronc. var. gratissima (Verbenaceae) 37

subsessile or briefly petiolate, densely pubescent at the inferior lamina, and glandu-
lous and glabrous by the central vein. The foliage is grayish, persistent, and with a
very fine texture. Inflorescences are spicate racemes or paniculate, with white,
zygomorphic fragrant flowers. The calix is gamosepale, tubular, four-lobed with a
larger anterior lobule, externally glabre, and 5 mm in length. The flowers have four
stamens, ovary with two locules, and one ovule in each locule. The fruits are dry
schizocarps and exceptionally drupaceous that in maturity separates in two meri-
carps. The seeds do not have endosperm (Domínguez 1928; Cabrera and Zardini
1978; Ragonese and Milano 1984; Rotman and Múlgura de Romero 1999; Alonso
and Desmarchelier 2015; Cané 2016; O’Leary et al. 2016).

Ethnobotanical information  A. gratissima is very used in homemade medicine.

Hieronymus (1882) has already reported the use of the infusion made of A. gratis-
sima cuttings to treat cold and stomach problems. Currently, the infusion is mainly
used as digestive, anti-catarrhal, sedative, or sudorific. It has also been commonly
used to alleviate symptoms associated with headache, bronchitis, and nervous sys-
tem disorders. Besides, the infusions made with the aerial parts are used to treat
varicose veins and dizziness. The infusion made from stems is reported as a treat-
ment for hypotension; those made from leaves, to sooth menstrual pains and against
hemorrhoids; and those combining flowers and leaves, as cardiotonic, stomachic,
carminative, and diaphoretic and to relieve stomachache (Toursarkissian 1980;
Andersen et al. 1997; Del Vitto et al. 1997; Rotman and Múlgura de Romero 1999;
Múlgura de Romero et al. 2002; Barboza et al. 2009; Sérsic et al. 2010; Zeni et al.
2014; Bernasconi Salazar et al. 2017).

Chemical data  The compounds found in the essential oil (EO) from the aerial
parts were α- and β-pinene, sabinene, α-terpinene, β-phellandrene, α- and β-thujone,
α- and β-caryophyllene, α-bisabolene, β-elemene, bicyclogermacrene, camphene,
cadinol, caryophyllene oxide, citral, copaenone, copaenol, chiysanthenyl acetate,
globulol, spathulenol, limonene oxide, limonene, linalool, viridiflorol,
10-­epicubebol, (E)-nerolidol, germacrene D, eucalyptol, pulegone, terpinyl acetate,
eugenol, myrcene, γ-terpinen-1-al- trans-pinocarveol, cis-pinocarveol, γ- and δ-
elemene, geranyl acetate, β-cubebene, globulol, α-humulene, alloaromadendrene,
γ-gurjunene, cubebol, elemol, germacrene B, and carotol. Other components that
have been found in the aerial parts were the sesquiterpenoids guaiol and bisabolol;
the triterpenoids amyrin, betulinic acid, oleanolic acid, and ursolic acid; the flavo-
noids apigenin-5-hydroxy-7,40-dimethyl ether, genkwanin, and luteolin-7,3-O,4-­
O-­trimethylether; the glycoside flavonol rutin; and the phenylethanoids verbascoside
and arenarioside (Soler et al. 1986; Barboza et al. 2009; Trovati et al. 2009; Benovit
et al. 2015) (Fig. 3.2).

Biological activities  The decoction, alcoholic extract, and hydroalcoholic extract

from the aerial parts tested against Enterococcus faecalis (19433), Escherichia
coli (11229), Rhodococcus equi (6939), Salmonella enteritidis (ATCC 11076),
and Staphylococcus aureus (ATCC 25923) showed inactivation capacity and/or
38 3 Cuyo

Fig. 3.2  Some chemicals found in essential oils (EOs) from aerial parts of Aloysia gratissima
(Gillies & Hook. ex Hook.) Tronc. var. gratissima

selective inhibition. The best results were attained with the alcoholic extract from
the fresh plant. Among the bacteria, Pasteurella multocida demonstrated more
sensibility to the three extracts, while S. aureus and E. faecalis presented higher
resistance (Souza and Wiest 2005). It was also reported that the hydroalcoholic
extract (30:70) has activity against B. subtilis (Duarte et al. 2004) and the alco-
holic extract against B. cereus (Demo et  al. 1998) and S. aureus (Oliva et  al.
2001). Furthermore, the EO has activity against Junín virus (responsible of hem-
orrhagic fever) and herpes simplex virus type 1 (HSV-1) (García et al. 2003). It
was reported that the EOs from the aerial parts have viricidal and nematicide
activity (Barboza et al. 2009).
The aqueous extract and its main component, the ferulic acid (46.95 mg 100 g−1),
attenuated quinolinic acid-induced hippocampus cell damage in mice when they
were pretreated with any of them (100  mg  kg−1). However, they did not prevent
clonic or tonic seizure incidence or its severity. The protection produced may
involve the glutamate transport modulation through an inhibitory effect on the
N-methyl-D-aspartate receptor (Zeni et al. 2014). The antidepressant activity appar-
ently involved N-methyl-D-aspartate (NMDA) receptors (Farahani et al. 2014).
Finally, an antiproliferative effect was reported. The infusion prepared with
leaves of A. gratissima (6 and 24 g l−1) collected in Rio Grande do Sul, Brazil, and
the corresponding essential oil obtained by hydrodistillation (0.25% on ethanol)
produced a decrease in the mitotic index on Allium cepa bulb cells after 24 h of
treatment indicating an antiproliferative activity (Hister et al. 2009). However, it has
to be considered that the essential oil composition differs according to the places the
species grows (Duschatzky et al. 2004; Ricciardi et al. 2005; Franco et al. 2007).
3.3  Lippia integrifolia (Griseb.) Hieron. (Verbenaceae) 39

3.3  Lippia integrifolia (Griseb.) Hieron. (Verbenaceae)

Synonymy  Lippia boliviana Rusby, Lippia boliviana Rusby var. angusta, Lippia
boliviana Rusby var. integrifolia Moldenke, Lippia integrifolia (Griseb.) Hieron.
var. beckii, Lippia turbinata Griseb. var. integrifolia Griseb.

Vernacular name  Pulco, poleo, inca-yuyo, incayuso, té del inca, manzanilla.

General information  L. integrifolia is a native species from Argentina that is on

risk of extinction. It grows in the provinces of CAT, COR, JUJ, LRI, SAL, SJU,
and TUC.

Botanical characteristics  L. integrifolia is a woody aromatic perennial shrub and

1–5 m high. Leaves are entire, green-light, 2–3 cm in length, and 4–6 mm wide,
ovoid-lanceolate in adulthood. The lamina border is slightly backwards curved with
a central prominent nerve in the inferior lamina. The flowers are white or pink-­
whitish, with a 5–6 mm corolla, originating in axillary globose-compressed clus-
ters. The fruit has two mericarps with exalbuminate seeds (Bassols and Gurni 1996;
Schiuma 1997; Rotman and Múlgura de Romero 1999; Zuloaga et al. 2008; Alonso
and Desmarchelier 2015).

Ethnobotanical information  In pre-Columbian America, L. integrifolia was con-

sidered as sacred and exclusively used by the upper Incas castes that thought that L.
integrifolia stimulated the development of mental powers. In the case of the original
settlers from the province of COR, the Diaguitas and Comechingones, they smoked
L. integrifolia without restrictions by castes, genre, or age. Currently, its popular use
is as nervous tonic, in cases of melancholy and sadness, neurasthenia and nerve
pain, and in cases of stress. The infusion made with leaves and flowers is used as
digestive, mostly when the disease has a nervous origin; to relieve the stomachache;
to treat empacho; as carminative; also as emmenagogue, anti-flu, and antitussive; in
bronchitis and flu; and as expectorant. The infusion prepared with the dried aerial
parts is used as stomachic, emmenagogue, tonic, and diuretic. It could be added to
the traditional beverage mate. It is also used for the elaboration of bitter nonalco-
holic beverages (Ratera and Ratera 1980; Toursarkissian 1980; Martínez Crovetto
1981; Del Vitto et al. 1997; Pascual et al. 2001; Barboza et al. 2009; Agüero 2017).

Chemical data  In the fresh EOs were found the compounds α-thujene, sabinene,
p-cymene, limonene, terpinen-4-ol, borneol, eremophyllene, γ-elemene,
β-caryophyllene, δ-elemene, α-humulene, african-1-en-6-ol, african-4,5-dione,
4,5-­seco africanone, spathulenol, S (+)-trans-nerolidol, 2-β-,9-α-2,6,6,9-­
tetramethyltricyclo[6.3.0.0 (2,4-)]undec-1-(8)-ene-6,7,11-dione, 1β-,2α-,8β-
2,6,6,10-tetramethylbicyclo[6.3.0]undec-5-en-2-ol(3α-hydroxy-6-­asteriscane),
1α-,7β-,9-a-1-hydroxy-3,6,6,9-tetramethylbicyclo[5.4.0]undec-3-en-­8 -on,
lippifoli1(6)-en-5-one, lippifoliane, 1,6-cis lippifolian-1α-ol-5-one, 1,6-trans
40 3 Cuyo

lippifolian-1-α-ol-5-one, lippifolil (6)-en-4-β-ol-5-one, trans-africanan-1α-ol,

african-5-en-1α-ol, β-caryophyllene oxide, 4,5-secoafricanan-4,5-dione, humulene
epoxide II, ketone 4,1,6-cis and 1,6-trans lippifolian-1α-ol-5-one, lippifoli1(6)-en-­
4α-­ol-5-one, integrifolian-1,5-dione, asteriscane, bicyclohumulendione, humulen-
dione, and trans humul (9E)-en-2,6-dione (Catalán et al. 1983, 1991, 1992; Dartayet
et  al. 1984; Terblanché and Kornelius 1996; Barboza et  al. 2009; Marcial et  al.
2016) (Fig. 3.3).

Biological activities  The choleretic and antispasmodic activity of the aqueous

extracts of L. integrifolia (250, 500 and 750 mg kg−1) was tested on rats. A dose of
500 mg kg−1 induced a significant and persistent increase (60.5%) in bile flow after
the first 15 min from the oral administration; the dose of 750 mg kg−1 was also effec-
tive. A significant increase of bile acids output and the antagonism of muscle con-
tractions induced by acetylcholine (Ach) and CaCl2 on rat isolated jejunum was
observed. Those effects were related to the content on mono- and dicaffeoylquinic
acids (0.10% w/v expressed as chlorogenic acid) and the possible inhibition of the
calcium influx into the cytoplasm (Speroni et al. 2003; Gorzalczany et al. 2008).
The aqueous extract (0.5, 1.0, 2.0 mg ml−1) has anti-inflammatory effects on
stomach cells (20–60%) and antiadhesive properties against Helicobacter pylori.
Infusions (24% w/w), decoctions (25% w/w), raw polysaccharides isolated from
decoctions (RPS 2.3%), and the phenolic fractions (PhF) stimulated cellular vital-
ity of human stomach cells (AGS cells). Those results could explain the popular
use of the herbal tea for digestive disorders (Marcial et  al. 2014). Also, it was

Fig. 3.3  Some chemicals found in Lippia integrifolia (Griseb.) Hieron

3.4  Minthostachys mollis (Griseb.) Epling (Lamiaceae) 41

reported a stimulation of innate immunity and potential radical scavenging by L.

integrifolia extracts (Marcial et al. 2014).
Furthermore, it was shown antimicrobial activity against Gram-positive and
Gram-negative bacteria (Mycobacterium tuberculosis) and fungi (Microsporum
gypseum, Trichophyton metagrophytes, T. rubrum). The EOs have antifungal activ-
ity against Aspergillus spp. (Bluma and Etcheverry 2008; Bluma et al. 2008a, b),
and the methanolic extracts showed a limited spectrum of action and a lower potency
against Microsporum canis, M. gypseum, Epidermophyton floccosum, Trichophyton
rubrum, and T. mentagrophytes (MICs between 250 and 1000 μg ml−1) (Muschietti
et al. 2005). Also, the organic and aqueous extracts have anti-trypanocide activity
against Trypanosoma cruzi and against Aedes aegypti (Gleiser et al. 2011). The EOs
have a moderate activity against mosquitos (Gleiser et al. 2011).
In vitro culture  L. integrifolia was micropropagated using nodal segments and
apices from a plant growing in a greenhouse as initial explants. They were trans-
ferred to vessels containing MS medium (Murashige and Skoog 1962) with 20 g l−1
sucrose, BAP (2.2 μM), and 7 g l−1 agar. The developed shoots were transferred to
rooting medium composed by MS half strength media with IAA (0.28 μM). Then,
the rooted plants were transferred to pots that were placed in a humidity chamber
first, and finally to a greenhouse. Also, the genetic variability was studied by ISSR
technique with the goal of developing a breeding program (Ianicelli et al. 2016).

Legal status  The Argentina Food Code contains some tea preparations and com-
mercial beverages containing incayuyo that are taken as aperitif (2013).

3.4  Minthostachys mollis (Griseb.) Epling (Lamiaceae)

Synonymy  Bystropogon kuntzeanum Briq., Minthostachys verticillata (Griseb.)

Epling, M. verticillata (Griseb.) Epling var. eupatorioides, b., Xenopoma verticillata
Griseb., M. mollis Griseb.

Vernacular name  Peperina, peperita, piperina, martin muña.

General information  Minthostachys verticillata (Griseb.) Epling (Epling 1939),

whose popular name is peperina, is nowadays defined in the Flora of Argentina as
Minthostachys mollis Griseb. (Zuloaga et al. 2008; van Baren et al. 2014); how-
ever, most of the literature still refers to peperina as M. verticillata. M. mollis is a
species with ethnobotanical, pharmacological, and commercial relevance that
grows as a bush in the central and northwestern regions of Argentina, in the prov-
inces of CAT, COR, JUJ, SAL, SDE, SLU, and TUC.  The leaves and aromatic
flower spikes have a distinct peppermint-like odor and are used as a condiment in
foods and as a food preservative.
42 3 Cuyo

Botanical characteristics  M. verticillata is an aromatic subshrub, 0.3–2.0  m in

height with highly branched pubescent quadrangular stems. Leaves are ovoid,
obtuse, with festooned edges, 1–5 cm in length. Flowers are white, small, and placed
as clusters in the axilla of the leaf. Fruits are small dry capsules with four tiny seeds
per fruit. Blooming is in summer. The fruiting period covers the months of March
and April (Núñez and Cantero 2000; Cariddi 2013).

Ethnobotanical information  Peperina is usually taken as an infusion for its attrib-

uted properties as digestive, or it is added, for its characteristic aroma and taste, to
the traditional beverage mate (Bandoni et al. 2002; González Pereyra et al. 2005;
Elechosa et al. 2007; van Baren et al. 2014). Also, it is added to yerba mate (Ilex
paraguariensis) for producing yerba mate compuesta and to liqueurs and aperitifs
(Valladares et al. 2002; Escobar et al. 2015). According to the dualistic classification
from Native Americans, it is considered a hot species (Schmidt-Lebuhn 2008).
Consequently, it is used in popular medicine to treat digestive diseases (empacho)
and as an aphrodisiac. Leaves mixed with flowers are used to prepare infusions with
recognized activities as stomachic, antispasmodic, antidiarrheal, antiemetic, anti-
rheumatic, carminative, sedative, hemostatic, and hypotensive. The infusion made
with dried leaf and stems is used as carminative, for improved fertility, and as sexual
stimulant (Del Vitto et al. 1997; Barboza et al. 2009).

Chemical data  Chemically, the oxygenated monoterpenes fraction represents

90% of the total M. verticillata EOs. The EOs from the aerial parts include pule-
gone, menthone, isomenthone, limonene, menthol, α-pinene and β-pinene, carvone,
carvacryl acetate, γ-terpinene, linalool, piperitenone, sabinene, myrcene, (E)-β-­
ocimene, thymol, camphene, p-cymene, α- and γ-terpinene, isopulegol, δ-cadinene,
β-caryophyllene, γ-gurjunene, and τ-cadinol (Fester and Martinuzzi 1950; Fester
et al. 1960; De Feo et al. 1998; Escobar et al. 2012). The EOs from the leaves have
β-caryophyllene: sesquiterpene and myrcene (monoterpene (E) β-ocimene).
However, in all cases the main components are pulegone (cyclo-hexanone, 5-methyl-­
2-(1-methylethylidene) (61%) and menthone (cyclohexanone, 5-methyl-2-(1-­
methylethyl) (27%) (Valladares et  al. 2002; Escobar et  al. 2012; Pellegrini et  al.
2017) (Fig. 3.4).

Biological activities  It was reported that M. verticillata extracts and its EOs have
antimicrobial, insecticidal, acaricidal, and antifungal properties (Primo et al. 2001;
Valladares et  al. 2002; Bluma et  al. 2008b; Palacios et  al. 2009; González and
Marioli 2010; Vogt et al. 2010a; Cariddi et al. 2011; Escobar et al. 2012; Escobar
et al. 2015; Pellegrini et al. 2017). Consequently, it was proposed their use in food
preservation and also as insecticides. The antimicrobial activity was demonstrated
against Aspergillus section Flavi, Streptococcus uberis (isolated from bovine masti-
tis), and P. larvae (responsible of American foulbrood in Apis mellifera). Aspergillus
section Flavi growth rate was tested at different EOs concentrations (0, 150, 300,
500, and 700 mg kg−1) being weakly inhibited by the EOs obtained by hydrodistil-
lation at all the water activities (0.982 and 0.955 aw) tested. The lowest ­concentration
3.4  Minthostachys mollis (Griseb.) Epling (Lamiaceae) 43

Fig. 3.4  Some of the chemical compounds found in Minthostachys verticillata (Griseb)

of these EOs showed growth rate stimulation in the aflatoxigenic strains, while the
highest concentrations reduced sporulation and vegetative mycelia in aflatoxigenic
growth zones (Bluma et al. 2008a). When the EOs and limonene were tested against
S. uberis, the minimum inhibitory concentration (MIC) values ranged from 14.3 to
114.5  mg  ml−1 (1.56–12.5% v/v) and the minimum bactericidal concentration
(MBC) from 114.5 to 229 mg ml−1 (12.5–25% v/v). MICs for limonene ranged from
3.3 to 52.5 mg ml−1 (0.39–6.25% v/v) and MBC from 210 mg/mL (25% v/v). The
aqueous extracts, namely, the water remaining after hydrodistillation (WRHD), and
decoctions (D) showed the highest antibacterial activities (González and Marioli
2010). Gram-positive bacteria were more sensitive to inhibition by the EOs than the
Gram-negative bacteria (Primo et  al. 2001). The antimicrobial capacities of the
aqueous extracts (5 mg of aerial parts per 100 ml distilled water) and the EOs deter-
mined against P. larvae showed a minimal inhibitory concentration (MIC) of
75  μg  ml−1 against P. larvae with inhibition zone diameters in a range between
10.0 mm and 26.0 mm for EOs. On the other hand, the aqueous extracts produced
100% inhibition (100% efficacy) (Pellegrini et  al. 2017). As WRHD is normally
discarded after the hydrodistillation procedure, the authors proposed that the WRHD
fraction could be reserved for apiaries sanitation, while EOs, for its lower biological
activity, could be used for cooking or in fragrances. However, analysis of toxicity on
honey bees as well as of WRHD delivery and/or application methods must be per-
formed (Pellegrini et al. 2017). The in vitro antiviral activity of the EOs was tested
against Suid herpesvirus type 1 (HSV-1) and Pseudorabies virus (PrV) and quanti-
fied by the plaque reduction essay with HEp-2 cells. The antiviral activity specifi-
cally affects HSV-1 and PrV multiplication at pulegone and EOs concentrations 20
to 30 times lower than the cytotoxic concentration values (CC50). Both inhibited
the replication of Suid herpesvirus type 1 in more than 50% and exerted an i­ nhibition
44 3 Cuyo

over 90% at a concentration of 100  μg  ml−1. The EC50 were 20.25  μg  ml−1 and
20.00  μg  ml−1, and the CC50 were 613 and 352  μg  ml−1 for EOs and pulegone,
respectively. Menthone and limonene had little and not selective antiviral action.
The therapeutic index (TI = CC50/EC50) were 30.3 for EOs and 17.6 μg ml−1 for
pulegone. The authors proposed the use of EOs and pulegone as therapeutic selec-
tive antiviral agents (Vogt et al. 2010a). As for its activity as insecticide, leaves of
M. verticillata that were affected by the leaf miner Chromatomyia platensis
(Brethes) (Diptera: Agromyzidae) and a cecidomyid gall insect (Diptera:
Cecydomyiidae) showed a decrease in pulegone concentration (42.8%) and an
increase in menthone concentration that almost duplicated the amounts found in no
mined leaves (25.12%). No variation was observed in galled stems. The results also
suggested a differential response according to the feeding habits of the insects
(Valladares et al. 2002). When tested against M. domestica, R (+)-pulegone showed
high insecticidal effects with LC50 1.7  mg  l−1, and menthone showed LC50
8.6 mg l−1, while limonene had LC50 6.2 mg l−1 (Palacios et al. 2009). There are
also references about the effects of D and EOs on the immunological system
(enhancing or modulating the immune response). The ability of D (210  μg  ml−1,
104 μg ml−1, 10.4 μg ml−1, and 1.04 μg ml−1) and EOs (0.96 μg ml−1, 0.48 μg ml−1,
48 μg ml−1, and 4.8 μg ml−1) to induce or inhibit in vitro basophil degranulation was
tested with human basophils from dust mite non-allergic and allergic patients. The
IL-13 levels were evaluated from lymphocyte cultures stimulated with only an aller-
gen or with a combination of the main monoterpenes found in the EOs (pulegone,
menthone, and limonene). IL-13 synthesis was quantified in supernatants of lym-
phocyte cultures from allergic patients stimulated with allergen (10,000 PNU ml−1)
alone or combined with EOs (6  μg  ml−1), pulegone (62  μg  ml−1), menthone
(60 μg ml−1), limonene (55 μg ml−1), or the combination of the three monoterpenes.
That combination and limonene alone were the most active to reduce IL-13 and to
stimulate cell proliferation. On the other hand, EOs and monoterpenes reduced the
μ-hexosaminidase release from basophils of allergic patients maybe acting as a
membrane stabilizer. The greatest inhibitory effect was for EOs (10 μg ml−1), pule-
gone (40 μg ml−1), menthone (40 μg ml−1), and limonene (20 μg ml−1). The inhibi-
tory effect with the EOs was highest than the one obtained with the single
monoterpenes or their combination. Furthermore, the anti-allergic effect was
reported to be higher than with desloratadine (Cariddi et  al. 2011). On the other
hand, lymphocyte-proliferating activity of D and EOs was tested and compared
with cellular expansion induced by phytohemagglutinin (PHA), Pokeweed mitogen
(PWM), and bacillus Calmette-Guérin (BCG) with a non-stimulated culture as con-
trol reference. The cytomorphological analysis showed mitogenic activity and
induced lymphocyte cluster and colony formation similar to PHA and PWM and
higher than CGB (p < 0.02). A 40% of the cells that proliferated were LT CD8 (+);
also the basophils from allergic individual resisted up to a concentration of
104 mg ml−1of D and 0.48 mg ml−1 of EO (González Pereyra et al. 2005).

Toxicity  Cytotoxicity and genotoxicity were tested against Vero cells, human
peripheral blood mononuclear cells (PBMCs), and mice bone marrow cells. No cyto-
toxicity was observed in Vero cells (10–1000 μg ml−1). The viability of PBMCs was
3.5  Acantholippia seriphioides (A. Gray) Moldenke (Verbenaceae) 45

72% with the highest concentration of EOs (1000 μg ml−1), and no signs of apoptosis
were found at the concentrations tested. EOs did not induce cytotoxicity in Vero cells
and human PBMCs. In addition, EOs (100–1000 μg ml−1) do not induced apoptotic
effects on human PBMCs. No clinical signs of toxicity, mortality, or changes in body
weights or food consumption were reported. There is no any toxicological effect in
the liver, kidney, or intestine of Wistar rats up to an EOs concentration of 7 g kg−1
feed for 90 days. The in vivo assay showed that EOs (25–500 mg kg−1) do not increase
the frequency of micronucleus in bone marrow cells of mice. Based on those results
where no cyto-genotoxic effect of EOs was found in vitro or, the authors conclude
that EOs are safe as therapeutic agent (Escobar et al. 2012).

Legal status  M. verticillata is among the nontoxic vegetal drugs authorized for
their use with medicinal purposes. It was accepted as an official drug in the FNA 6th
edition.

3.5  A
 cantholippia seriphioides (A. Gray) Moldenke
(Verbenaceae)

Synonymy  Lippia seriphioides A. Gray, Lippia foliolosa Phil.

Vernacular name  Tomillo, tomillo andino, tomillo de la sierra, tomillo del campo,
tomillo mendocino, tomillo macho, tomillo de olor, tomillo hembra, ñancuñán,
tomatillo silvestre.

General information  It grows in Chile, Bolivia, and in Argentina in the provinces

of BAI, CHU, LPA, MEN, NEU, RNE, SCR, SJU, and SLU in arid environments
characterized by stony or saline soils, from sea levels up to 1000 m.a.s.l. (Van Baren
et  al. 2015). The intense overexploitation of medicinal and aromatic plants has
placed this species at risk of extinction (Delucchi 2006; Elechosa et al. 2009). It is
used as flavor of foods and as a condiment in replacement of the authentic thyme
(Thymus vulgaris L.) for its typical high content of thymol and carvacrol (Ruiz Leal
1972; Roig 2001; Gastaldi et al. 2016). Its propagation could be achieved uneasily
through seeds or cuttings, but the most common way in nature seems to be the spon-
taneous layering (Strasser et al. 2002; Strasser and Fernández 2006).

Botanical characteristics  A. seriphioides is an aromatic bush 30–60 cm in height,

branched from the basis, with 2–6 mm sessile leaves, 3–5 lobed in the apex, narrow
towards the base, and with reddish spherical glands more abundant on the ­underside.
Leaves are pubescent or subglabrous, opposite, sessile, and with a fasciculate appear-
ance due to its axillar short braquiblast (van Baren et al. 2015). The white inflores-
cences are clustered at the end of the branches. The calix is tubular with two-­toothed
or entire lobules and 2.5–3.0 mm in length with white simple hairs. The corolla has
five uneven lobules with a cylindric tube 3.0–3.5 mm in length. The fruit is formed
by two fused achenes, with mericarps, and around 2 mm in length (Caro 1982).
46 3 Cuyo

Ethnobotanical information  The infusions prepared with the aerial parts are used
as a stomachic, gastrointestinal, and diaphoretic and against the influenza. Infusions
made with leaves and stem are taken as aphrodisiac (Del Vitto et al. 1997; Ratera
and Ratera 1980; Toursarkissian 1980; González and Molares 2004).

Chemical data  Hydrodistillation from aerial parts yielded between 0.57 and
2.46% (v/w) EOs that have thymol, p-cymene, carvacrol, γ-terpinene, citral, and
piperitone (Fig. 3.5). On the basis of the 13 main compounds of the EOs determined
by GC-FID-MS, 4 chemotypes were determined: carvacrol, thymol, cis- and trans-­
dihydrocarvone, and linalool-geraniol. The co-occurrence of different chemotypes
in a same population was found indicative of a pattern of variation throughout the
natural distribution area of this species (Van Baren et al. 2015).

Biological activities  The infusion, tinctures of dry plant material, and the EOs
have antioxidant activity (determined by the DPPH assay) attributed to its content
on thymol and carvacrol (Gastaldi et al. 2016). The plant EOs have antioxidant
and antimicrobial activity (Ruffinengo et al. 2005; Fuselli et al. 2007; Gillij et al.
2008; Lima et al. 2011). The A. seriphioides EO showed a high acaricidal effect,
causing high mite mortality at low oil concentration, but at the same time it had a
low selectivity ratio (LD50 mite = 1.2 μl per cage, ratio selection 1.3 for 48 h of
treatment). The main compounds of this EO are thymol, carvacrol, and orthocy-
mene. Thymol and carvacrol are effective in control of Verroea destructor
(Ruffinengo et al. 2005).

Fig. 3.5  Some of the chemical compounds found in the essential oil from Acantholippia seriphi-
oides (A. Gray) Moldenke
3.6  Achyrocline satureioides (Lam.) D.C. (Asteraceae) 47

3.6  Achyrocline satureioides (Lam.) D.C. (Asteraceae)

Synonymy  Achyrocline vargasiana DC., Gnaphalium satureioides Lam. candi-

cans (Kunth.).

Vernacular name  Macela, marcela hembra, marcela, marcela del campo, marcel-
ita, planta de la abeja, vira-vira, vira-vira guazú, yateí caá.

General information  A. satureioides is one of the most spread medicinal species

from Argentina; its popularity comes from its use for the treatment of a variety of
disorders, principally gastrointestinal and respiratory diseases (Toursarkissian 1980;
Ratera and Ratera 1980; Gattuso et al. 2007). The species grows mostly in r­ angelands
and modified soils in the provinces of BAI, CAT, CHA, COR, COS, ERI, JUJ, LPA,
MIS, SAL, SDE, SFE, SLU, and TUC (Barboza et al. 2009). Also, it is an attractant
to butterflies which participate in its propagation. A. satureioides is also much
appreciated for their delicate perfume with a light trace of anise, although the flavor
is astringent. The stems, leaves, and flowers contain resins, essential oils, tannins,
and other bitter principles (Ratera and Ratera 1980). For all of that, marcela is use-
ful both for the pharmaceutical and for the cosmetics and bitter beverages industries
(Del Vitto et al. 1997; Gattuso et al. 2007; Barboza et al. 2009; Retta 2014).

Botanical characteristics  A. satureioides is a perennial herbaceous species and

about 0.30–1.20 cm in height. It has stiffened stems covered by short white hairs.
Leaves are grayish, sessile, alternate, linear, elliptic, or lanceolate, with an entire
margin, and densely tomentose on the underside, 3–5 cm in length. The inflores-
cences are small capitula distributed in terminal glomerulus with 3–6 marginal
feminine filiform flowers and 1–3 central tubular hermaphrodite flowers. Anthers
are sagittate; style is short and truncated by the apex. Fruits are small brown achenes
with a white pappus with faint hairs and a naked receptacle. Blooming is by the end
of summer (Cabrera and Zardini 1978; Gattuso et  al. 2006, 2008; Alonso and
Desmarchelier 2015).

Ethnobotanical information  A. satureioides was very popular among native com-

munities that had been taken it as a digestive tisane. Nowadays, the aerial parts have
a popular use in Argentina, Uruguay, Brazil, and Paraguay as a decoction or infusion
for treating miscellaneous diseases. It is considered beneficial in cases of
­gastrointestinal disorders, as antidiabetic, digestive, antibacterial, antispasmodic,
anti-inflammatory, antitumor, and antidiarrheal, to treat asthenia, and as tonic and
stimulant. The mixture of dried inflorescences and stems is used as sedative, antisep-
tic, antispasmodic, for inflammation, and to aid and promote menstruation. The infu-
sion prepared with the leaves is administrated as antitussive in bronchitis and the
mixture of leaves and flowers as digestive, carminative, antispasmodic, sedative,
anti-asthmatic, antitussive, and hypoglycemic. The infusion prepared with the flow-
ers is used to treat asthma and as analgesic, antispasmodic, and anti-inflammatory.
48 3 Cuyo

As an external prescription, leaves are applied as anti-inflammatory in compresses

and washes (Ratera and Ratera 1980; Toursarkissian 1980; Alonso and Desmarchelier
2015; Agüero 2017).

Chemical data  This species has a high concentration of polyphenols and flavo-
noids (Fig. 3.6). In the dried entire plant, it was identified achyrofuran, quercetin,
caffeic acid, chlorogenic and isochlorogenic acids, galangin, 3-methoxy galangin,
quercetin-3-methylether, 2, 4′, 4-trihydroxy-6-methoxychalcone, and 7,
4′-dihydroxy-­ 5-methoxyflavone. Quercetin, luteolin, 3O-methyl quercetin, and
3,7-dimethoxy-5,8-dihydroxyflavone were found in the dried inflorescence (Braga
et al. 2006; Barboza et al. 2009; Grassi-Zampieron et al. 2009a; Carini et al. 2014;
Retta 2014; Carini et al. 2015; Oliveira de Souza et al. 2018).
The inflorescences EOs contain monoterpenes (<7.3%) and sesquiterpenes (73.8%)
(Fig.  3.7). Their main components were caryophyllene (39.7%), δ-canadiene

Fig. 3.6  Some polyphenols and flavonoids detected in Achyrocline satureioides (Lam.) DC
3.6  Achyrocline satureioides (Lam.) D.C. (Asteraceae) 49

Fig. 3.7  Main components of the essential oils from Achyrocline satureioides (Lam.) DC

(9.4%), and α-copaene (14.2%). On the contrary, in Brazil the main components
were monoterpenes (>56.4%) (Labuckas et al. 1999).
Biological activities  It was reported the antioxidant and free radical scavenging
activities of the A. satureioides aqueous and methanolic extracts (Desmarchelier
et al. 1998; De Souza et al. 2002; Grassi-Zampieron et al. 2009b). The antioxidant
activity was confirmed by  in silico evaluation (PASS analysis) (Salgueiro et  al.
2016). The ability of A. satureioides major compounds (such as the flavonoids lute-
olin, quercetin, 3-O-methylquercetin, and achyrobichalcone) to protect rat brain
homogenates from lipid damage was reported (Rivera Megret et al. 2013). The con-
tent on those compounds was related to the hepatoprotective, immune-stimulant,
cytotoxic, vasodilatory, antiherpetic, antispasmodic, relaxant, anti-ulcerative, anti-
microbial, antioxidant, molluscicidal, antiviral, anti-inflammatory, and mutagenic
activities attributed to the species.
The effect on skin lipoperoxidation was tested using a topical nanoemulsion
made with A. satureioides extract or with pure quercetin on porcine skin from the
back part of the ear. The nanoemulsion was prepared with A. satureioides inflores-
cences (7.5 w/v) in ethanol (80%) at a total amount of 100 μg of quercetin. The
level of quercetin retained in the skin was higher with the A. satureioides extract-­
containing nanoemulsion than with the quercetin-containing nanoemulsion. Also,
the thiobarbituric acid-reactive species (TBA-RS) assay showed a better result
with the extract-containing nanoemulsion. The higher effectivity to avoid lipoper-
oxidation from the A. satureioides extract-containing nanoemulsion was attributed
to the synergic effect of the compounds in the extract (Zorzi et al. 2015). An anti-
oxidant hydrogel containing A. satureioides extract-loaded nanoemulsion (total
flavonoids content, 1 mg ml−1) was formulated to prevent oxidative stress on the
skin (Balestrini et al. 2016).
The immunosuppressant, antioxidant, and anti-inflammatory activities of the
dried inflorescences have also been attributed to the presence of flavonoids, mainly
quercetin and its derivatives (Desmarchelier et  al. 1998; Rivera Megret et  al.
2013). However, when flower extracts were obtained by supercritical fluid extrac-
tion, its antioxidant activity did not show dependence on the phenolics concentra-
tion, suggesting the presence of other non-phenolic substances with antioxidant
50 3 Cuyo

properties (Peruchi et al. 2005). The maximum antioxidant activity (81.6 ± 5%)

was obtained at 250 bar and 30 °C, and the maximum amount of phenolics content
(85  ±  3  mg gallic acid equivalent g−1 of extract) was recovered at 200  bar and
30 °C (Peruchi et al. 2005). As for their mechanism of action in tissue inflamma-
tion, the infusion (0.06–0.24 μg ml−1 quercetin equivalent) reduced phytohemag-
glutinin (PHA)-induced proliferation and production of interferon (IFN)-γ and
interleukin (IL)-4. It also slightly increased the spontaneous generation of reactive
oxygen species (ROS) from a concentration of 0.06 μg ml−1 quercetin equivalent
and inhibited N-formyl-­Met-Leu-Phe (fMLP)-induced ROS generation as well as
spontaneous and fMLP-­induced IL-8 production between 0.0012–0.03  μg  ml−1
quercetin equivalents (Cosentino et al. 2008). Additionally, it was reported that the
treatment of male Wistar rats with A. satureioides aqueous and ethanolic extracts
reduced the neutrophil influx and the secretion of leukotriene B4 (LTB4) and cyto-
kine-induced neutrophil chemoattractant 1 (CINC-1) in the exudates and the num-
ber of rolling and adhered leukocytes in the mesentery postcapillary venules,
neutrophil L-selectin, 𝛽2-integrin, and toll-like receptor (TLR-4) expression and
the oxidative burst, but did not cause an alteration in the morphology and activities
of the liver and kidney. Together, the data showed that A. satureioides extract
inhibited neutrophil functions related to the innate response and did not cause
systemic toxicity (Barioni et al. 2013).
Another interesting activity of the hydroalcoholic extract of A. satureioides (14.8
equivalents of caffeic acid and 3 mg ml−1 quercetin) is the improvement of sleep
quality in patients with obstructive sleep apnea syndrome (OSAS) that have received
an oral administration of the extract. As sleep apnea might enhance oxidative stress
through a reduction of antioxidant capacity of the blood due to hypoxia, the positive
effect could be related to the antioxidant and anti-inflammatory actions attributed to
the polyphenolic content of the extract (Pedemonte et al. 2013).
To test the neuroprotective effect, a model of permanent middle cerebral artery
occlusion (pMCAo) with 2,3,5-triphenylltetrazolium chloride (TTC) was assessed
in rats. When a decoction of dried flowers (2% w/v) was administrated to the rats
during the 21st days previous to the treatment with TTC, the functional deficit
caused by the tetrazolium salt was reversed and the cerebral infarct volume dimin-
ished at a plasma level of 60.4 ± 58.2 ng ml−1 quercetin. That effect was partially
attributed to the content in flavonoids, mainly quercetin (1.14 ± 0.19 mg ml−1) and
luteolin (Rivera Megret et  al. 2013). An infusion of the aerial parts (2% w/v)
­triggered a differentiation effect in PC12 cells at 10 and 20 μg ml−1 of total polyphe-
nols. In that work it was demonstrated an increase of the number of neurites per cell,
the percentage of cells with neurites, the number of fusiform cells, and the percent-
age of differentiation, which translated into an increase in long neurites. The changes
induced resembled to those produced by nerve growth factor treatment. However,
the differentiation features observed would likely not only be related to flavonoids
(Blasina et al. 2009).
The popular use of the aqueous extract of A. satureioides as a hepatoprotective
and digestive agent could be mediated through its antioxidant and choleretic activi-
ties (Kadarian et al. 2002; Rahimi 2015). Additionally, extracts of A. satureioides
3.6  Achyrocline satureioides (Lam.) D.C. (Asteraceae) 51

whole plant material were significantly active in in-house type 2 diabetes model on
genetically diabetic db/db mice (oral dose 20  mg  kg−1 q.d). The hypoglycemic
activity was attributed by the authors to the compound achyrofuran as its oral
administration (20 mg kg−1) significantly lowered blood glucose levels (Ruffa et al.
2002; Carney et al. 2002). It was proposed the inclusion of the extracts in food for-
mulations to treat (or prevent) type-2 diabetes and to ease the digestion process
(Peruchi et al. 2005).
The achyrobichalcone, isolated from A. satureioides (Holzschuh et  al. 2010;
Carini et al. 2012), could also represent a promising chemotherapeutic biomolecule
considering its similarity with other cytotoxic bi-chalcones (Carini et al. 2014). In
general, the effects (attributed to the flavonoid content) were inhibition of cell pro-
liferation, cell cycle arrest, apoptosis, and inhibition of angiogenesis, prevention of
migration/metastasis, and overcoming multidrug resistance, alone or in combina-
tion with commonly used chemotherapeutic drugs (Oliveira de Souza et al. 2018).
Moreover, the flavonoid extract potentiated the activity of temozolomide, a chemo-
therapeutic drug. The positive results of these preclinical studies suggest a promis-
ing use of these flavonoids for the development of new anticancer drugs (Oliveira de
Souza et al. 2018).
The antimicrobial effect against Staphylococcus strains isolated from patients
with youthful acne and from clinical materials (prosthesis, etc.) was tested with a
sterile decoction (5% w/v final concentration) prepared from leaves from samples
collected in the province of Córdoba. The inhibitory effect was demonstrated on
coagulase-negative non-hemolytic strains (95%). Consequently, the topic applica-
tion of the decoction to reduce the staphylococcal population was proposed (Calvo
et al. 2006). The decoction of A. satureioides aerial parts was also active against S.
aureus, E. coli, and A. niger (Anesini and Pérez 1993). The synergistic antibacterial
activity of different components of the ethanolic extract (quercetin, 23-methyl-6-O-
desmethylauricepyrone, and 3-O-methylquercetin) was also reported (Joray et al.
2013). It was also referred an activity (14.3 mm diameter of inhibition) of the etha-
nolic inflorescences extract on Streptococci viridians strains found in the oral cavity
(Massori et al. 2017). The aqueous, freeze-dried, and spray-dried extracts prepared
from inflorescences were tested for their activity on pathogenic intestinal flora of
male Wistar rats. The freeze-dried extract prepared from the 80% ethanol (v/v)
extractive solution exhibited a broad spectrum of antimicrobial activity (100–
200  mg  ml−1) being the antibacterial effects of the extracts higher than those of
amoxicillin (used as a positive control) when tested against Bacillus cereus (ATCC
9634) and Staphylococcus aureus (ATCC 1901). Total flavonoid content was
132 mg g−1 for the freeze-dried extract, 129.7 mg g−1 for the spray-dried extract, and
54.23 mg g−1 for the freeze-dried aqueous extract. The reported results also sug-
gested that 80% ethanol (v/v) is more efficient for extracting the three flavonoid
aglycones and chalcone from the inflorescences than water alone (Moresco et al.
2017). The n-hexane and dichloromethane extracts of the aerial parts contained,
besides the compounds already mentioned, prenylated phloroglucinol α-pyrones
that showed antimicrobial activity (against E. coli ATCC35218, S. aureus VISA, S.
aureus MSSA, E. faecalis ATCC29212, and yeast sp.) that was related by the
52 3 Cuyo

authors to the 3-methyl-2-butenyl chain attached to the phloroglucinol nucleus

(Casero et al. 2014).
On the other hand, an antiherpetic activity was also informed (Bidone et  al.
2015). To locally treat HSV-1 infection, the ethanolic extract of A. satureioides was
incorporated into nanoemulsions (polydispersity index <0.1, mean droplet size
200  nm) with flavonoid association efficiency of almost 100%. The antiherpetic
activity of the nanoemulsion (307.79 ± 10.23 μg ml−1 quercetin, 194.50 ± 7.91 μg ml−1
luteolin, 780.35 ± 31.42 μg ml−1 3-O- methyl quercetin) tested by the viral plaque
number reduction assay on Vero cells monolayers was significantly higher (IC50
1.4  ±  0.88  μg  ml−1) than that corresponding to the ethanolic extract (IC50
14.07 ± 3.46 μg ml−1) (Bidone et al. 2014; Bidone et al. 2015).
As for the anti-plasmodial effect, it was reported that the treatment with A. satu-
reioides had control of Trypanosoma evansi parasitemia in Wistar female rats. The
number of trypomastigotes forms diminished both with an oral administration dose
(1.5 ml kg−1) of the EO (around 33% decrease) or the nano-encapsulated EO (80%
decrease). On the other hand, the nano-encapsulated essential oils showed a reduc-
tion of the intense inflammation response caused by T. evansi and resulted more
protective than the EOs on the liver and kidney (Do Carmo et al. 2015). Furthermore,
at a concentration of 250 μg ml−1, the ethanolic extract showed 100% of lysis on
bloodstream of T. cruzi in vitro (Rojas de Arias et al. 1995).
It was reported a relevant inhibition of Macrophomina phaseolina growth with
A. satureioides aqueous, hexane, and chloroform extracts (1000 μg ml−1). The hex-
ane and chloroform extracts were also inhibitory against Fusarium graminearum, F.
verticillioides, and Sclerotium rolfsii. Nevertheless, the inhibition of F. solani was
minimal (< 20%) in the conditions reported (Vogt et al. 2010a, b).
In veterinary, it was showed that the decoction from inflorescences (75 mg ml−1)
has inhibitory activity on bacteria isolated from bovine mastitis after 24 h of con-
tact. From the bacteria isolated, 20 were Gram-positive (14 Staphylococcus aureus,
5 Streptococcus dysgalactiae, 1 Streptococcus uberis), and 3 were Gram-negative
(1 Escherichia coli, 1 Klebsiella sp., and 1 Pseudomonas aeruginosa) (Rocha
Sperotto et al. 2012).
As for its potential interest for the food industry, hydroalcoholic extract of A.
satureioides (0.5 and 1%) was effective in decreasing the lipid oxidation of
salami during storage without interfering with the acceptance of the product at a
0.5% concentration (Campagnol et  al. 2011). Other authors also reported the
antimicrobial activity of A. satureioides decoctions and hydroalcoholic extracts
against bacteria of interest in food (S. aureus, E. faecalis, S. enteritidis, E. coli)
being more sensitive the Gram-positive (Staphylococcus and Entherococcus)
bacteria than the Gram-negative (Salmonella and Echerichia) (Mota et al. 2011).
The crude hydroalcoholic extract (5% w/v hydroalcoholic extract) deactivated all
strains of Salmonella spp. resistant to antibiotics isolated from raw material from
swine and poultry (Maciel et al. 2017).
In vitro cultures  A. satureioides can be regenerated in vitro in Murashige and
Skoog (MS) ¾ media with benzilaminopurine (BAP, 1 mg l−1) and 3% w/v sucrose
References 53

with uni-nodal segments (5–10 mm) as explant. Once the plant developed, rooting
was induced by adding indole acetic acid (IAA, 1 mg l−1). After 84 days of culture,
the yield of rooted plantlets obtained was 62.35%. After transferring to pots in
greenhouse, the survival was 77% after 150 days (Gattuso et al. 2007).

Toxicity  The species is widely consumed by South America population without

any report of toxicity. However, several studies were performed in order to assess
that information. The maximum tolerated dose, determined by a sub-chronic toxic-
ity study in mice and rats with two administration routes (oral and intraperitoneal),
was 5 g kg−1 without any evidence of toxicity in the liver or kidney (Rivera et al.
2004). Further studies have shown that the cold aqueous extract (10–50 μg ml−1)
have low toxicity; it did not induce apoptotic effects on human peripheral blood
mononuclear cells and did not show in  vitro genotoxicity against Vero cells at
10–50  μg  ml−1; however, at high concentrations the micronucleus assay showed
cytotoxicity (Sabini et  al. 2013). When the same authors tested the hot aqueous
extract effect, they found that it was more toxic than the cold one; however, it did
not induce apoptosis. As for the cytogenotoxic effect, it was positive on bone mar-
row of mice (Cariddi et al. 2015). On the other hand, a cold aqueous extract (0.5,
1.0, 2.0, 3.0, and 4.0 mg ml−1 of mineral water) was used to study cytotoxicity by
the Allium cepa L test. Only the higher concentrations induced cytotoxicity, but the
effect was reversible and not associated with mutagenicity (Sabini et al. 2011).
Other studies have demonstrated that the decoction showed a certain cytotoxic
activity on human lymphocytes at a concentration of 5% w/v (Calvo et al. 2006) and
that the methanolic extract has a toxic effect against Hep G2, a human hepatocel-
lular carcinoma cell line (Ruffa et al. 2002). Finally, the A. salina bioassay showed
a LD50 of 2.06  mg  ml−1, which is considered very low for plant extracts. When
different extract concentrations (30, 60, 150, and 300 μg ml−1) were tested on human
lymphocytes, none of them caused damage to DNA of cells, suggesting that the
extracts have low potential genotoxicity (Salgueiro et al. 2016).
Legal status  A. satureioides is an official vegetable drug in the Brazilian
Pharmacopoeia (Retta et  al. 2012). A quantification of the active markers was
­validated in order to propose it for its pharmacopoeia quality control in Argentina
(Retta et al. 2007).

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Chapter 4
The Pampa: An Infinite Plain

4.1  Introduction: The Pampa

The Pampa plain (Fig. 4.1) is an extensive surface with undulations (from ancient
dunes) that comprise the province of BAI; North East of the province of LPA; South
of the provinces of COR, SFE, and ERI; and sections of the provinces of SLU,
RNE, and MEN. It limits to the North with the Mesopotamian region and the CHA
plain, by the West with the Sierras Pampeanas and the Cuyo region, by the South
with the Colorado River and Patagonia, and by the East with the Atlantic Ocean, the
Río de la Plata, and the Paraná River. The plain is interrupted towards the South by
the systems of Sierras de Tandilia and Sierra de la Ventana (500–1300 m height).
Sierras de Tandilia extends from the city of Mar del Plata to Las Flores stream; its
highest height is La Juanita Hill (524 m). Sierra de la Ventana extends from the
Guaminí lagoon towards the Atlantic coast; its highest height is the Tres Picos hill
(1239 m). Sierras of Lihuel Calel are placed in the province of LPA.
In the Pampa plains prevails a temperate humid to subhumid climate with warm
summers. The rains are distributed throughout the year, without a rainy season
(600 mm at the SW and 1100 mm at the NE). By the West there is an extensive
system of freshwater or saline lakes and lagoons sporadically linked to each other.
The characteristic winds of the region are Pampero and Sudestada. Pampero wind
is cold and dry, comes mainly in summer from the Southwest bypassing the cordil-
lera de los Andes, and produces rain, temperature fall, and large clouds of dust.
Sudestada wind is cold and humid and comes from the Atlantic Ocean and Río de
la Plata producing floods on the right margin of Río de la Plata and in the south of
the littoral region. The Pampa is one of the most fertile areas in the world; the soils
are rich with a high content of organics and nutrients. The farming regions are par-
ticularly susceptible to flooding during the thunderstorms.
The Pampa plain is an important agro-industry region (agriculture and livestock
production). The agricultural capacity is remarkable; the agricultural economy is
based on the cultivation of wheat, corn, flax, oats, barley, rye, sunflower, potatoes,

© Springer Nature Switzerland AG 2019 63

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_4
64 4  The Pampa: An Infinite Plain

Fig. 4.1  A landscape from the Pampa plain. PG Alvarez personal collection

peanut, and soybean. More recently, vineyards established in the Buenos Aires
province. There is also horticultural production.
Much of the area is also used for cattle breeding for meat (Charolais, Hereford,
and Aberdeen Angus) and dairy (Holando-Argentino breed). There is also intense
pig (Duroc Jersey, Poland China, and Berkshire), equine, and sheep cattle and
avian production.
Fishing exploitation extends from the San Antonio cape to the South. Hake, from
the Atlantic Ocean, and shad, from the rivers, are the main fished species. As for
mining there are exploitations of granite and limestone basins. The industrial activ-
ity of the region is outstanding, ranging from petrochemical, automotive to textile.
Among the fauna, the great animals are today threatened (puma, wildcat, pampa
gray fox, pampas deer, skunk and ferret), there are other mammals (plain vizcacha,
southern mountain cavy, coypu, armadillos, weasels), birds (nandú or rhea, southern
screamer, partridges, elegant crested tinamou, birds of prey, burrowing owl), and
birds associated with aquatic environments (herons, Rallidae spp., white faced ibis,
white stork, neotropic cormorant, etc.). Some of the wetlands from the region are
important centers for migratory birds from the Northern hemisphere. The humid
Pampa ecosystem is one of the richest grazing areas in the world.
As for the flora, the original formations are temperate rangelands mainly of
Gramineae (genus Nassella, Piptochaetium, and Andropogon like Bromus, Stipa,
Poa, Paspalum, Eragrostis, etc.), halophile rangeland, dune grasslands, forest com-
munities restricted to ravines, and cords of shells (cordones de conchillas) (Fig. 4.2).
Today, the ceibo (Erythrina crista-galli) and the willow (Salix humboldtiana), part
of the primitive vegetation of the region, only grow in the riparian zones. Other spe-
cies, like the quebradillo (Acanthosyris spinescens) and the jarilla (Larrea spp.),
grow in the steppe regions. The Pampas are bounded by the drier Argentine Espinal
grasslands, which form a semicircle around the north, west, and south of the Humid
Pampas. In the Pampas from ERI, the grasslands associate arboreal elements such
as Prosopis alba, P. affinis, Celtis tala, and Phytolacca dioica (Cabrera 1976;
Oyarzábal et al. 2018).
4.2  Erythrina crista-galli L. var. crista-galli (Fabaceae) 65

Fig. 4.2  Plant communities from the Pampa eco-region. PG Alvarez personal collection

4.2  Erythrina crista-galli L. var. crista-galli (Fabaceae)

Synonymy  Coladendron crista-galli (L) Kuntze, Erythrina fasciculate Benth.,

Erythrina lauriflora Jacq., Erythrina pulcherrima Tod., Erythrina speciosa Tod.,
Micropteryx crista-galli (L.) Walp., Micropteryx lauriflora (Jacq.) Walp.,
Micropteryx fasciculata (Benth.) Walp.

Vernacular name  Seibo, ceibo, ceibo macho, seibo entrerriano, choipo, cockspur
coral tree (Eng.).

General information  The scientific name of the species describes the color and
aspect of its flower; the word erythros comes from the Greek and means red color;
crista-galli meaning is “the red crest of roosters.” In 1942, E. crista-galli flower was
named the national flower from Argentina (national decree 138474). It has also been
chosen as the national flower from Uruguay. The native name of this species is sui-­
nandi (Guaraní language) that means parrot’s food. E. crista-galli flowers are vis-
ited for its nectar by hummingbirds, Icterus pyrrhopterus (boyerito alférez), and
other birds. The beauty of the flowers promotes its use to decorate streets and parks
(Fig.  4.3). It is usual to see epiphytes such as ferns, cacti, orchids, and the
Bromeliaceous Tillandsia aeranthos growing on its branches (Chebez and Masariche
2010). E. crista-galli grows in the provinces of BUE, CHA, COS, ERI, FOR, JUJ,
LPA, MIS, SAL, SDE, SFE, and TUC. Other two Erythrina species that grow in
Argentina are E. falcata (ceibo de Jujuy) and E. dominguezii (ceibo rosado). E.
falcata has red flowers lighter than those from E. crista-galli and grows in the
woods of TUC, SAL, and JUJ and in the MIS forest. E. dominguezii has pink or
salmon-red flowers and grows in the woods from CHA and forests from SAL
(Haene 2007; Lozano and Zapater 2010; Cané 2016). It also grows in Bolivia,
Brazil, Chile, Paraguay, and Uruguay.
Besides its medicinal uses, the wood is used to make bows for percussion instru-
ments, horse saddles, and orthopedic devices (Abeddini et al. 2000), the pulp is used
66 4  The Pampa: An Infinite Plain

Fig. 4.3  Erythrina crista-galli L. var. crista-galli (Fabaceae). Branches with leaves and flowers
(a), detail of the flowers (b), fruits (c), flowers and fruits (d). MA Alvarez personal collection

to paper production for its color and lightness, and the extract of the flowers is used
as a fabric dye (Demaio et al. 2015). Finally, recent research (Enciso et al. 2017) has
demonstrated that a mixture of anthocyanins extracted from ceibo flowers can be
used as a sensitizer to produce low-cost dye-sensitized solar cells.
Botanical characteristics  E. crista-galli (cockspur coral tree) is a deciduous tree,
4–10 m high, with a trunk diameter by the base of around 1 m, with a gray and rough
bark. The trunk is short and reduced; the crown is wide and irregular. Branches are
arched, glabrous, and with stings. Leaves are deciduous, alternate, pinnate-­trifoliate,
3–11 cm in length, and 1.5–5.5 cm in width. The leaflets have entire margins. Stems
and petiole could have stings and thorns. Flowers are zygomorphic, hermaphrodite,
butterfly-shaped, fleshy with a red bell calix, and a corolla with five petals. One of the
petals is the longest, elliptic, and 3–4 cm in length (the standard), the lateral two are
smaller (the wings), and the other two are fused (the keel or carina) wrapping ten
stamens 3.5–5.0 cm in length. That architecture gives to the flowers an aspect from
which derives the name of the species, crista-galli. Flowers can be single or gathered
in foliose branches. Fruits are curved legumes, 10–20 cm in length and 1.0–1.5 in
width, dehiscent, and brown or black with lighter spots. This species has 1–6
­kidney-shaped seeds per legume, 10–19 mm in length. The trees gather and grow in
groups called ceibales in the banks of rivers and streams; they are highly tolerant to
waterlogging (Larré et  al. 2013). Seeds are, in those cases, transported by water.
Blooming is from late spring up to mid-summer; fruit ripening starts in August
(Cabrera and Zardini 1978; Lahitte et al. 1999; Díaz Cillio 2008; Sharry et al. 2011).

Ethnobotanical information  Leaves, flowers, and bark are used in folk medicine.
In the eighteenth century, the Jesuit monk Florian Paucke reported the use of the
ceibo bark decoction by the Mocovíes, native people from SFE, to heal puma bites
4.2  Erythrina crista-galli L. var. crista-galli (Fabaceae) 67

(Demaio et al. 2015). Other uses are in dermatosis (to heal wounds), asthma, hemor-
rhoids, and also as antiseptic and narcotic. The decoction of bark is prescribed to
treat urinary, vaginal, and respiratory tract infections. The decoction of leaves is
taken for their attributed hypnotic, sedative, anti-hemorrhoidal, and astringent activ-
ities (Martínez Crovetto 1964; Tourkarkissian 1980; Maier et al. 1999; Etcheverry
et al. 2003). An emollient syrup is made with the flowers for treating cough, colds,
catarrh, bronchitis, etc. (Ratera and Ratera 1980; Barboza et al. 2009; Sharry et al.
2011; Alonso and Desmarchelier 2015).

Chemical data  Erythrina isoquinoline alkaloids are distributed in all the plants
(Fig. 4.4). Erythraline was determined as the major alkaloid in leaves, while in fruits
was found erythraline and erythrinine. Fruit wall tissue appears to be the major site
of the spirocyclic alkaloid biosynthesis (Maier et al. 1999). Seeds produced eryso-
dine as main constituent. Also, they have erythramine, erysonine, erysopine, eryso-
vine, and hypaphorine (Deulofeu et al. 1947; Orekhov 1955).
The essential oil (EO) obtained by hydrodistillation of fresh flowers was mainly
made up of linalool (53.9%), spathulenol (9.6%), and α-terpineol (8.4%) (Fig. 4.5)
(Lamarque et al. 1998).
The leaves have the isoquinolines, crystadine and crystamidine (Fig. 4.6), and
oleanolic acid, erythrinine, erysodine, and erythrosine (Ito et al. 1973).

H3C
H
O
O
O O

N
O O
O N
N
H3C O O
HO

erythraline erythrinine erysodine

CH3
O
H3C
H3C N CH3
O
–
O H N
O
O

O O
N
O
H
NH

erythramine hypaphorine erysovine

Fig. 4.4  Some isoquinoline alkaloids from Erythrina crista-galli L. var. crista-galli
68 4  The Pampa: An Infinite Plain

Fig. 4.5  Some compounds found in the Erythrina crista-galli L. var. crista-galli essential oils

Fig. 4.6  Other chemical compounds found in leaves and flowers of Erythrina crista-galli L. var.
crista-galli

Flowers owe their red color to anthocyanin pigments, in particular pelargonidin

and cyanidin 3-glucosides and to the alkaloids erythristemine,
11𝛽-methoxyglucoerysodine, erysothiopine, 11𝛽-hydroxyerysodine-glucose, and
11-hydroxyerysotinone-rhamnoside (Fig.  4.6) (Merlugo et  al. 2015; Enciso et  al.
2017). Apigenin-7-O-rhamnosyl-6-C-glucoside was isolated as a new flavonoid
glycoside, and luteolin-6-C-glucoside was isolated for the first time from Erythrina
crista-galli (Ashmawy et al. 2013).
4.2  Erythrina crista-galli L. var. crista-galli (Fabaceae) 69

On the other hand, seeds have arginine, alanine, and aspartic acid and E. crista-­
galli lectins (ECA) that bind selectively to glycolipids with Galβ4GlcNAc and
Fucα2Galβ4GlcNAc terminal sequences (Teneberg et al. 1994).
Biological activities  The extracts from the dried aerial parts have been reported to
have antibacterial, analgesic, and anti-inflammatory activities. Also, dried leaf
extracts have antibacterial, psychotropic, and CNS depressor activity (Barboza et al.
2009; Alonso and Desmarchelier 2015).
The methanolic and ethanolic bark extracts have antibacterial activity against S.
aureus and B. subtilis and antifungal activity against Candida albicans and C. krus-
sel. On the other hand, extracts from the aerial part extracts were active against B.
subtilis (Mitscher et al. 1988; Vivot et al. 2006, 2007, 2012).
As for its activity on the central nervous system (CNS), the extracts and purified
fractions from leaves have a depressor effect in CNS and on the spontaneous motor
activity in rats (Etcheverry et al. 2003). A sedative effect could be attributed to the
isoquinoline alkaloids (Ito et al. 1973).
The aqueous leaf extracts had analgesic effect in mouse with a effect was at a
dose of 1000 mg kg−1 and an antinociceptive effect of 53%. It was also described a
curarizing effect of the alkaloids from seeds and bark (Deulofeu 1959).
The dichloromethane extract from leaves showed significant anti-inflammatory
effect ( 51% inhibition) in mouse 3 h. post administration; also, the test of edema in
mouse ears showed a 72% inhibition (Miño et al. 2002). The methanol crude extracts
from leaves protected from hemolysis induced by 2,2′-azobis (2-amidinopropane)
(AAPH) through inhibition of erythrocyte lipid peroxidation (Victol et al. 2015).
The in vitro antiparasitic activity of the ethanolic extracts from bark and leaves
of E. crista-galli L., E. dominguezii Hassl., and E. falcate Benth. was evaluated on
Trypanosoma cruzi trypomastigotes forms and Leishmania amazonensis, L. infan-
tum, and L. braziliensis promastigotes (Yaluff et  al. 2015). Against Leishmania
strains, only E. falcata Benth. leaf extracts showed activity; however, bark extracts
of the tree species have anti-leishmania activity. On the other hand, only E. falcate
Benth. leaf and bark extracts showed inhibitory activity against both strains of T.
cruzi. Another study reported antimalarial activity from three phenolic compounds
(phaseollidin, sandwicensin, and lonchocarpol A) from the stem bark of E. crista-­
galli (Tjahjandarie et al. 2014).
A matrix containing E. crista-galli lectin (ECA) supported the undifferentiated
growth of human embryonic stem cells (hESC) and human-induced pluripotent stem
cells (hiPSC). Those results suggested a potential application of ECA as a matrix for
culture and differentiation of human pluripotent stem cells (Mikkola et al. 2013).
The aqueous and aqueous methanol extracts and its fractions showed significant
estrogenic activities evaluated by Arabidopsis thaliana pER8: GUS reporter assay
and by their effect on the proliferation of the estrogen-responsive cell line MCF-7
(Ashmawy et al. 2013).
The aqueous extract of Erythrina showed cytotoxic activity against breast and
lung cancer cell lines (RathiSre et al. 2015). The superior cytotoxic activity of the
aqueous extract over the aqueous methanol extract is attributed to the presence of
70 4  The Pampa: An Infinite Plain

low amounts of erythrinan and benzylisoquinoline alkaloids in higher amounts

(Ashmawy et al. 2013).
Toxicity  Oral toxicity of the leaf infusion and the methanolic extract from the alka-
line infusion in rats was negative at a dose of 2000 mg kg−1 (Etcheberry et al. 2004).
Seeds are toxic probably as a result of its alkaloid content.

4.3  Phytolacca dioica L. (Phytolaccaceae)

Synonymy  Phytolacca arborea Moq., nom.nud, P. populifolia Salisb., Pircunia

dioica (L.) Moq., nom. illeg., Phytolacca dioica L. var. ovalifolia Chodat.

Vernacular name  Ombú, umbú, calamaic, morolawén, zumbí.

General information  Its massive enlarged base may grow to 4  m in diameter,

spreading above the ground, making the tree looking as standing on a dune. The
base and the trunk have water storage tissues accumulating up to 80% of water,
which could be an excellent fire-resistant adaptation to the intense grassland fires
that are common in this region. Since P. dioica derives from herbaceous ancestors,
its trunk has an unusual secondary growth rather than true wood. As a result of its
fast growth, its wood is soft and spongy. Since the sap is poisonous, it is not grazed
by cattle, and it is immune to locust and other pests. P. dioica grows in south of
Brazil, Paraguay, Uruguay, and Argentina. In Argentina it is found in the provinces
of BAI, CHA, COS, DFE, ERI, FOR, MIS, SAL, SFE, SLU, and TUC. Although it
is a species native from the provinces of FOR, COS, and MIS, it is characteristic
from the Pampa plain. It is an ideal tree for shade, but it has to consider its huge
diameter at its base (>3 m). It is used in parks and squares (Díaz Cillio 2008). The
flowers are visited by Diptera and Hymenoptera during the day and by Lepidoptera
in the night. In Spain the ombú is called Bella sombra and umbra tree in English-­
speaking countries.

Botanical characteristics  P. dioica is an 8–12 m in height tree with a huge crown,

a thick trunk, and large roots. As it is not ligneous, it cannot be considered as a true
tree but rather a huge herbaceous plant (Chebez and Masariche 2010). Leaves are
simple, fleshy, petiolate, alternate, caducous, glabrous, elliptic, or ovate, 5–13 mm
in length and 2–8 cm in width, with a yellow entire margin and a pinkish petiole.
Flowers are dioecious, apetalous, white or yellow, distributed in pendulous branches,
7–12 cm diameter. Male flowers have 20–30 stamens. Feminine flowers have 8–14
carpels, free in the upside during blooming. The fruit is a lobulated green-yellowish
berry with one black seed (Fig. 4.7). Blooming is from November to February; fruc-
tification is from November to May (Cabrera and Zardini 1978; Lahitte et al. 1999,
Alonso and Desmarchelier 2015).
4.3  Phytolacca dioica L. (Phytolaccaceae) 71

Fig. 4.7  Phytolacca dioica L., Phytolaccaceae. Tree (a), leaves and fruits (b), pendulous flower
clusters (c). MA Alvarez personal collection

Ethnobotanical information  The vernacular name ombú is a derivation from the

Guaraní name ïmboú that means “tree that attracts the rain.” The dried entire plant
is used as febrifuge, vulnerary, vermifuge, and antiseptic. The infusion made with
leaves and root bark is taken as emetic, antirheumatic, antiseptic, astringent, and
vulnerary. The dried leaf is considered as a purgative. Leaves are used in case of
cephalgia, as wound healer and coagulant. The bark ashes are used to produce soap
for their saponin content (Tourkarkissian 1980; Martínez Crovetto 1981; Quiroga
et al. 2001).

Chemical data  Ribosome-inactivating proteins (RIPs) are RNA N-β-glycosidases

that are classified into three types (1, 2, and 3) on the basis of the number and type
of their constituting polypeptide chains. P. dioica leaves and seeds contain at least
four type 1 RIPs, namely, PD-L1, PD-L2, PD-L3, and PD-L4. The expression of
RIPs in leaves has a seasonal and age regulation (Di Maro et al. 2012). Leaves
also have the flavonol ombuoside (Fig. 4.8); fresh flowers have fructose. The dried
seeds contain the RIP PD-S2 with a probable role as defense protein and/or as a
supplier of essential amino acids during seedling development (Parente et  al.
1993; Di Maro et al. 1999, 2012). Autumn leaves also have a secretory class III
peroxidase belonging to the plant heme-dependent peroxidase family (Guida
et al. 2011).
72 4  The Pampa: An Infinite Plain

Fig. 4.8  Ombuoside from

Phytolacca dioica L.
leaves

Biological activities  The extracts of the entire plant have molluscicidal, anti-­
inflammatory, antifungal, and antibacterial effects (Blanco et  al. 1998). Also, the
RIP D2 has demonstrated an in  vivo antiviral activity in two pathogen systems,
tobacco mosaic virus (TMV), Nicotiana tabacum, and tobacco necrosis virus
(TNV), Phaseolus vulgaris. It was presumed that the antiviral activity is expressed
only when D2 and the virus are in the same cell. Moreover, the oxidative burst in
D2-treated leaves could result from the acceleration of the hypersensitive response.
On the other hand, D2 cannot translocate into the phloem or induce systemic
acquired resistance (SAR) signals (Faoro et al. 2009).
The acid hydrolysate from P. dioica berries containing triterpenoid sapogenins,
such as phytolaccagenins (PG), has antifungal potency when tested against Candida
albicans ATCC 10231, Cryptococcus neoformans ATCC 32264, and clinical isolates
of C. albicans and C. neoformans (Di Liberto et al. 2010). PD-L4 is a type 1 RIP
isolated from summer leaves; a recombinant version (rPD-L4) was tested against E.
coli, P. aeruginosa, S. aureus, and the plant pathogen A. tumefaciens. Both the rPD-
L4 as its denaturized version demonstrated antimicrobial activity at a concentration
of 3 μM. The PD-L4 peptides obtained by chemical fragmentation were also tested.
A cryptic peptide, involving residues 40–65 of full-length protein (PDL440–465 pep-
tide), showed a significant antimicrobial activity on the different bacterial strains
tested. Since cationic antimicrobial peptides (CAMPS) are of interest, the authors
tested the activity of the PDL440–465 peptide towards HaCat and CaCo-2 human cells
lines. PDL440–465 triggered an anti-inflammatory response in lipopolysaccharide
4.4  Salix humboldtiana Willd. var. humboldtiana (Salicaceae) 73

(LPS)-treated CaCo-2 cells. By in silico analysis, homologous cryptic peptides in

other type 1 RIPs from Phytolaccaceae were identified with a possible key role
during seedling development, a process strongly susceptible to attack by pathogens
(Pizzo et al. 2015). Therefore, the Phytolacca RIPs have a potential use in crop
plant biotechnology to increase resistance to fungal and virus pathogens (Di Maro
et al. 2009).
Toxicity  When aqueous leaf and berry extracts (50, 100, 200 mg kg−1) were admin-
istered to Wistar rats for 14  days, a progressive reduction in the serum levels of
platelets, white blood cells, neutrophils, and monocytes was observed concomi-
tantly with a dose-dependent increase of lymphocytes and a reduction of eosino-
phils. As for enzymes, a decrease in serum activity of alkaline phosphatase, alanine
transaminase, and aspartate transaminase was reported. Those alterations were
accompanied by a significant reduction in body weight, absolute organ weight, and
organ-body weight ratio. All the abovementioned alterations could be an indication
of toxicity; hence, their use as an oral medicine would not be safe (Ashafa et al.
2010; Ashafa et al. 2011).

4.4  Salix humboldtiana Willd. var. humboldtiana (Salicaceae)

Synonymy  Salix chilensis Molina var. crispa, Salix magellanica Lam.

Vernacular name  Sauce colorado, sauce criollo, sauce chileno, sauce isleño,
sauce amargo, treique, cheique, reique, huayao.

General information  S. humboldtiana is the only Salix species native from South
America. It is a marsh tree widely distributed in Argentina that grows along rivers,
lakes, and lagoons in the provinces of BAI, CAT, CHA, CHU, COR, COS, DFE,
ERI, FOR, JUJ, LPA, MIS, NEU, RNE, SAL, SDE, SFE, SJU, and TUC. The wood
is light and soft. It is used for rural constructions to make boxes, packaging, toys,
oars, matches, poles, and fences (Ratera and Ratera 1980; Barboza et  al. 2009;
Sharry et al. 2011).

Botanical characteristics  It is a dioecious species, 15–25  m in height and

40–80 cm in diameter. Its foliage is deciduous; the crown is conic or rounded with
fallen foliage and pendulous branches (Fig. 4.9). The bark is grayish-brown, rough,
cracking in thick and irregular pieces. Leaves are simple, light green, alternate,
3–10 cm in length, and 0.6–0.8 cm in width; they are lanceolate, glabrescent, with a
toothed-serrate margin. Petiole is 0.5–1.0 cm in length; leaves and petiole are gla-
brescent. Inflorescences are pendulous terminal aments, 4–10  cm in length. The
masculine flowers are green-yellowish; the 5 mm in length calix has six unequal
stamens sustained by lanceolate-ovate bracts. The feminine flowers are green; the
calix is 2–3 mm in length, with a superior unilocular ovary, with multiple ovules.
74 4  The Pampa: An Infinite Plain

Fig. 4.9  Salix humboldtiana Willd. var. humboldtiana. Tree (a) and leaves (b). MA Alvarez per-
sonal collection

Fruits are small brown-greenish ovoid capsules, dehiscent by 2–4 valves, with many
seeds surrounded by copious white cottony hairs (Di Sapio and Gattuso 1992; Cané
2016; Agüero 2017).

Ethnobotanical information  The vernacular name huayao derives from wayaw in

Quechua language. The bark is used as a substitute of quinine. Also, it is referred
the use of the bark stem against influenza and dermatosis and as antirheumatic,
antispasmodic, sedative, febrifuge, astringent, and tonic (Tourkarkissian 1980; Di
Sapio and Gattuso 1992; Líate and Hurrell 1994; Barboza et al. 2009; Sharry et al.
2011). A preparation of boiled leaves and pieces of bark is administered to treat
diarrhea (Muñoz et al. 2000; Bourdy et al. 2004). The ashes from the bark are used
to treat dermic affections. As a shampoo, it is used as a hair revitalizing (Dimitri
et al. 1997; Lahitte 1997).

Chemical data  The leaves have flavonoids, coumarin, tannins, salicylic acid, and
the alkaloid salicine (Fig. 4.10) (Líate and Hurrell 1994).

Biological activities  The  antimicrobial activity of the aqueous extract of aerial

parts against Brucella canis and Staphylococcus aureus was reported (Toso et al.
2006). S. humboldtiana leaf extract showed stronger activity against Plasmodium
vinckei in vitro (99% inhibition at 10 mg ml−1) than in vivo (32% at 1000 mg kg−1)
(Muñoz et al. 2000). The bitter bark has febrifuge and tonic, sedative, and antispas-
modic activities. Salicine from bark has antirheumatic properties (Demaio  et  al.
2015, Medina et al. 2015). The well-known salicylate derivatives responsible for the
febrifuge and anti-rheumatism activity contained in the genus Salix justify the tra-
ditional uses of this species.
4.5  Grindelia pulchella Dunal var. discoidea (Hook. & Arn.) A. Bartoli & Tortosa… 75

Fig. 4.10 Chemical
structure of the alkaloid
salicine from Salix
humboldtiana Willd. var.
humboldtiana

4.5  G
 rindelia pulchella Dunal var. discoidea (Hook. & Arn.)
A. Bartoli & Tortosa (Asteraceae)

Synonymy  Grindelia anomala DC., Grindelia discoidea Hook. & Arn.

Vernacular name  Quilcha amarilla, botoncillo, pichana amarilla.

General information  G. pulchella produces a crude resin similar to that obtained

with G. camphorum in the Arizona desert (Tortosa et al. 2000); henceforth, the use
of Grindelia resins as alternatives to nonrenewable petroleum-based substances
was suggested (Timmermann and Hoffmann 1985). G. pulchella grows in the prov-
inces of BAI, CHU, COR, DFE, ERI, LRI, MEN, SDE, SLU, and TUC (Barboza
et al. 2009).

Botanical characteristics  G. pulchella is a shrub, 25–70 cm high, with lanceolate

or oblong alternate leaves 1.5–5.5 cm long, glabrous, toothed, with long petioles.
The apical capitula (heads) have dimorphic yellow flowers, the marginal flowers are
feminine, ligulate (sometimes absent), and the central flowers are tubular and her-
maphrodite. The receptacle is glabrous; fruits are thick smooth or slightly striated
glabrous achenes. G. pulchella has a characteristic secreting system composed of
glandular trichomes in the epidermis of stems, leaves, and involucral bracts (Cabrera
and Zardini 1978; Bartoli et al. 2011).

Ethnobotanical information  Most of the Grindelia species are used in folk medi-
cine as antispasmodic and diuretic. The aerial parts are used as febrifuge and anti-
rheumatic (Barboza et al. 2009).

Chemical data  The essential oils from the aerial parts contain the sesquiterpenes
aromadendrene, α-bisabolene, and γ-cadinene; the monoterpenes bornyl acetate,
camphene, and carvone; the diterpene cordobic acid; and the flavonol pachypodol
(Barboza et al. 2009). Also G. pulchella produces grindelic acid and resins composed
76 4  The Pampa: An Infinite Plain

of a mixture of labdane diterpenoids (Figs.  4.11 and 4.12) (Timmermann and

Hoffmann 1985).

Biological activities  Grindelic acid possesses antimicrobial activity towards sev-

eral phytopathogenic fungal and bacterial strains (Hernández et al. 2005). Grindelic
acid hydroxylated derivatives, the acids 6-β-hydroxygrindelic, 6-α- hydroxygrin-
delic, and 3-β-hydroxygrindelic, were successfully obtained by fungal bioconver-
sion from grindelic acid (Hernández et  al. 1997, 2002; Orden et  al. 2005) and
showed bioactive properties towards Tenebrio molitor larvae species and phyto-
pathogen fungi and bacteria. Also, the ethanolic and hydro-methanolic extracts
from aerial parts showed feed-deterrent effects (200 μg cm−2) on Epilachna paenu-
lata Germ (Coleopteran-Coccinellidae) larvae (IA 23.1 and 80%, respectively).
The aqueous, methanolic, and hexane extracts inhibited Avena sativa germination
and to a lesser extent Raphanus sativus germination. The aqueous extract
(2  mg  ml−1) has antibacterial activity (MIC  >  4) against E. coli ATCC 25922
(Palacios et al. 2007).

Fig. 4.11  Chemical compounds from Grindelia pulchella Dunal var. discoidea (Hook. & Arn.)
A. Bartoli & Tortosa essential oils
4.6  L arrea spp. (Zygophyllaceae) 77

Fig. 4.12  Grindelic acid O

and labdane from
Grindelia pulchella Dunal CH3
CH3 O
var. discoidea (Hook. & OH
CH3
Arn.) A. Bartoli & Tortosa

H3C CH
3

grindelic acid labdane

In vitro cultures  Calli cultures were established from leaves of G. pulchella seed-
lings in MS media (Murashige and Skoog 1962) with indolbutiric acid (IBA) and
benzylamine purine (BAP) at different concentration relationship (Hernández et al.
2005). Grindelane diterpenes were produced in all the callus lines established, while
grindelic acid was only produced in the lines growing in MS media with 20.0 μM
IBA and 18.0 μM BAP as plant growth regulators. Cell suspension cultures were
initiated from friable calli and maintained in the same conditions on rotary shaker.
However, they only produced grindelic acid that was released to the culture media.
When elicitation was performed with CuSO4 (1.0–2.0 mM) and DMSO (1 μl ml−1
culture), the terpenoid compound production increased at expense of cell growth
(Hernández et al. 2000, 2005).

4.6  Larrea spp. (Zygophyllaceae)

Larrea spp. are widespread distributed along the country (Pampa plains, Monte,
Chaco, Patagonia) being generally known as jarrillas (Morello 1958; Ezcurra et al.
1991). Larrea species inhabit arid ecosystems, with low temperature, or temperature
fluctuations, low absolute humidity, and high solar radiation (Alonso and
Desmarchelier 2015). Moreover, isolated populations of L. nitida and L. divaricata
occur in arid and semiarid areas of Chile, Bolivia, and Peru (Hunziker et al. 1972).
The infusions prepared with jarrillas have a strong bitter taste and a typical odor.
Botanical Characteristics
Larrea species rarely exceed 2–5 m height and can be differentiated by their leaf
characters. Larrea nitida has compound leaves, while L. divaricata and L. cuneifo-
lia have only two leaflets with a small filiform mucro. Flowers are solitary and axil-
lary, with five partially fused green sepals and five imbricated-free yellow petals.
Filaments are partially fused with a basal scale, forming a cup-shaped receptacle
where nectar is accumulated. The penta-carpellate, globose, and pubescent ovary is
superior, with a basal nectariferous tissue. The style and the stigma are simple in all
the species; the presence of an early female stage (protogyny) has been reported
(Simpson et al. 1977). The fruit is a schizocarp dehiscent into five one-seeded meri-
carps (Rossi et al. 1999).
78 4  The Pampa: An Infinite Plain

4.6.1  Larrea cuneifolia Cav.

Synonymy  Neoschroetera cuneifolia (Cav.) Briq., Schroeterella cuneifolia (Cav.)

Briq.

Vernacular name  Jarilla, jarilla macho, jarilla crespa, jarilla nortesur.

General information  It grows in the provinces of BAI, CAT, CHU, COR, LPA,
LRI, MEN, NEU, RNE, SAL, SDE, SJU, SLU, and TUC. Stems are used to roof
construction and also as fuel. Leaves are used as tinctorial (Ratera and Ratera 1980;
Ladio and Lozada 2009).

Ethnobotanical information  It is widely used as anti-inflammatory; for exter-

nally treating wounds; as diaphoretic, oxytocic, and emmenagogue; and for foot-
bath and rheumatism (Ratera and Ratera 1980; Barboza et  al. 2009; Ladio and
Lozada 2009).

Chemical data  The dried leaves have 11 flavonoid aglycones (Fig. 4.13): querce-
tin 3,7,3′,4′-tetramethyl ether (retusine), quercetin 3,7-dimethyl ether, quercetin
3,3′-dimethyl ether, quercetin 7, 3′-dimethyl ether (rhamnazin), quercetin 3′-methyl
ether (isorhamnetin), kaempferol 3,7-dimethyl ether (kumatakenin), kaempferol
3-methyl ether (isokaempferide), kaempferol ‘I-methyl ether (rhamnocitrin),
kaempferol, luteolin 7,3′-dimethyl ether (velutin), luteolin 3′-methyl ether (chrys-
oeriol), apigenin 7-methyl ether (genkwanin), apigenin, and dihydromyricetin
3′,5′-dimethyl ether (dihydrosyringetin) (Valesi et  al. 1972). Also, the flavonols
quercetin 3,7,3′-trimethyl ether and quercetin 7,3′,4′-trimethyl ether were deter-
mined (Sakakibara et al. 1976). Also, the leaves have the protein asparagine (Valesi
et al. 1972).

Biological activities  L. cuneifolia Cav. methanol and chloroform extracts have lar-
vicidal activity against Culex quinquefasciatus larvae (Batallán et  al. 2013). The
ethanol extract has antibacterial activity against Gram-positive (Amani et al. 1998;
Quiroga et al. 2001) and Gram-negative bacteria (Zampini et al. 2007). The aqueous
extracts have antioxidant properties (Carabajal et al. 2017). L cuneifolia Cav. infu-
sion (0.1 g ml−1), decoction (0.1 g ml−1), and tincture (0.1 g ml−1 alcohol 96°) were
tested against Fusarium subglutinans, F. thapsinum, F. bothii, and F. graminearum
sensu stricto. All the strains showed a greater susceptibility to tinctures, except in
the case of F. thapsinum that was mostly inhibited by the infusion. Decoctions were
more active than infusions against F. subglutinans, F. bothii, and F. graminearum
sensu stricto (Jiménez and Sgariglia 2010). Ethanolic extracts of leaves and bark
were tested against Salmonella enteritidis (Gram-negative), Enterococcus faecalis
(Gram-positive), and two strains of agricultural relevance, Xanthomonas axonopo-
dis pv. phaseoli (Gram-negative) and Clavibacter michiganensis (Gram-positive).
4.6  L arrea spp. (Zygophyllaceae) 79

CH3 CH3
O
O CH3
OH
O

O O O O
H3C H3C

CH3
O OH
OH O OH O

retusine rhamnazin
CH3
O
OH OH

HO O O O

OH O
OH O OH O

isorhamnetin kumatakenin

CH3
O
OH O CH3
OH
O
O O
H3C
HO O
OH
OH OH O

isokaempferide velutin
CH3
O
OH O
OH

HO O

O O
OH
OH O OH

chrysoeriol genkwanin

Fig. 4.13  Some of the flavonoids found in Larrea divaricata Cav. leaves

S. enteritidis was inhibited with all the extracts (MIC 106 to 220  μg  ml−1). Leaf
extracts were active against C. michiganensis, E. faecalis, and S. enteritidis. The
phytopathogenic bacteria were more resistant (MIC 216–228  μg  ml−1) (Lorenzo
et al. 2018). A patent (KR101348471B1) was published with a formulation based on
L. cuneifolia Cav. for preventing and treating estrogen-related menopausal symp-
toms (hot flashes, osteoporosis, venous thrombosis, and any of the symptoms of
atrophic vaginitis).
80 4  The Pampa: An Infinite Plain

4.6.2  Larrea divaricata Cav.

Synonymy  Schroeterella divaricata (Cav.) Briq., Neoschroetera divaricata (Cav.)

Briq., Larrea monticelli Perrone & Caro.

Common name  Jarilla, jarilla hembra, chamanilla, jarilla del cerro, yarilla.

General information  L. divaricata is widespread distributed in savannas along the

country, mostly in the provinces of BAI, CAT, CHU, COR, LPA, LRI, MEN, NEU,
RNE, SAL, SDE, SJU, SLU, and TUC. It is a nectar-producing xenophile bush that
attracts bees.

Botanical characteristics  L. divaricata is a bush, 2–3  m high, with woody and

flexible stems. Leaves are opposite, waxy, coriaceous, and bifoliate, with the knife-­
shaped leaflets united at the base. The small yellow solitary flowers are located in the
foliar axils. They have five pubescent sepals, five yellow petals, ten reddish stamens
distributed in two whorls, dithecous dehiscent anthers, and a five-carpel ovary. The
spheroidal fruits have five hairy mericarps that break at maturity into units contain-
ing each one a smooth reniform seed (Cabrera and Zardini 1978; Ratera and Ratera
1980; Alonso and Desmarchelier 2015; Cané 2016; Bernasconi Salazar et al. 2017).

Ethnobotanical information  L. divaricata Cav. is widely used in Argentine in

folk therapy as anti-inflammatory (Del Vitto et al. 1997) or antirheumatic (Ratera
and Ratera 1980; Pedernera et al. 2006). Aerial parts (leaves and tender branches)
are taken as a tea as anti-inflammatory; for treating external wounds; as diaphoretic,
febrifuge, emmenagogue, oxytocic, odontalgic, antitussive, antirheumatic, and
hypotensive; or for treating and preventing varicose veins. Also, it is used as rube-
facient, for footbaths, as antihyperlipidemic, against gastric disturbances, and peptic
ulcer. For its balsamic properties is taken to reduce mucus. The decoction with addi-
tion of salt is a good deodorant and foot antiperspirant (Mabry and Bohnstedt 1997;
Tourkarkissian 1980; Goleniowski et  al. 2006; Stege et  al. 2006; Barboza et  al.
2009; Ladio and Lozada 2009; Agüero 2017; Sen and Chakraborty 2017).

Chemical data  It was reported the presence of the nontriterpenoid larreagenine,

nordihydroguaiaretic acid (NDGA), α-thujene, myrcene, limonene, E-ocimenone,
thymyl acetate, γ-elemene, spathulenol, β-oplopenone, α-cadinol, eugenol, carva-
crol, limonene, and thymoquinone (Waller and Gisvold 1945; Boiteux et al. 2018).
Also, apigenine-7-methylether, 3,4′-dihydroxy-3′,4-dimethoxy-6,7′-cyclolignan
and aromadendrene oxide were identified (Vogt et al. 2013). The resin that covers
the leaves contains flavonoid aglycones, several lignans, and NDGA (Saigler et al.
1974; Barboza et al. 2006) (Fig. 4.14).

Biological activities  Extracts of L. divaricate display immunomodulatory

(Franchi-Micheli et al. 1986; Schreck et al. 1992; Anesini et al. 1996a), antimicro-
4.6  L arrea spp. (Zygophyllaceae) 81

CH3

O HO
CH3
O
CH3 CH3
HO

CH3 CH3
OH
HO
H3C CH3 OH

larreagenin nordihydroguaiaretic acid

OH HO

γ-elemene spathunelol α-cadinol

thymoquinone β-oplopenone

Fig. 4.14  Chemical compounds from Larrea divaricata Cav.

bial (Quiroga et  al. 2001; Stege et  al. 2006), antitumoral (Anesini et  al. 1996b,
1999), and anti-ulcerogenic activities (Gisvold and Thaker 1974; Anesini and Perez
1993; Amani et al. 1998; Quiroga et al. 2001, 2004; Pedernera et al. 2006; Davicino
et al. 2007).
The aqueous extract has tumorstatic capacity (Anesini et al. 1997); the methanol
and dichloromethane extracts have in vitro cytotoxicity (Bongiovanni et al. 2007).
The anti-inflammatory activity, attributed to NDGA, was displayed in cotton pel-
let granuloma and adjuvant induced arthritis models in rats. The methanolic extract
has anti-ulcerogenic and anti-inflammatory effect in ethanol- and HCl-induced
ulcers in rats. In the first case, the inhibition was dose dependent. Moreover, the cold
aqueous extract, infusion, and decoction had inhibitory activity at 0.04–0.1 mg l−1
against clarithromycin and metronidazole susceptible and resistant Helicobacter
pylori strains, which was attributed to flavonoid activity (Stege et al. 2006). Those
results validate the popular use of L. divaricata Cav. as an anti-inflammatory agent
with gastric anti-ulcerogenic properties (Franchi-Micheli et al. 1986; Schreck et al.
1992; Pedernera et al. 2006).
82 4  The Pampa: An Infinite Plain

The leaf aqueous extract was found to have both proliferative (at low concentra-
tions) and antiproliferative action on murine lymphoma cell line (BW 5147). The
proliferative effect was related to leukotriene modulation, while the antiproliferative
effect was related to the modulation of both hydrogen peroxide and nitric oxide. The
same extract induced proliferation in a murine lymphocytic leukemia cell line (El-4)
by the presence of nordihydroguaiaretic acid (NDGA) (Davicino et al. 2011a, b).
Hence, a fraction with low NDGA quantity was proposed as a potential therapy for
lymphoma and leukemia treatment (Martino et al. 2013).
The antimicrobial and antimitogenic activity of the aqueous extracts was
explained by interference on the arachidonic acid metabolism via lipoxygenase
(Anesini and Perez 1993; Anesini et al. 1999). The extracts in vitro inhibited the
transcription and/or replication of the viruses HIV-1, several herpes types, and Junín
(Garcia et al. 2003; Konigheim et al. 2004; Bernasconi Salazar et al. 2017). On the
other hand, immunization with L. divaricata crude extracts triggers the production
of antibodies that cross-react with proteins from Gram-negative bacteria including
Pseudomonas aeruginosa favoring its opsonophagocytosis by murine macrophages,
which was found promissory for developing a potential vaccine against P. aerugi-
nosa (Canale et al. 2018). An association of L. divaricata Cav. and Coffea arabica
extracts is commercialized for hair growth. It was reported that the spry formulation
improved the overall hair volume and appearance, increased hair thickness, induced
hair growth, and decreased hair loss without adverse local reactions in patients with
no cicatricial alopecia after a 3-month treatment. The amount of dandruff decreased,
but it was attributed to the inhibitory effect of C. arabica on Malassezia furfur
(Alonso and Anesini 2017).
The ethanolic and chloroformic extracts of aerial parts showed in vitro antifungal
activity against P. notatum, Lenzites elegans, Schizophyllum commune, Trichoderma
spp., Ganoderma applanatum, Pycnoporus sanguineus, Fusarium oxysporum, and
A. niger and the yeasts S. carlsbergensis and Rhodotorula spp. (Quiroga et al. 2001).
The chloroformic extract showed the highest antifungal activity, attributed to
apigenine-­ 7-methylether, NDGA, and 3,4′-dihydroxy-3′,4-dimethoxy-6,7′-
cyclolignan being this last compound the most active both in vitro (Fusarium gra-
minearum MIC 15.6 μg ml−1) as in vivo. The authors proposed the management of
wheat seedling blight and crown rot by using L. divaricata Cav. formulations (Vogt
et al. 2013). Finally, L. divaricata extracts have also shown activity against the phy-
topathogens Monilinia fructicola, the most destructive pre- and postharvest patho-
gen in stone fruit worldwide (Boiteux et al. 2018), Botrytis cinerea (Hapon et al.
2017), Fusarium graminearum, F. solani, F. verticillioides, and Macrophomina
phaseolina (Vogt et al. 2013).
In vitro culture  In vitro cultures of L. divaricata were established (Palacio et al.
2006). The addition of precursors (l-phenylalanine, cinnamic acid, ferulic acid, and
sinapic acid) affected the yields of the compounds of medicinal interest (NDGA,
p-coumaric acid, ferulic acid, and sinapyl alcohol). Feeding l-phenylalanine at 0.5,
1.0 and 3.0  mM concentration resulted in an increase of NDGA of up to
301.35 ± 1.19, 285.23 ± 28.44, and 190.53 ± 19.50 μg g−1 DW, respectively. At a
4.6  L arrea spp. (Zygophyllaceae) 83

concentration of 3  mM, l-phenylalanine increased p-coumaric acid concentration

from 47.43 ± 9.01 to 1157.28 ± 47.79 μg g−1 DW. The addition of 0.5 μM cinnamic
acid enhanced growth but not NDGA yields. The addition of cinnamic acid (1 and
1.5 μM), ferulic acid (0.1, 0.5, and 1 mM), and sinapic acid (0.1, 0.5, and 1 mM)
resulted toxic for the cells and the production of NDGA and the phenylpropanoids
were suppressed. The sinapyl alcohol content was not affected by any of the precur-
sors tested (Palacio et al. 2011). Calli, both with and without organogenesis, pro-
duced NDGA and quercetin, as did plantlet and wild plants. Cell suspension cultures
produced NDGA, p-coumaric acid, ferulic acid, and sinapyl alcohol. When undif-
ferentiated cultures were induced to produce adventitious shoots, the phenolic pro-
duction increased significantly. The capacity of undifferentiated tissues to produce
secondary metabolites is very limited, but when the undifferentiated cultures under-
went organogenesis, the yields usually increased (Alvarez 2014). However, plant-
lets regenerated from adventitious shoots of L. divaricate calli did not show the
same phenolic pattern as wild plants, with 3.6- and 5.9-fold lower levels of NDGA
and quercetin, respectively (Palacio et al. 2012).

4.6.3  Larrea nitida Cav.

Synonymy  Larrea balsamica (Molina) I.M.  Johnst., Covillea nitida (Cav.) Vail,
Mimosa balsamica Molina.

Common name  Jarilla crespa, jarilla de la sierra, jarilla fina.

General information  L. nitida mainly grows in the pre-Andes mountainous area

from the province of SAL to the North to the province of CHU to the South. It could
be found in the provinces of BAI, CAT, CHU, LPA, LRI, MEN, NEU, RNE, SAL,
and SJU (Kiesling 2003; Agüero et al. 2011).

Ethnobotanical information  The infusion of the dried leaf and stem is used as
emmenagogue. A not specified part is referred to as anti-inflammatory, choleretic,
antiperiodic, emmenagogue, sudorific, stimulant, balsamic, and vulnerary
(Tourkarkissian 1980; Ladio and Lozada 2009; Barboza et al. 2009).

Chemical data  In the aerial parts—2′4′-dihydroxychalcone, quercetin-3,3′,7-­

trimethyl ether, NDGA, 3′ methyl-nordihydroguaiaretic acid (MNDGA),
4-[4-(4-hydroxy-phenyl)-2,3-dimethyl-butyl]-benzene-1,2-diol, norisoguaiacine,
and ferulic acid (Fig. 4.15). Two epoxylignanes were isolated: compound 4 meso-
(7S,8S,7′R,8′R)-3,4,3′,4′-tetrahydroxy-7,7′-epoxylignan and compound 5
(7S,8S,7′S,8′S)-3,3′,4′-trihy droxy-4-methoxy-7,7′-epoxylignan. In the leaves:
asparagine. In the volatile compounds from aerial parts and propolis predominates
monoterpenoid compounds, mainly o-cymene and limonene (Agüero et al. 2011).
84 4  The Pampa: An Infinite Plain

Fig. 4.15  Chemical compounds from Larrea nitida Cav. aerial parts

Biological activities  Leaf and stem extracts showed antitumoral activity. Twigs
and leaf resin have antioxidant activity. Propolis and extracts from aerial parts of L.
nitida Cav. growing in the mountain valleys of Iglesia and Calingasta (SJU) have
antifungal activity attributed to MNDGA, NDGA, and (4-[4-(4-hydroxy-phenyl)-
2,3-dimethyl-butyl]-benzene-1,2-diol) (Agüero et al. 2011).

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Chapter 5
Mesopotamia-Paranaense Forest

5.1  Introduction: The Mesopotamia

The Northeast region of Argentina comprises two subregions: the Chaco region and
the Mesopotamia. Mesopotamia is located throughout the northeast of the country
and involves the provinces of Misiones (MIS), Corrientes (COS), and Entre Rios
(ERI). It is surrounded by two mighty rivers, Paraná and Uruguay, whose
Northeastern tributaries are the Iguazú, San Antonio, and Pepirí-Guazú Rivers. The
Iguazú Falls, declared “World Natural Heritage of Humanity,” is a system of 275
waterfalls in the middle of the Paranaense Forest, located 17 kilometers from the
mouth of the Iguazú River in the Paraná River, in the borderline between the prov-
ince of MIS and Brazil (Fig. 5.1). The total wide of the falls is 27 km, and the aver-
age height is 60–80 m, with 1800 m3 s−1 water flow; two-thirds of the falls are in
Argentina (https://iguazuargentina.com/es/parque-nacional-iguazu). The Paraná
and Uruguay Rivers flow downstream to the south up to the Rio de la Plata where
they empty, crossing the provinces Chaco (CHA), COS, ERI, and Santa Fe (SFE).
The Delta of Paraná marks the boundary between SFE and ERI and occupies the
southern coasts of these provinces. The Uruguay River is the border between
Argentina (the eastern border of the provinces MIS, COS, and ERI) and Uruguay.
Both subregions, Mesopotamia and Chaco, have rich and varied flora and fauna,
especially in the subtropical protected forest sectors of MIS and in the forests and
more temperate parks of COS and ERI, such as the Iberá Wetlands natural reserve.
In the provinces of MIS and COS, the soil is brick colored for the presence of a
mineral called laterite, composed of aluminum, silica, and large amounts of iron
oxide, which gives different shades of red to the soil.
It has a subtropical weather with high temperatures and intense rainfalls all along
the year, even in winter. The southern zone of the Mesopotamian subregion is hotter,
more humid and rainier than the northern, during summer. In this subregion the
climate is warm and humid to the north and temperate and dry to the west; the pre-
cipitations decreased from east to west.

© Springer Nature Switzerland AG 2019 91

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_5
92 5  Mesopotamia-Paranaense Forest

Fig. 5.1  The Iguazú Falls. Province of Misiones, Argentina. MA Alvarez personal collection

Agriculture and livestock are the most outstanding economy resources of the
region. Crops, corn, rice, beans, cassava, citrus, cotton, sugarcane, tobacco, and
especially tea and yerba mate are the most relevant. As for livestock, there are
breeding of cattle, horses, and pigs. The main industries are timber and steel and the
exploitation of iron deposits (MIS) and hydroelectric energy (COS: Iberá, Paraná
Medio, and Yaciretá-Apipé power stations).

5.2  The Paranaense Forest

This ecoregion, also known as ecoregion of the Misiones Forest, is part of the
Atlantic Forest, one of the most relevant biodiversity world hotspots that contain
about 7% of the world species and a high degree of endemism (Di Bitetti et  al.
2005). The climate is humid subtropical or tropical. There are plenty of rivers and
streams. In the Paranaense Forest is placed the Guaraní aquifer (up to 1000 m depth)
one of the world highest fresh water reservoirs.
As for the fauna, there are more than 500 bird species described for the region,
mammals such as Panthera onca (jaguar), Puma concolor (puma), Leopardus par-
dalis (ocelot), Tapirus terrestris (tapirus), Mazama spp., Nasua nasua (coati),
Speothos venaticus (vinegar fox), armadillo (Chlamyphoridae and Dasypodidae),
and various species of primates. Among the reptiles, there are Eunectes murinus
(green anaconda) and Caiman latirostris (overo yacare caiman). The region has
more than 50 amphibian species.
The Paranaense Forest is an extremely rich habitat with rich soils; the main veg-
etation type is semi-deciduous forests. Some of the plant species from the region are
Aspidosperma polyneuron, Ilex paraguariensis, Nectandra spp., Euterpe edulis,
Ocotea spp., and Astronium balansae (Oyarzabal et al. 2018). Within the area is the
Parque Nacional Iguazú that has 53 to 73 arboreal species per hectare, 85 species of
5.3 The Jesuit Missions 93

orchids, and more than 3000 species of vascular plants (1/3 from the total plant
vascular species from Argentina) (Placci and Giorgis 1994; Zuloaga et  al. 2000;
Giraudo et  al. 2003). Among the plant species used in popular medicine are
Aspidosperma quebracho-blanco Schltdl. (Apocynaceae), Anadenanthera
­colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul (Fabaceae) Aristolochia trian-
gularis Cham. Et Schlecht. (Aristolochiaceae), Blechnum occidentale L. var. occi-
dentale (Blechnaceae), Bulnesia sarmientoi Lor. Ex Griseb. (Zygophyllaceae),
Baccharis articulata (Lam.) Pers., Baccharis trimera (Less.) DC, Cecropia pachys-
tachya Trecul. (Cecropiaceae), Chenopodium ambrosioides L. (Chenopodiaceae),
Heteropterys angustifolia Griseb. (Malpighiaceae), Ilex paraguariensis A. St.-Hil
var. paraguariensis (Aquifoliaceae), Lepidium bonariense L. (Brassicaceae),
Maytenus ilicifolia Mart. ex Reissek (Celesteraceae), Passiflora caerulea L.
(Passifloraceae), Plantago tomentosa Lam. (Plantaginaceae), Phoradendron liga
(Gill) Eichl. (Viscaceae), Schinus molle L. (Anacardiaceae), Sida cordifolia L., Sida
rhombifolia L. (Malvaceae), and Tabebuia impetiginosa (Mart. ex DC.) Standl.
(Bignoniaceae).

5.3  The Jesuit Missions

The Jesuit Order arrived to the territory in 1608 and established settlements that
were inhabited by Guarani natives in the borders of present-day Paraguay, Brazil,
and Argentina. The missions acted as a state within a state in which the native
people, guided by the Jesuits, remained autonomous and isolated from Spanish
colonists and Spanish rule. In the missions the natives found protection from
enslavement (bandeirantes) and the forced labor of encomiendas. Under the leader-
ship of both the Jesuits and native caciques, the reductions achieved a high degree
of autonomy within the Spanish colonial empire. With the use of native labor, the
reductions became economically successful. In 1767, the Jesuits were expelled
from the Guaraní missions and the Americas by order of the Spanish King, Charles
III (Woodrow 1985; Anonimous 2017). In Argentina, the reductions settled in the
province of MIS were San Ignacio Mini, Nuestra Señora de Santa Ana, Nuestra
Señora de Loreto, Nuestra Señora de la Candelaria, Santa María la Mayor, Corpus
Christi, Santos Apóstoles Pedro y Pablo, San José, San Javier de Yaguaraities,
Nuestra Señora de la Concepción, and Santos Mártires, which were named World
Heritage Sites by UNESCO in 1983. In the province of COS were established the
Nuestra Señora de Yapeyú, Santo Tomé, La Cruz, and San Carlos Caapi reductions.
In the province of Córdoba, la Manzana Jesuítica (Jesuit Block) and Estancias de
Córdoba were named World Heritage Sites in 2000.
Nuestra Señora de Loreto was founded by fathers José Cataldino and Simón
Masseta to the left of the Paranapanema River. In 1613 they moved it to the margin
of the Yabebirí stream. It was the birthplace of the first printing press in South
America, founded by father José Serrano and father Neumann (MIS).
94 5  Mesopotamia-Paranaense Forest

Fig. 5.2  San Ignacio Miní Mission, Misiones, Argentina. MA Alvarez personal collection

San Ignacio Miní (Fig. 5.2) was also founded by father Cataldino and Masseta in
the Guirá in 1610. It moved for the first time to the right margin of Yabebirí stream
in 1631 and finally in 1660 to the place where its ruins are today (MIS).
Nuestra Señora de Santa Ana, founded by fathers Romero and C. de Mendoza in
1633 near Yacuiba stream, moved to the East of Paraná River in 1638 and finally in
1660 to the place where its ruins stand today (MIS).
Nuestra Señora de la Candelaria, funded in 1637  in Ibicuí by fathers Roque
González and Romero, was funded again in Caazapá mini (today Brazil). In 1667 it
was moved to the Paraná River coast to the Southwest of La Candelaria. It was con-
sidered the capital of the missions (MIS).
Santa María la Mayor, funded in 1628 by fathers Diego de Boroa and Claudio
Ruyer in the right bank of Iguazú River, in 1637 moved south to the site where its
ruins are today.
5.4  Cecropia pachystachya Trecul (Cecropiaceae) 95

The urban layout of the missions is attributed to father Antonio Sepp. The village
was divided into two equal parts. The church and the courtyard of the fathers were
in the center of the plaza. The streets ran parallel to the sides of the square. The main
street stretched from the main entrance to the plaza and the church. At the side of
the temple was the school or residence, on the other side the cemetery and the large
room (coty guazú). The other sides were occupied by the houses of the natives. A
house for the visitors was placed far from the houses of the natives and the cabildo
in front of the plaza. Without pre-established location were the jail, the school, and
a hospital. The layout was symmetrical, straight with north-south or east-west ori-
entation (Anonimous 2017).
In the schools, the curriculum was the Ratio Studiorum. In addition to Christian
doctrine, writing, and reading, the Jesuits considered important the teaching of sing-
ing and music (lyre, violin, harp) (Anonimous 2017).
From the commercial point of view, it behaved like a communal stay, with yerba
mate as the only safe sale product that gave them large profits since it replaced alco-
hol. Other crops were cotton, sugarcane, and wheat and groves of oranges, olives,
pears, vineyards, apple and peach orchards, and walnut trees. The Jesuits introduced
the silkworm to the region. They had 200,000 head of cattle in Yapeyú. The reduc-
tion manufactured glasses, tin vessels, mechanical watches, compasses, mirrors,
violins, bugles, harps, guitars, and astronomical lenses with rock crystals. They dis-
covered that the Itacurú stone had large amounts of iron, which allowed them to
produce farming tools and bells. They have also built dams and irrigation channels.
In January 1768 Carlos III from Spain banished the Jesuits from Paraguay. At the
beginning of the 19th century the reductions are already abandoned ruins (Woodrow
1985).

5.4  Cecropia pachystachya Trecul (Cecropiaceae)

Synonymy  Cecropia adenopus Mart. ex Miq., Cecropia adenopus Mart. ex Miq.

var. vulgaris, Cecropia lyratiloba Miq., Cecropia adenopus Mart. ex Miq. var. lyra-
tiloba, Cecropia adenopus Mart. ex Miq. var. macrophylla, Cecropia glauca Rojas
Acosta, Cecropia catarinensis Cuatrec., Cecropia cinerea Miq.

Vernacular name  Ambay, ambai, ambahi, ambaí, amba-í, ambahú, am-baiba,

ambay guazú, palo lija, palo de lija, umbaúba, embaúva (Amorin and Rosow 1988;
Alonso and Desmarchelier 2015).

General information  It is a medium-size tree that grows in groups or isolated in

temperate regions next to the riverbanks and into the Neotropical rainforest in the
provinces of CHA, COS, FOR, MIS, and SFE. Also, it could be found in the south
of Brazil and Paraguay (Consolini et al. 2006).
96 5  Mesopotamia-Paranaense Forest

Botanical characteristics  C. pachystachya is a dioecious tree, 10–20  m height,

with knotty hollow branches, and latex in the bark. The leaves are light wide palm-­
shaped; lamina is peltate, palmately compound, divided in long leaflets, dark green
on the upper side and silver gray and hairy on the lower side. Leaflets are entire,
obtuse, sometimes acuminated, with a petiole. The primary and secondary nerves
are brownish yellow, prominent on the underside. The petiole is 10–30 cm length,
cylindrical, 4–11 mm diameter, rigid, and brittle (ANP 1978). The flowers are small,
dioecious, distributed in digitated big fleshy receptacles (Fig. 5.3). The fruits are
little, reddish, edible achenes. The species that grew in the temperate region of the
country, the Espinal biogeographic region, is smaller (up to 1 m height).

Ethnobotanical information  C. pachystachya or ambay has been traditionally

used by Guaraní indigenous medicine (Gupta 1995). The oldest reference to ambay
is by Dominguez and Soto (1925) describing a syrup made with bark and leaves of
ambay that was prescribed as bequic and expectorant. Its use as expectorant, cardio-
tonic, to treat cough and catarrh is cited by Ratera and Ratera (1980) and Martínez
Crovetto (1981). Hypoglycemic, diuretic, and sedative activity was also reported
(Soraru and Bandoni 1978). There are some references to its use against eye infec-
tions. The trunk aqueous extract is used against cutaneous bores and as antidiarrheal
(Barboza et al. 2009; Alonso and Desmarchelier 2015).

Chemical data  The more relevant chemicals isolated from the aerial parts of the
plant are orientin, isoorientin, rutin, chlorogenic acid, and pomolic acid. In dried
leaves luteolin-C-glycosides, orientin, isoorientin, vitexin, isovitexin, catechin,
protocatechuic acid, isoquercitrin, β-sitosterine, stigmast-4-en-3-one, α- and

Fig. 5.3  Cecropia pachystachya Trecul (Cecropiaceae). Parque Nacional Iguazú, Misiones,
Argentina. MA Alvarez personal collection
5.4  Cecropia pachystachya Trecul (Cecropiaceae) 97

β-amyrin, β-sitosterol, cecropine, cecropinine, and arachidonic acid; the amino

acids alanine, glycine, serine, valine, isoleucine, asparagine, proline, aspartic acid,
phenylalanine, glutamic acid, and arginine; the sugars glucose, fructose, and galac-
tose; and the glucosides ambain and ambainin were found (Fig.  5.4). Also, the
plant contains oxidases, mucilages, and saponins. The bark has cecropine and tan-
nic acid. The chloroform/methanol (9:1) extraction from the bark yielded
fisetinidol-3-O-β-D-xylopyranoside (C20H22O9) or anadanthoside as the main com-
pound (Piacente et al. 1999, Costa et al. 2011).
In roots, there are β-sitosterol and tormentic acid. In roots and stems, there are
2α-acetoxy-3β,19α-dihydroxy-11α,12α-epoxyursan-28,13β-olide,3β-acetoxy-­
2α,19α-dihydroxy-11α,12α-epoxyursan-28,13β-olide and 2-O-acetyleuscaphic
acid.

Fig. 5.4  Some chemical compounds isolated from Cecropia pachystachya Trecul
98 5  Mesopotamia-Paranaense Forest

Biological activities  Effects of C. pachystachya aqueous extracts as anti-­

inflammatory, cardiotonic, sedative, and antioxidant have been associated with their
high level of antioxidant compounds (Velázquez et al. 2003; Consolini and Migliorisi
2005; Consolini et al. 2006; Schinella et al. 2008; Uchoa et al. 2010; Aragão et al.
2013). Chlorogenic acid, the main phenolic compound of the aqueous extract, plays
a major role in the protective effect to H2O2 challenge (Mendonça et al. 2016). The
leaf aqueous extract resulted beneficial against behavioral and biochemical modifi-
cations induced in mice by chronic unpredictable stress. Apparently, its phenolic
compounds could modulate glutathione peroxidase involved in the prevention of
stress-induced oxidative damage in mice hippocampus. Therefore, it was proposed
the use of ambay aqueous preparations for treating depressive disorders (Gazal et al.
2014).
The effect on cardiac rhythm differs between plants from temperate regions and
those from Neotropical regions; extracts from the first ones produced tachycardia,
while those from the second ones did not (Consolini and Migliorisi 2005). However,
both decreased rat spontaneous locomotion and exploratory behavior producing a
similar sedative effect. The central inhibition attributed to flavonoids (Consolini et al.
2006) could be the reason of ambay efficiency to treat cough and asthma. A stronger
hypotensive effect and an in vivo cardiotonic activity mediated by an anticholinergic
effect of extracts from Neotropical specimens were also informed. The analgesic and
central depressor effect was also described (Perez-Guerrero et al. 2001).
In a chronic renal failure model in rats, the aqueous extract attenuated hyper-
tension and renal lesions by decreasing angiotensin converting enzyme activity in
the renal cortex, which could also be mediated by flavonoids (Maquiaveli et al.
2014). The hexane extract from leaves showed anti-inflammatory activity
(Hikawczuk et al. 1998).
The methanolic extract of leaves was tested in normal and alloxan-induced dia-
betic rats (dose: 80 mg kg−1). Normal rats showed a better hypoglycemic activity on
the oral glucose tolerance test, while diabetic rats had a significantly higher hypo-
glycemic effect (< 116 mg ml−1 glucose after 12 h, 68% of blood glucose reduction).
That effect was partially attributed to chlorogenic acid and C-glycosylated flavones
(Ivorra et  al. 1989); however, the mechanism of action has not been proposed
(Aragão et al. 2010).
The methanolic extract of dried leaves inhibited quorum sensing when the bio-
sensors Chromobacterium violaceum ATCC 31532 and E. coli pSB403 were used
(Brango-Vanegas et al. 2014), but did not inhibit bacterial growth. Thus, the extract
and the isolated compounds could be used in the development of antipathogen drugs
or for antifouling coatings (Brango-Vanegas et al. 2014). The extracts did not have
a diuretic activity on rats (Consolini and Migliorisi 2005).
Finally, an anti-leishmanial activity (Mello Cruz et al. 2013) was attributed to the
flavonoids from the ethanol leaf extract (5% w/v) from Salvador, Bahia (Brazil). The
ethyl acetate-methanol soluble fraction diminished L. amazonensis (L.) promasti-
gote axenic growth/survival (70% inhibition in 96  h), inhibited arginase activity
(IC50 7 μg ml−1), and altered the mitochondrial kinetoplast DNA (K-DNA) array. As
Leishmania arginase participated in the maintenance of the infection in mammalian
5.5  Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul (Fabaceae) 99

hosts, its inhibition by C. pachystachya extracts was of paramount relevance.

Orientin (luteolin-8-C-glucoside), an arginase inhibitor (IC50 15.9 lM), was the
major component of the fraction; lower concentrations of quercetin, another arginase
inhibitor, were detected as well (Mello Cruz et al. 2013).
Toxicity  The infusions prepared with dried leaves (9.2% of crude extract) did not
show any genotoxic effect in animals. On the other hand, a neurotoxic effect was
found as the aqueous extract induced DNA damage on brain tissue independently of
the dose (500–2,000 mg kg−1) (Mendonça et al. 2016). The extract did not show
mutagenic activity in five S. typhimurium strains tested (TA100, TA98, TA97,
TA102, and TA1535).

Interactions  The extracts are synergic with digoxin (Domínguez and Soto 1925).

Legal status  C. pachystachya was included in the Argentinian National

Pharmacopeia sixth edition (1978). It is there established that “ambay is the dried
leaf from C. adenopus Martius and its varieties in different states of fragmentation
with no more than 2 % of foreign organic material.” In 2004 it was introduced in the
primary healthcare system of the province of Misiones.

5.5  Anadenanthera colubrina (Vell.) Brenan var. cebil

(Griseb.) Altschul (Fabaceae)

Synonymy  Acacia cebil Griseb., Anadenanthera macrocarpa (Benth.) Brenan,

Piptadenia cebil (Griseb.) Griseb., Piptadenia macrocarpa Benth., Piptadenia mac-
rocarpa Benth. var. cebil, Piptadenia hassleriana Chodat, Piptadenia hassleriana
Chodat var. fruticosa, Piptadenia macrocarpa Benth. f. puberula, Piptadenia mac-
rocarpa Benth. f. rupestris, Piptadenia macrocarpa Benth. f. microcarpa, Piptadenia
macrocarpa Benth. var. vestita, Piptochaetium macrocarpa Benth. var.
plurifoliata.

Vernacular name  Cebil, cebil colorado, cebil moro, curupaih, curupay, kurupá,
yopo, vilca, cohaba.

General information  A. colubrina is a caducous tree that grows on rocky hill-

sides in well-drained soil, often near rivers, in the provinces of CAT, CHA, COR,
COS, ERI, FOR, JUJ, MIS, SAL, SDE, and TUC. Its multiple uses include leather
tanning, as fuel (firewood), to make charcoal and bleach (natural soap), and for
rural construction. For its rapid growth, its condition as a nitrogen fixer, and its
resistance to strong wind, it has importance for the establishment of agroforestry
systems (Justiniano and Fredericksen 1998). A. colubrina flourishes between
September and November, and the fruit matures from August to September (Lorenzi
and Matos 2002).
100 5  Mesopotamia-Paranaense Forest

Botanical characteristics  A. colubrina is a tree 5–30  m tall, with an irregular

crown, and a 30–80 cm diameter trunk, covered by a rough, grayish, and thorny
bark. Leaves are bipinnately compound with opposite leaflets, 4–6 mm length. The
white flowers are gathered in panicles with globular spikes. The fruits are flattened,
rigid, glabrous, brilliant-brown dehiscent legumes containing 5–10 dark seeds
(Fig.  5.5). Seeds are brown, smooth, and laterally compressed (Viglione and
Vallejo 2011). Heartwood is pale brown and darkens to reddish brown with darker
colored veins on exposure. Sapwood is yellow, brown, or light pink (Lorenzi and
Matos 2002).

Ethnobotanical information  It was used by the native communities of Wichis,

Comechingones (Córdoba), Ranqueles, and Mbayas, among other native inhabit-
ants, as hallucinogen in medical rituals for its psychoactive principles (tryptamine-­
derived alkaloids) (Schultes and Hofmann 1993; Pérez Gollán and Gordillo 1993;
Alonso and Desmarchelier 2015).
The bark and seed powder was snuffed; semi-roasted seeds were used to make
the tribe kurupáih or kurupa (a spell according to Montoya) that produced ecstasies
when inhaled as snuff (rapé). Those Guaraní natives that owned the secret to pre-
pare kurupá were called kurupadyaras (Amorín 1974). The Mosetene ethnic group
from Bolivia used the bark to tan leather (Vargas & Quintana). Currently, it is used
in folk medicine to treat dysentery, as vermifuge and antipyretic. Bark preparations
are used as abortive, astringent, depurative, antitussive, and expectorant and for
healing infections (Toursarkissian 1980). The gum is prescribed for respiratory

Fig. 5.5  Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul (Fabaceae). Parque
Nacional Iguazú, Misiones, Argentina. MA Alvarez personal collection
5.5  Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul (Fabaceae) 101

problems. Seed are abortive and also used in case of heart complications and for
treating the disease called susto (Martínez and Pochettino 1999). A syrup formula-
tion is taken for treating cough, pertussis, and bronchitis. A bark macerate is used in
cases of inflammation and leukorrhea. The alcoholic preparation is applied on exter-
nal wounds, as hemostatic and cicatrizing (Matos 1997; Palmeira et al. 2010). The
bark releases a resin helpful in cases of skin infections (Mors et al. 2000). The fruit
is poisonous (Agra et al. 1996).
Chemical data  The main active ingredient is calcium bufotenate. The dried aerial
parts contain alnusenol, anadanthoflavone, lupenone, lupeol, betulinic acid, α- and
β-amyrin, β-sitosterol, stigmasterol, apigenin, 4-hydroxybenzoic acid, and cinnamic
acid. The heartwood contains dalbergin, 3,4,5-dimethoxy-3-dalbergione, and
5-methoxy-N-methyltryptamine. The dried bark has anadanthoside (®setinidol-­3-­
O-b-D-xylopyranoside), gallic acid, caffeic acid, chlorogenic acid, rutin, catechin,
quercetin, isoquercetin, luteolin, apigenin, and cinnamic acid. The gum has galac-
tose, arabinose, and acidic heteropolysaccharides (Fig.  5.6). The seed contains
bufotenin and N, N-dimethyltryptamine (Fig. 5.7) (Piacente et al. 1999; Barboza
et al. 2009; Leite et al. 2010).

Biological information  The ethanolic extracts (2024 μg ml−1) of A. colubrina are

a natural source of antibacterial compounds (flavonoids, tannins, and terpenes)
(Reginatto et al. 2001). The extract was effective against the broad-spectrum resis-
tant bacteria E. coli 27 (isolated from surgical wounds), E. coli ATCC10536,

Fig. 5.6  Chemical compounds from Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.)
Altschul
102 5  Mesopotamia-Paranaense Forest

Fig. 5.7  Indole alkaloids

from Anadenanthera N N

colubrina (Vell.) Brenan

var. cebil (Griseb.) Altschul HO

NH NH

bufotenin N, N-dimethyltryptamine

Staphylococcus aureus 358 (isolated from surgical wounds), and S. aureus

ATCC25923 (Figueredo et  al. 2013). Also, it was effective against Pseudomona
aeruginosa (MIC 0.5 mg ml−1) (Freitas Lima et al. 2014).
As antifungal it was effective against Candida albicans ATCC 76485 (MIC
0.31 mg ml−1) and C. albicans ambulatorial lineages (MIC 1.0 mg ml−1). Synergic
effect was observed with fluconazole but not with nystatin (Nunes et al. 2015).
The aqueous (10% w/v) and ethanolic (12.5% w/v DMSO) extracts have in vitro
anti-parasite action against larvae and nymphs from Rhipicephalus microplus
(Boophilus) (Lopez da Silva Filho 2007). R. microplus is considered the most
important tick parasite of livestock from the world. The aqueous extract was effec-
tive (85% mortality) against ticks at a concentration of 8.26  mg  ml−1 during an
exposure of 12 h. The ethanolic extract was effective (85% mortality) at 1.56, 6.25,
and 12.5 mg ml−1 concentrations. A. colubrina is also efficient in agronomic crop
pest control (Zanon de Moura et al. 2012). The leaf and bark aqueous extracts were
active against Meloidogyne incognita, one of the main cotton pathogen nematodes,
at 52.48 g l−1 and 70.72 g l−1 concentrations (Lima Fonseca et al. 2017). On the other
hand, the extract affected the viability of the larval stage and influenced the fecun-
dity of the lepidopteran Spodoptera frugiperda (Lepidoptera: Noctuidae) (Zanon de
Moura et al. 2012).
The leaf ethanolic extract (5, 10, 20 mg ml−1) showed allelopathic effect by inter-
fering with the germination of Brassica chinensis and Lactuca sativa (Goncalves
Silva et al. 2010).

5.6  Ilex paraguariensis A. St.-Hil var. paraguariensis

(Aquifoliaceae)

Synonymy  Ilex mate A. St.-Hil., nom. superfl., Ilex curitibensis Miers, Ilex domes-
tica Reissek var. glabra, Ilex sorbilis Reissek, Ilex theaezans Bonpl. ex Miers, Ilex
paraguariensis A. St.-Hil. f. latifolia, Ilex paraguariensis A. St.-Hil. f. parvifolia,
Ilex paraguariensis A. St.-Hil. var. dentata, Ilex paraguariensis A. St.-Hil. var. ido-
nea, Ilex curitibensis Miers var. gardneriana, Ilex paraguariensis A.  St.-Hil. f.
domestica, Ilex paraguariensis A. St.-Hil. f. sorbilis, Ilex paraguariensis A. St.-Hil.
5.6  Ilex paraguariensis A. St.-Hil var. paraguariensis (Aquifoliaceae) 103

f. confusa, Ilex paraguariensis A.  St.-Hil. f. dasyprionata, Ilex paraguariensis

A. St.-Hil. var. ulei, Ilex paraguariensis A. St.-Hil. f. glabra, Ilex paraguariensis
A. St.-Hil. var. usitata, Ilex paraguariensis A. St.-Hil. var. genuina.

Vernacular name  Yerba mate, mate, té de los Jesuitas, yerba del Paraguay, erva-­
mate, congonha, chá dos Jesuitas, thé du Paraguay, maté, ka’a, caá-mirim.

General information  In Argentina several species of Ilex, e.g., I. argentina Lillo

(roble  tucumano, palo de yerba, árbol de yerba), I. brevicuspis Reissek (caona,
cauna, siete sangrías), and I. dumosa var. Guaranina Loes (yerba señorita, caábera,
caáchiri, caáverá), grow. However, the most popular is I. paraguariensis A. St.-Hill
which is popularly taken as a beverage in Argentina and in neighboring countries
(Uruguay, South Brazil, and Paraguay). In Argentina, I. paraguariensis grows in the
provinces of COS and MIS where the plantations and processing facilities are
located. A study demonstrated that plants that grow in plantations have a higher
content of phenolic acids that those that grow in natural forests (Burris et al. 2012a).
Some adulterants are I. dumosa, I. theezans, I. brevicuspis, I. conocarpa, and I.
microdonta (Alonso and Desmarchelier 2015). Total RNA from I. paraguariensis
was sequenced; multiple sequence analysis allowed estimating the existence of
32,355 genes and 12,551 gene variants or isoforms. Members of more than 100
metabolic pathways were identified and categorized. Several transcription factors
and genes, involved in heat and oxidative stress, pathogen response, disease resis-
tance, and hormone response, among others, were identified. The authors also pro-
vided a collection of over 10,800 simple sequence repeats accessible to the scientific
community (Debat et al. 2014).

Botanical characteristics  I. paraguariensis is a perennial dioecious tree, 3–8 m

height. The bark is brown with white spots. The branches are glabrous, with peren-
nial alternate coriaceous green leaves, 4–10 length, a remarkable central nerve and
serrated margin. Petioles are up to 15 mm length (Fig. 5.8). Flowers are white, uni-
sexual, with four petals. The fruit is a drupe, purple reddish with 4–5 seeds.

Ethnobotanical information  The species was highly appreciated by the Guaraní

natives that called it caá mate. When the Spanish colonizers arrived, they adopted
the beverage. It was cultivated in the missions founded in the seventeenth century
by the company of Jesus. Today it is the most popular beverage in Argentina,
Uruguay, Paraguay, and south of Brazil. The leaves and small pieces of petiole are
used to prepare an infusion, the mate, which is consumed mostly in a recipient, also
called mate, using a metallic straw, the bombilla. It could also be taken cold (tereré)
or as a tea in a cup (mate cocido). The popular use attributes to the infusion proper-
ties against influenza and scurf, as tonic, refrigerant, diuretic, digestive, stimulant,
choleretic, eupeptic, hypolypemic, and hepatoprotective. The infusion made with
the aerial parts is used as galactagogue, cicatrizing, and in cases of cephalea or fever
(Toursarkissian 1980; Gupta 1995; Cansian et al. 2008).
104 5  Mesopotamia-Paranaense Forest

Fig. 5.8  Ilex paraguariensis A.  St.-Hil var. paraguariensis (Aquifoliaceae). San Ignacio Miní
­mission, Misiones, Argentina (a and b), and in Buenos Aires Botanical Garden (c). MA Alvarez
personal collection

Chemical data  The leaves contain saponins, alkaloids, and phenolics (Fig. 5.9).
Among the phenolic compounds are caffeic acid, chlorogenic acid,
3,4-­dicaffeoylquinic acid, 3,5-dicaffeoylquinic acid, 4,5-dicaffeoylquinic acid, and
5-caffeoylquinic acid. The flavonoids are quercetin, rutin, and kaempferol. The
purine alkaloids are caffeine (1,3,7-trimethylxanthine), theobromine
(3,7-­dimethylxanthine), and theophylline (1,3-dimethylxanthine). The saponins
identified were matesaponin, matesaponin 2 (ursolic acid 3-O-[β,D-­
glucopyranosyl(1  →  3)[α,L-rhamnopyranosyl(1  →  2)]]α,L-arabinopyranosyl]
(28  →  1)-β,D-glucopyranosyl ester), matesaponin 3 (ursolic acid 3O[β,D-­
glucopyranosyl(1 → 3)α,L-arabinopyranosyl](28 → 1)[β,D-glucopyranosyl(1 → 6)
β,D-glucopyranosyl] ester), matesaponin 4 (ursolic acid 3,O-[β,D-glucopyranosyl
(1  →  3)[α,L-rhamnopyranosyl (1  →  2)]]α,L-arabinopyranosyl] (28  →  1)[β,
D-glucopyranosyl(1  →  6) β,D-glucopyranosyl] ester), matesaponin 5, guaicin B,
and nudicaucin (Martínez et  al. 1997; Saldaña et  al. 1999; Saldaña et  al. 2002;
Taketa et al. 2004; Barboza et al. 2009; Burris et al. 2012a). Also, several vitamins
such as ascorbic acid, nicotinic acid, and beta carotenes, and minerals as iron, mag-
nesium, manganese, calcium, copper, potassium, zinc, and sodium were found
(Moreau 1973; Alikardis 1987; Clifford and Ramirez-Martinez 1990; Ashihara
1993; Gosmann et al. 1995; Kraemer et al. 1996; Ricco et al. 1991; Schenkel et al.
1996; Vera García et al. 1997; Paroul et al. 2002; Filip and Ferraro 2003; Filip et al.
2001; Cansian et al. 2008).
5.6  Ilex paraguariensis A. St.-Hil var. paraguariensis (Aquifoliaceae) 105

HO OH

O
HO
O OH
O O
HO O
OH
HO
O O

O O

HO OH
HO OH

OH OH OH

3,4-dicaffeoylquinic acid 3,5-dicaffeoylquinic acid

O
O O

N N
N N N N H

O N N N N
O N O N

caffeine theobromine theophylline

HO

HO O
OH
O

HO O
HO O

OH
OH OH

OH OH O

chlorogenic acid kaempferol

Fig. 5.9  Some chemical compounds found in Ilex paraguariensis A. St.-Hil var. paraguariensis.
(Filip et al. 1998)

Biological activities  The depurative, stimulant, and diuretic effect of yerba mate
was attributed to its high purine content (Baltassat et al. 1984) and the antioxidant
activity to its polyphenol content (Filip et al. 2000). Caffeine was reported as the
responsible compound for the stimulant properties.
It was found that yerba mate produced a dose-dependent increase in alert or
attentive wakefulness, a decrease in non-REM sleep, and shallow sleep episodes
in adult cats (Felis domesticus) that orally received 0, 2.5, 5.0, 10.0, or 30.0%
106 5  Mesopotamia-Paranaense Forest

I. paraguariensis solution over a period of 5 minutes. The authors proposed the

use of different varieties or products for treating excessive sleepiness and for peo-
ple that need to be awake for long periods (Falconi et al. 2013).
Antioxidant activity was tested by the inhibition of the luminal-induced chemi-
luminescence assay (Actis-Goretta et al. 2002), the inhibition of 2,2′-thiobarbituric-­
reactive substances production in liposomes (Filip et al. 2000), the peroxidase-like
activity (Anesini et  al. 2006), the ferric thiocyanate method (Bastos et  al. 2006;
Turner et al. 2011), and with 6-hydroxy-2,5,7,8- tetramethychroman-2-carboxylic
acid. The extract obtained by spray drying also showed a strong free radical scaven-
ger activity (Berté et al. 2011).
Leaf extracts were found to increase bile flow, have anticancer effect, trigger
inhibition of LDL oxidation, and have a potential therapeutic and/or prophylactic
use in cardiovascular disorders (Bracesco et al. 2003; Ruxton et al. 2007).
Assays performed with supercritical carbon dioxide extracts of I. paraguariensis
on adult male Wistar rats with acute induced-peritonitis inhibited the neutrophil
recruitment in the circulating blood (Pasquali et al. 2011). It also reduced acute lung
inflammation in mice exposed to cigarette smoke (Lanzetti et al. 2008).
The treatment of rats with 1 g kg−1 body weight of instant yerba mate during
8 weeks attenuated gain weight and adiposity, decreased epididymal fat pad weight,
and restored cholesterol, triglycerides, LDL cholesterol, and glucose serum levels
(Arçari et al. 2009; Morgan-Martins et al. 2013). Besides, an inhibition of athero-
sclerosis progression in cholesterol-fed rabbits was informed (Mosimann et  al.
2006). The methylxanthine fraction increased lipogenesis of the muscle and hepatic
glycogen synthesis and lipolysis in adipose tissue, while the saponin fraction
enhanced lipogenesis in adipose tissue and also fecal fat excretion in Wistar rats (De
Resende et al. 2015). I. paraguariensis showed hypolipemic effect in healthy indi-
viduals with normo- or dyslipidemia, reduced LDL cholesterol, non-HDL choles-
terol, and apolipoprotein B blood levels. Also, it reduced the LDL-C/HDL-C ratio
and increased HDL-C (Morais et al.). Besides, it lowered LDL-C in hypercholester-
olemic individuals (De Morais et al. 2009). The I. paraguariensis aqueous extract
modulated the expression of genes related to obesity such as those that codify for
TNF-α, IL-6, leptin, CCR2, CCL2, angiotensinogen, PAI-1, adiponectin, PPAR-γ2,
PGC-1α, and UCP1 (Arçari et  al. 2009). Also, it regulated adipogenesis in a
β-catenin-dependent manner (Arcari et  al. 2013a). Finally, it was reported an
improvement of glucose tolerance in obese animals and a modulatory effect on
insulin-related target genes in the liver of animals with a high-fat diet (Arcari et al.
2011, 2013b). Caffeine combined with the activities of the polyphenols and flavo-
noids was proposed of the responsible of the thermogenic activity observed (Ruxton
et al. 2007).
Antimicrobial activity of the aqueous extracts from commercial yerba mate
against pathogenic E. coli O157:H7 was reached at relatively low concentrations (5
to 10 mg ml−1) supporting the use of yerba mate as a bio-based antimicrobial for
foods and beverages. On the contrary, neither the crude ethanolic extract nor the
saponin fraction, both at a 100 mg ml−1 concentration, showed antimicrobial activ-
ity (Burris et al. 2012b).
5.6  Ilex paraguariensis A. St.-Hil var. paraguariensis (Aquifoliaceae) 107

The anticancer activity of the infusions showed an inhibition of viability and

cell growth of human colon adenocarcinoma cells SW480 and their metastatic
derived SW620 cells (Pérez et al. 2014). The saponin inhibited colon cancer cells
(HT-29) proliferation activating a specific intracellular apoptosis pathway and
increased the expression of the pro-apoptotic protein Bax, decreased the expression
of anti-­ apoptotic protein Bcl-2, and subsequently activated caspase- 3
(Puangpraphant et al. 2011). Also, an inhibitor effect on CaCo-2 adenocarcinoma
cells growth was reported (González de Mejía et al. 2010).
The higher bioavailability of compounds in industrially processed samples (har-
vesting, zapecado, drying, and aging) than in green leaves was attributed to the
conditions of temperature, humidity, cell disruption, and mechanical impact during
the industrial process (Turner et al. 2011).
Beyond its traditional use, yerba mate has the potential to be used in the cosmetic
and food industries. The increasing interest for yerba mate and its sub-products in
the United States, Asia, and Europa, where it is exported, led to the development of
a spray-drying technology that allow their conservation for long periods. The recon-
structed extract had an IC50 of 2.52 mg ml−1, high catalase-like activity, and a high
concentration of caffeic acid (1.54 mg g−1), 5-caffeoylquinic acid (91.40 mg g-1),
rutin (5.38 mg g−1), and total phenolics (178.32 mg g−1) showing that the technology
could be useful since it maintained the antioxidant activity and polyphenolic con-
tent (Berté et al. 2011).
Recently, the use of yerba mate as negative oral contrast in magnetic resonance
cholangiopancreatography was explored using phantom devices (100 g l−1 infusion
and 3:4, 1:2, and 1:4 dilutions). It was found that it produced an innocuous contrast
for the patient, was effective for the elimination of the signal of the gastrointestinal
tract, and improved the visualization of the anatomical structures of the biliary tract
(Martin et al. 2006).
Toxicity  The reports about toxicity of yerba mate are contradictory. Some studies
reported an increased risk of bladder cancer and renal cell carcinoma associated
with mate drinking in humans (Vassallo et al. 1985; De Stefani et al. 1998, 2007)
and mutagenic and clastogenic activities in cell culture (Fonseca et al. 2000). The
aqueous extract was genotoxic for E. coli, mutagenic for Salmonella typhimurium,
and in vitro cyto- and genotoxic for lymphocytes (Wnuk et al. 2009). On the other
hand, other reports established that I. paraguariensis is not genotoxic to liver, kid-
ney, and bladder cells. Animals that received different doses of the aqueous extract
showed unaltered levels of DNA damage. Moreover, animals that received regular
ingestion of mate increased DNA resistance to H2O2-induced DNA strand breaks
and improved DNA repair (Miranda et al. 2008). Also, it was reported that neither
the infusions nor concoctions of I. paraguariensis showed cytotoxicity in the
Artemia salina test (Bidau et al. 2004).
It was referred that at high doses, yerba mate causes insomnia, nervous excita-
tion, gastritis, nausea, and colitis.
108 5  Mesopotamia-Paranaense Forest

In vitro culture  In vitro cultures (Ross et  al. 2017) and micropropagation of I.
paraguariensis were achieved in 1/4 MS culture medium with the addition of 3%
sucrose and BAP (0.44  μM). Different chemical and physical culture conditions
affected the anatomy, histology (architecture, leaf vein pattern, leaf thickness, sto-
matal index, etc.), and physiological behavior of plants, being the temporary immer-
sion system the most conservative treatment (Luna et al. 2017).

Legal status  Ilex paraguariensis was included in the 6VIth edition of the FNA and
in the Argentina’s Alimentary Codex. I. paraguariensis also figures in the British
Herbal Pharmacopoeia (1983–1996), Martindale (30th ed.), German Monographies
commission E, List II from the European Council, and Ayurveda Pharmacopeia and
is approved as dietary supplement by the FDA. In the United States, yerba mate is
listed as GRAS (generally recognized as safe) (Newall et al. 1996).

5.7  Tabebuia impetiginosa (Mart. ex DC.) Standl.

(Bignoniaceae)

Synonymy  T. avellanedae Lorentz ex Griseb., Tabebuia ipe (Mart.), Handroanthus

avellanedae (Lorente ex Griseb) Kuntze, H. impetiginosus (Mart. Ex DC.) Mattos,
H. heptaphylla (Vell.).

Vernacular name  Lapacho, lapacho rosado, lapacho dorado, palo de arco, taheebo.

General information  T. impetiginosa grows in America from Mexico to the North

and center of Argentina. Particularly, in Argentina it is found in the provinces of
CAT, JUJ, SAL, and TUC. The wood is used as timber and to obtain a yellow dye
(Bandoni et al. 1972; Tyler 1987). The native and exotic species of the genus are in
general very required as ornamental, due to the striking brightness of its flowers. It
is highly appreciated as timber for its density (higher than water), insect resistance,
and durability (Mechler 2009). It is also called the “new mahogany,” being the sec-
ond most expensive timber and the most logged species in Brazil (Bonfim Silva-­
Junior et  al. 2017). Hence, the species merit greater protection than provided by
current generic forestry laws in Brazil and other Amazonian countries (Schulze
et al. 2008). A well-curated but unfinished genome assembly was made providing a
useful tool for genetic and functional analysis (Bonfim Silva-Junior et al. 2017).

Botanical characteristics  T. impetiginosa is a caduceus tree, 5 to 50 m high, with

a diameter of up to 1  m. The crown is stratified; leaves are opposite, petiolate,
10–14  cm length, 5–7 foliolate, with reticulate venation; the lower leaflet is the
smallest, the terminal the largest. Leaflets are narrow, lanceolate, or elliptical, with
an entire or slightly crenate margin. The gray brownish or yellowish external bark is
suberified. The internal bark is pink whitish, 0.5–1.0  cm width. The flowers are
short panicles, 15 cm length; calix is greenish, campanulate to tubular, 2.0–2.5 cm
5.7  Tabebuia impetiginosa (Mart. ex DC.) Standl. (Bignoniaceae) 109

length, in general 5-lobed, looking like a trumpet. The corolla is pink or red, tubular
by the base, and expanded in the upper section, 7–10 cm length; the outside surface
is either glabrous or pubescent. Blooming take place in spring (August–September).
The fruits are dark-brown dehiscent capsules, 15–38  cm long, 0.9–1.5  cm wide,
smooth, with two lateral sutures, covered by numerous scales, with a persistent
calix. The fruits contain numerous winged white seeds, 2–3 cm length. Plants are
hermaphrodite (Gilman and Watson 1994; Grose and Olmstead 2007; Mechler
2009; Alonso and Desmarchelier 2015).

Ethnobotanical information  In the pre-Columbian America, it was used by dif-

ferent native communities to treat diseases such as arthrosis, fever, and digestive
and circulatory diseases and as a magical component for treating mal de ojo, etc.
Later on, infusions from the aerial parts were used as antibacterial and the decoc-
tions of the stem bark as astringent, vulnerary, and antiseptic. Also, in popular medi-
cine it is taken to treat external diseases as ulcers, eczema, psoriasis, hemorrhoids,
candidiasis, and skin tumors. Besides, it is used as antianemic, hypotensive, revital-
izing, hypoglycemic, abortive, and anti-infective urinary laxative in diseases of
respiratory tract (whooping cough) and as an antitumor (leukemia). Finally, it has
applications as antinociceptive, anti-dermatogen, antiviral, antitrypanosome, anti-
fungal, antibacterial, and antitumor (Ratera and Ratera 1980; Toursarkissian 1980;
Barboza et al. 2009; Alonso and Desmarchelier 2015).

Chemical data  The main active principle from the species of this family is lapa-
chol, a quinone compound (Fig. 5.10). Other compounds found in T. impetiginosa
are flavonoids, tannins, and other quinones. The dried heartwood contains, besides
lapachol, the quinoids 1-hydroxyanthraquinone and 2-methyl anthraquinone. The
dried bark contains the iridoid monoterpene ajugol, 6–0-(4-hydroxybenzoyl). Also,
it contains anisaldehyde and the quinoid naphtho (2,3b) furan-4,9-dione, 2-acetyl
(Fig. 5.11). The stem bark contains lapachol, α-and β-lapachone, the lignans cyclo-
olivile, and aryltetraline-type lignans, and furanonaphtoquinones. The mayor vola-
tile compounds isolated from dried inner bark are 4-methoxybenzaldehyde,
4-methoxyphenol, 5-allyl-1,2,3-trimethoxybenzene (elemicin), 1-methoxy-4-(1E)-

O O O
O

O
O
OH O

lapachol α-lapachone β-lapachone

Fig. 5.10  Chemical structure of lapachol and other quinones found in Tabebuia impetiginosa
(Mart. ex DC.) Standl
110 5  Mesopotamia-Paranaense Forest

OH

O
O
OH HO
O
O O
HO
HO

HO OH

OH

ajugol 2-methoxy-4-methylphenol eugenol

Fig. 5.11  Chemicals found on the dried inner bark of Tabebuia impetiginosa (Mart. ex DC.)
Standl

1-propenylbenzene (trans-anethole), and 4-methoxybenzyl alcohol, 2-methoxy-­4-

methylphenol and eugenol (Park et al. 2003; Barboza et al. 2009).

Biological activities  Dried plant material (inner bark) is commercialized to pre-

pare infusions, pills, and syrups. Impetiginosa means that it was used to treat impe-
tigo, an infection of the skin caused by Staphylococcus or Streptococcus. Also its
activity was referred as antiulcer, leishmanicidal, anti-carcinogenic, anti-edema,
anti-inflammatory, antimalarial, antiseptic, antiviral, bactericide, fungicide, pesti-
cide schistosomicide, antinociceptive, and hypocholesterolemic in hyperlipidemic
mice (Barboza et al. 2009). Some studies have shown that the preparations made
with bark and leaf and the dried bark have antimicrobial activity, while leaves have
antioxidant activity. In vitro antimicrobial activity assays were made testing ethano-
lic extracts from leaves, stem bark, and branches against Staphylococcus aureus
ATCC 25923, Salmonella enterica ATCC 1307, Acinetobacter baumannii ATCC
17978, Enterococcus faecalis ATCC 29212, Pseudomonas aeruginosa ATCC
27853, and Escherichia coli ATCC 25922. The crude ethanolic extract of leaves has
moderate antimicrobial activity against S. aureus (MIC ≥312.5 μg ml−1) and a very
weak inhibitory activity against S. epidermidis (≥ 625 μg ml-1). For Acinetobacter
calcoaceticus and Enterococcus faecalis, MIC was ≥1 μg ml-1 (Moreira Vasconcelos
et al. 2014). In vitro studies with the ethanolic and methanolic extracts and the infu-
sion were made to test their free radical scavenging activity (DPPH), reducing
power and inhibition of lipid peroxidation in brain homogenates. In all cases the free
radical scavenging ability was negative (Suo et al. 2013; Vasconcelos et al. 2014).
Lapachol, the main active principle of the plant, was associated with activities
such as antiulcer, leishmanicidal, anticarcinogenic, anti-edema, anti-inflammatory,
antimalarial, antiseptic, antiviral, bactericide, fungicide, and pesticide schistosomi-
cide (Hussain et al. 2007; Rodrigues de Almeida 2009; Silva 2009).
The methanolic extract from commercial dried inner bark was successively par-
titioned into hexane, chloroform, ethyl acetate, butanol, and water-soluble fractions,
and then they were tested against Helicobacter pylori ATCC 43504. The hexane,
5.8  Allophylus edulis (A. St.-Hil., A. Juss. & Cambess.) Hieron. ex Niederl.… 111

ethyl acetate, and chloroform fractions had a strong antibacterial activity. The com-
ponents present in the chloroform fraction 2-(hydroxymethyl) anthraquinone,
anthraquinone-2-carboxylic acid, and lapachol were separately studied being
2-hydroxymethyl anthraquinone at a dose of 0.1 mg per disc, the one with the stron-
gest activity against H. pylori (Park et al. 2006).
The antioxidant activity of the volatile compounds isolated from the dried inner
bark showed a dose-dependent inhibitory activity at a concentration of 1 mg ml−1.
Among the components isolated from the extract, 2-methoxy-4-methylphenol and
eugenol had the strongest antioxidant activity. The extract appeared to be more
active in the hexanal/hexanoic acid assay than in the conjugated diene assay (Park
et al. 2003).
Toxicity  The ethanol extract of leaves and stem bark did not show cytotoxicity;
however, the extract of twigs were cytotoxic (LC50 546.7 μM). The cytotoxicity
was determined in four human tumor cell lines (MCF-7, NCI-H460, HeLa, and
HepG2) and also in non-tumor cells (porcine liver primary cells, PLP2) (Pires et al.
2015). On the other hand, the genotoxic potential of the flower extracts (100, 300,
and 500 mg kg−1 of body weight) on the blood and liver cells of male Wistar rats was
evaluated after 24 h of administration. A significant increase on DNA damage was
determined at the two highest doses mostly on liver cells; however, in both cases the
response was dose-dependent (Lemos et al. 2012). When genotoxicity was tested in
D. melanogaster by the somatic mutation and recombination test (SMART), it was
shown that both lapachol and β-lapachone induced apoptosis by generating oxygen-­
reactive species affecting cell cycle checkpoints. However, the results showed that
T. impetiginosa bark and stem powder dissolved in pure water were toxic but not
genotoxic by themselves, having a potential carcinogenicity activity. More experi-
ments are being performed in order to analyze interactions with chemotherapeutic
drugs (de Sousa et al. 2009).

In vitro cultures  In vitro propagation was achieved in MS culture medium with

Gamborg vitamins (MSG) (Murashige and Skoog 1962; Gamborg et al. 1968) and
Woody Plant Medium (WPM) (Lloyd and McCown 1980). The optimal conditions
were WPM supplemented with 20 μM BA and 1 μM IBA (MR: 4.37) and 40 d for
rooting in 1/2 MSG using 50 μM IBA induction (83% rooting) (Larraburu et al. 2012).

5.8  Allophylus edulis (A. St.-Hil., A. Juss. & Cambess.)

Hieron. ex Niederl. (Sapindaceae)

Synonymy  Allophylus edulis (A.  St.-Hil., A.  Juss. & Cambess.) Hieron. ex
Niederl. var. gracilis, Allophylus pauciflorus Radlk. var. rojasii, Allophylus edulis
(A. St.-Hil., A. Juss. & Cambess.) Hieron. ex Niederl. var. rosae, Schmidelia edulis
A. St.-Hil., A. Juss. & Cambess., Urvillea seriana Griseb.
112 5  Mesopotamia-Paranaense Forest

Vernacular name  Chal chal, coloradillo, conhú, kokú, picazú-rembiú (wild pigeon
food), pitanga, guacú or huacú, chichita, bacú, cocú, codocoypu, coquy, cuquito,
jocú, pucancho, puca-puca, sena, huaquito, vacú, vaquito (Ferrucci 2004).

General information  The species is endemic to British Guiana, Brazil, Paraguay,

Bolivia, Uruguay, and Argentina. In Argentina it is found in JUJ, SAL, TUC, CAT,
FOR, CHA, MIS, COS, SFE, ERI, and BAI to the Martín García Island. It grows in
humid and shady places, in serrano pastureland, and in the Chaqueño forest in the
transition with the Yungas in Valle de Lerma (SAL). This plant is commonly used as
a source of tannins and essential oils (Resico, 2007). Sometimes the wood is used
as fuel (Tortorelli, op. cit.: 536). Aloja de chalchal is a fermented sweet drink made
with the fruits (Ferrucci 2004). Fruits are appreciated by birds, inambues, agutíes,
tangaráes, pigeons, thrushes, and Artibeus lituratus (Bodrati and Haene 2004). It is
also considered an ornamental species. It was tested the development of the species
in petroleum-contaminated soils; since roots and eophyll structure showed little
variation, the authors concluded that A. edulis has the potential to be used for phy-
toremediation (Nogueira et al. 2011).

Botanical characteristics  It is a perennial tree or shrub, 4–15 m high, 10–50 cm

trunk diameter, and a thin bark. Bark is covered by scales. Branches are chestnut-­gray,
with a smooth bark and linear lenticels. Leaves are alternate, composite, trifoliate,
with sessile or subsessile oblanceolate or elliptic foliates, mucronate, toothed-serrated,
with an acute apex, and without stipules. Petiole is 1–4 cm length. Inflorescences are
spiciform or sub-spiciform, puberulous, or pubescent, with a 1.0–4.5 cm peduncle and
pubescent bracts. The male flower is 8 mm × 8 mm, with four free concave uneven
green sepals, ovate the two external and subcircular the two internal; with four free
white petals, with ovate 1 mm limb and a 0.77 mm nail and with long hairs in the
internal surface particularly in the margins; and with seven stamen that surround the
superior ovary. The feminine flowers are 2–4 mm long, with 1.25–2.00 mm glabre
sepals; 1.25–2.5 ciliated petals, with a bifid scale, a gold-yellow or orange glabrous
nectary disc; and 1.5–3.0 mm stamens in the staminate flower, smaller in the pistillate
flower, pubescent filaments, and ovary with disperse glandular hairs. The fruit is sub-
globose or ovoid, 8 mm diameter, reddish when ripe. The seeds are subglobose, 4 mm
diameter, brown with a small aril. Blooming is in September–October. Fructification
is in November–January (Ferrucci 2004; Malizia et al. 2018).

Ethnomedical information  Traditionally, the leaves were used as anti-­

inflammatory and for gastrointestinal disorders (Körbes 1995; Fatori Trevizan et al.
2016). Also, the leaves and stems were used in popular phytotherapy as a digestant
and for hepatitis treatment (Amat and Vajia 1991). In the province of MIS, leaves
are added to tereré, a beverage made with I. paraguariensis. Also, this is used in
cases of digestive disorders. It was reported the use of leaves and stems by the
Polish migrants in Misiones in cases of slow digestion, stomach and liver refreshing
(humoral medicine), stomach cooling (humoral medicine), liver pains, hepatitis,
­liquid retention in organism (diuretic), and empacho; ocasionally,  the fruits are
taken as snack (Kujawska and Pieroni 2015). The decoction and the infusion made
5.8  Allophylus edulis (A. St.-Hil., A. Juss. & Cambess.) Hieron. ex Niederl.… 113

with the leaves are used as analgesic in cases of cardiac pain, cancer, and stomach-
ache (Rondina et al. 2006).

Chemical data  It has phenolic compounds, including bergenin, quercetin 3-O-(2”-

O-galloyl)-glucoside, vitexin-2”-O-α-rhamnoside, isovitexin, vicenin, vitexin and
quercetin-3-O-glucuronide dihydroisocoumarin, L-quebrachitol,
6,7-­epoxycaryophyllene, spathunelol, sitosterone, lupeol, 2-oxo-13-hidroxy-neo-
cleroda-3,14-diene, and sitosterol. Viridiflorol is the main component of the EO
(30.88%) (Fig.  5.12). Other compounds, such as α-thujene, caryophyllene,

Fig. 5.12  Chemical compounds found in Allophylus edulis (A.  St.-Hil., A.  Juss. & Cambess.)
Hieron. ex Niederl
114 5  Mesopotamia-Paranaense Forest

β-selinene, bicyclogermacrene, δ-cadinene, β-atlantol, α-cadinol, and andledol,

were present in concentrations ranging from 3.02% to 7.87%. Leaves also contain
cyanogenic compounds and alkaloids (Bandoni et  al. 1972; Ariswa et  al. 1989;
Rastogi and Mehrotra 1990, 1993; Yajía et al. 1999; Díaz et al. 2008; Ragonese et
Milano, op. cit.: 1984; Fatori Trevizan et al. 2016).

Biological activities  A. edulis extract have pharmacological activities such as anti-­

hepatotoxic, negative ionotropic effect, inhibition of angiotensin converting enzyme
(ACE) and β-glucuronidase, cytotoxicity to KB cells, antioxidant activity, and anti-
microbial effect against S. aureus, as well as low toxicity (Ariswa et  al. 1989;
Hoffmann-Bohm et al. 1992; Matsunaga et al. 1997; Tirloni et al. 2015a, b).
The EO obtained by hydrodistillation of fresh leaves and the isolated viridiflorol
were in vitro tested as antioxidant using DPPH and ABTS reagents and as antimi-
crobial against Mycobacterium tuberculosis H37Rv (ATCC 27294). The samples
exhibited moderate effect in both assays, comparable to those of the commercial
antioxidants butylated hydroxytoluene (BHT) and ascorbic acid. As for the antimi-
crobial activity, MIC values were measured with respect to two standard antituber-
cular (anti-TB) drugs, isoniazid (MIC 0.05  mg  ml−1), and rifampicin (MIC
0.01 mg ml−1). The EOs (MIC 157.5 mg ml−1) and viridiflorol (MIC 190.0 mg ml−1)
exhibited promising activity. As for the anti-inflammatory activity, viridiflorol pro-
duced a significant inhibition of paw edema caused by carrageenan, and the oral
administration of the EOs (30 and 100 mg kg−1) inhibited leukocyte migration by
62 ± 5% and 35 ± 8%, respectively, 4 h after the carrageenan injection. Additionally,
the oral administration of viridiflorol (3 and 30 mg kg−1) also inhibited leukocyte
emigration by 71 ± 5% and 57 ± 3%, respectively, in a dose-dependent manner. The
authors have found that the pure viridiflorol and EOs showed anti-inflammatory
activity in comparable doses, concluding that viridiflorol is responsible, at least in
part, for the anti-inflammatory action of the EOs (Fatori Trevizan et al. 2016).
The ethanolic extracts of twigs were deterrent against aphids (Myzus persicae,
Epilachna paenulata, Rhopalosiphum padi) and have a lower deterrent activity
against chewers. Fractionation resulted in a lower activity against aphids and a higher
one against chewers. A synergism against aphids and an antagonism against chewers
in the crude extract are responsible of the activities demonstrated (Díaz et al. 2014).
Toxicity  Preliminary assays showed an increase in liver weight only at a dose of
5 g kg−1 of ethanolic extract suggesting hepatotoxicity. Hence, the authors consid-
ered that the extract has low toxicity (Tirloni et al. 2015a, b).

5.9  Passiflora caerulea L. (Passifloraceae)

Synonymy  Passiflora caerulea L. var. regnellii, Passiflora caerulea L. var. glauca,

Passiflora caerulea L. var. imbricata, Passiflora caerulea L. var. angustifolia,
Passiflora caerulea L. var. glaucophylla.
5.9  Passiflora caerulea L. (Passifloraceae) 115

Vernacular name  Granadilla, mburucuyá (fly hatchery), maracuyá (Portuguese),

flor de la pasión (blue passion flower), pasionaria.

General information  The name of the species comes from the first observations
made by the Spanish conquerors that arrived to America. They saw the symbol of the
crucifixion of Christ in the flower, hence the name of pasionaria (passion flower). In
the polychrome filaments, they saw the crown of thorns and the five wounds of
Christ; the three styles were the nails and the stamens the hammers that sank these
nails. The sepals and petals, in number of ten, would represent the Apostles. The two
that are not represented are Judas and Peter, one for his betrayal and the other, for
having denied Christ. P. caerulea is a bulbous vine that grows in BAI, CAT, CHA,
COR, COS, DFE, ERI, FOR, JUJ, LPA, LRI, MEN, MIS, SAL, SDE, SFE, SJU,
SLU, and TUC. There were reported 18 species growing in Argentina. Among them,
the species with medical properties are P. caerulea, P. capsularis, P. cicinnata, P.
edulis, P. elegans, P. foetida, P. misera Kunth, P. mooreana, P. palmatisecta, P.
suberosa, P. tenuifolia, and P. umbilicata (Barboza et al. 2009).

Botanical characteristics  It is a woody, climbing plant with a glabrous stem, with

tendrils to climb. The glabrous leaves are alternate, sectate palmate with five lanceo-
late lobes, dark green on the adaxial side, and pale bluish green on the abaxial one,
and an entire margin. The petiole has nectaries and small and caducous stipules. The
large flowers are solitary, hermaphrodites, with a calix with 4–5 free sepals and 4–5
free petals, with peduncles. The crown has filaments (radii), blue on the apex, white
on the center, and purple on the base, five free or fused stamens, and a nectary
(Fig. 5.13). The fruit is an ovoid fleshy berry, 6 cm-length, orange on the outside and
an edible red endocarp. The obcordate or cuneate seeds have an intense perfume

Fig. 5.13  Passiflora

caerulea L. flower. MA
Alvarez personal collection
116 5  Mesopotamia-Paranaense Forest

(Gupta 1995; Lahitte et al. 1999; Deginani 2001; Şesan et al. 2016). The striking
corolla functions as attractant to pollinators by visual and olfactory stimuli (Varela
et al. 2016).

Ethnobotanical information  The dried aerial parts are used to prepare an infusion
(usually 1%), with attributed properties as sedative, hypotensive, cardiotonic, anti-­
spasmodic, antiscorbutic, diuretic, anti-icteric, and emmenagogue, and to fight alco-
holism. The decoction of the aerial parts is used to treat urinary and respiratory tract
infections, diarrhea, catarrh, and pneumonia. Leaves, flowers, fruits, and stems are
used as sedative, anxiolytic, cordial, anthelmintic, calmative, eupeptic, diuretic,
emmenagogue, and contraceptive. The leaf is used as hypotensive, bradycardic, anti-
cephalgia, and sedative. Fruits are taken as diuretic, antiscorbutic, and anti-­jaundice.
Roots are considered as antispasmodic, antiphlogistic, toxic, narcotic, emmena-
gogue, sedative, contraceptive, and anthelmintic. The infusion or the syrup made
from the root is taken against pneumonia. Finally seeds are considered hypnotic.
Sometimes the infusion is associated with Melissa officinalis, Tilia spp., Piper
methysticum, or Valeriana officinalis for treating insomnia and with Crataegus oxy-
acantha to treat tachycardia and palpitations (Martínez Crovetto 1981; Barboza
et al. 2009; Petenatti et al. 2014; Alonso and Desmarchelier 2015).

Chemical data  The dried aerial parts contain caffeic acid, coumarin, umbellifer-
one, harman (passiflorine,1-methyl-9H-pyrido[3,4-b] indole), chrysin
(5,7-­dihidroxiflavone), tetraphyllin-4-sulphate, and epitetraphyllin- B4-sulphate.
Leaves contain gynocardin and the flavone schaftoside. Vitexin and isoorientin were
reported in leaves and calli. Greenery stems have steroids, saponins, peroxidases,
and heterosides. Fruits contain glucose, fatty acids, the flavone chrysin, cyanogenic
glucoside sulphate tetraphyllin 8–4-sulphate, and epitetraphyllin B-4-sulphate
(Fig. 5.14). The alcoholic extracts contain more bioactive compounds than petro-
leum ether extracts; saponins were only determined in ethanolic extracts. Maximum
flavonoid content was 12.82 mg Q g−1 DW (Farag et al. 2016; Şesan et al. 2016).
The intense aroma from flowers were attributed to sesquiterpenes secreted by the
periantum, α-copaene, 2(E),6(E)-farnesol, D-germacrene, (−)-α-cubebene, and to
the ester methyl-linolenate (Martínez Crovetto 1981; Seigler et al. 1982; Speroni
et al. 1996; Pereira et al. 2004; Áquila et al. 2005; Barboza et al. 2009; Varela et al.
2016; Hadas et al. 2017).

Biological activities  The ethanolic extract from leaves had higher anticonvulsant,
analgesic, anti-inflammatory, antipyretic, and antioxidant activities than the aque-
ous extract (Şesan et al. 2016; El-Askary et al. 2017; Hadas et al. 2017). After frac-
tionated extraction from the ethanolic extract, the  ethyl acetate fraction was the
most active. The isolated compounds were lucenin, 2,4-hydroxycinnamic acid, and
chrysin 6-C-β-D-glucoside. The biological activities were attributed to C-glycosyl
flavones and hydroxycinnamic acid derivatives (El-Askary et al. 2017). The antibac-
terial activity was endorsed to flavonoids (Anesini and Perez 1993; Dhawan et al.
2004; Pérez Ibáñez et al. 2017).
5.9  Passiflora caerulea L. (Passifloraceae) 117

Fig. 5.14  Some of the chemical compounds determined in Passiflora caerulea L.

Chrysin (1  mg  kg−1) induced significant anxiolytic behavior in mice, without
myorelaxant effect (dose range 0.6–30 mg kg-1) suggesting that it was an anxiolytic
devoid of sedative or muscle-relaxing collateral effect, unlike diazepam (Wolfman
et al. 1994; Dhawan et al. 2004; Duke et al. 2009; Feliu-Hemmelmann et al. 2013).
It was reported that chrysin prevented the expression of tonic-clonic seizures
118 5  Mesopotamia-Paranaense Forest

induced by pentylenetetrazol by acting as a ligand for central as well as peripheral

benzodiazepine receptors. The content of benzodiazepine-like compounds in P. cae-
rulea was assumed by the authors (Medina et al. 1990).
Other reported activities were as phytoestrogens with potential clinical and ther-
apeutic utilizations against physiological and biochemical effects of aging (Dhawan
et al. 2002) and as anthelmintic and acaricidal (Duke et al. 2009).
In vitro culture  In vitro cultures of P. caerulea were established. The in  vitro
response of different explants and biotypes on MSG culture medium with 1 mg l−1
of benzyl adenine (BA) as plant growth regulator produced calli or regenerated the
plant (Nakayama 1966; Severin et al. 2011).

Legal status  P. caerulea and P. incarnata were approved for human use by the
ANMAT (Res. 2673/99, annex III).

5.10  Blechnum occidentale L. var. occidentale (Blechnaceae,

Pteridophyta)

Synonymy  Blechnum glandulosum Kaulf. ex Link, Blechnum suburbicum Vell.,

Blechnum glandulosum Kaulf. ex Link var. elongatum, Blechnum caudatum Cav.,
Blechnum occidentale L. var. caudatum, Blechnum occidentale L. var. pubirhachis,
Blechnum occidentale L. var. lacerata, Blechnum cognatum C. Presl, Blechnum car-
tilagineum Schkuhr, Blechnum appendiculatum Willd., Blechnum meridionale
C.  Presl, Lomaria campylotis Kunze, Mesothema campylotis (Kunze) C.  Presl,
Blechnum campylotis (Kunze) J.  Sm., Blechnum mucronatum Fée, Blechnum
lomarioides Sodiro, Blechnum scaberulum Sodiro, Blechnum glandulosum Kaulf.
ex Link var. pallida, Blechnum flaccisquama A. Rojas.

Vernacular name  Doradilla, hammock fern, blechnum fern.

General information  Blechnum L. is a worldwide distributed genus with around

50 species in America. They are land ferns, sometimes epiphyte or scandant, with
numerous scales and often fibrous roots. They grow in rainy jungles, mountain for-
ests, ravines, savannahs, wastelands, along rivers, and streams being rarely found in
dry regions (Tryon and Tryon 1982; Kramer et al. 1990). B. occidentale is found
from the south of North America to South America; in Argentina there are 20–21
taxa mainly in the provinces of JUJ, MIS, SAL, and TUC (Ponce 1996; Durán 1997).

Botanical characteristics  B. occidentale is a perennial, terrestrial, or rare rupicolas

(epipetric) species with an upright decumbent stiff with bicolor scales. The fronds
are monomorphic, subdimorphic, or dimorphic with a petiole with triangular scales,
mainly in the base. The leaf blade is pinnate at the base becoming pinnatisect towards
5.11  Maytenus ilicifolia Mart. ex Reissek (Celesteraceae) 119

the apex, truncated at the base, glabrous or with hyaline trichomes. The rachis is
glabrous or with septate trichomes. Pinnae are in 13–28 pairs, 1.2–9.5 cm length ×
0.6–1.6 cm wide with a free venation; veins are undivided or 1–4 bifurcated; pinnae
have an entire to serrate margin, not articulated to the rachis. Sori are placed on a
long or rarely short linear vascular commissure, parallel and close to the main vein
of each segment, sometimes extending beyond the commissure. Sporangia have a
three-rowed stalk with ellipsoidal to spheroidal monolete spores (kidney-shaped
grains with a single elongate scar) (Dittrich et al. 2007; Pessôa Santiago et al. 2014;
Dittrich et al. 2015).

Ethnobotanical information  In Brazil this fern has been used to treat inflamma-
tory and pulmonary diseases, urinary infections, and liver diseases (Barros and
Andrade, 1997). In Argentina the entire plant was used to treat lung and urinary
disorders. A not specified part was taken as an infusion and decoction to dissolve
renal calculi and for treatment of nervous hysteria (Barboza et al. 2009).

Chemical data  The dried leaf contains rhodoxanthin, lutein epoxide, lutein,
adonixanthin, zeaxanthin, violaxanthin, and antheraxanthin (Fig. 5.15) (Barboza
et al. 2009).

Biological activities  The methanolic extract from air-dried and powdered blades
was tested on male Wistar rats or Swiss Webster mice for its activity as anti-­
inflammatory (paw edema model in mice) and antinociceptive (writhing test). The
administration resulted in an inhibition of the inflammatory effect caused by car-
rageenan and in an antinociceptive effect in acetic acid-induced writhing and for-
malin tests. The authors suggested that those effects are related to the arachidonic
acid cascade and/or modulation of pro-inflammatory molecule production (Nonato
et al. 2009). The entire plant had hypothermic, diuretic, and antibacterial activity
(Barboza et al. 2009).

Toxicity  The acute toxicity of the methanolic extract was tested giving to mice a
100 or 1000 mg kg-1 intraperitoneal dose. After 14 days any death was produced.
Besides, any motor performance alteration was not detected (Nonato et al. 2009).

5.11  Maytenus ilicifolia Mart. ex Reissek (Celesteraceae)

Synonymy  Celastrus spinifolium Larrañaga, Monteverdia ilicifolia (Mart. ex

Reissek) Biral, Maytenus aquifolium Mart., Maytenus hassleri Briq., Maytenus
ilicifolia Mart. ex Reissek f. angustior Briq., Maytenus pilcomayensis Briq.

Vernacular name  Pus pus, congorosa, concorosa, quebrachillo, molle morotí,

cangorosa, capororoca, molle espinoso, cancerosa, sombra de toro.
120 5  Mesopotamia-Paranaense Forest

CH3
CH3 CH3 CH3 H3C O

CH3 CH3 CH3 CH3

O
CH3

rhodoxanthin

OH
OH

OH OH

lutein zeaxanthin

OH
O
OH

O O
HO HO

violaxanthin antheraxanthin

Fig. 5.15  Chemicals found in Blechnum occidentale L. var. occidentale

5.11  Maytenus ilicifolia Mart. ex Reissek (Celesteraceae) 121

General information  The genus Maytenus include about 200 American species
of which 11 are in Argentina and 77  in Brazil (Hurrell and Bazanno, 2003). M.
ilicifolia is native from South Brazil, Paraguay, Bolivia, Uruguay, and Argentina
where it is mainly found in the provinces of BAI, CHA, COS, ERI, FOR, MIS,
SAL, and SFE. It grows in subtropical temperate climates, with well-drained clay
soils and a high content of organic matter, up to 1200–2000 m.a.s.l (Hurrell and
Bazanno 2003).

Botanical characteristics  M. ilicifolia is a dioecious perennial shrub, up to 5 m

height (up to 10  m height in the Mata Atlántica). The root is long and pivoting
(Brussa Santander and Grela González 2007). Leaves are simple, entire, coriaceous,
with alternate phyllotaxy, a short petiole and a lanceolate laminae (2–7 cm length
and 1.5–3.0 cm width). The leaf apex is acute with toothed thorny margins (2–7
pairs), the leaf base is cuneate or rounded, the midrib is bright green more promi-
nent in the abaxial side which is paler (Fig. 5.16). The inflorescence is solitary or in
axillar yellow fascicles with reddish bracts. Flowers are small, actinomorphic, with
a small pedicel, five reddish sepals united by the base, five free yellow ovate or
entire or dented petals, five stamen, five emarginated discs, and two loculate ovary
with four included or emerging ovules. The fruit is a reddish ovoid or ellipsoid 1 cm
length capsule with a thin aryl containing 1–4 reddish seeds. Blooming is during
spring and fructification in summer; fertilization is entomophile (Cabrera 1965;
Hurrell and Bazzano 2003; Duarte and Debur 2005).

Fig. 5.16  Maytenus

ilicifolia Mart. ex Reissek,
Celesteraceae. Buenos
Aires, Botanical Garden.
MA A Personal collection
122 5  Mesopotamia-Paranaense Forest

Ethnobotanical information  The infusion made with the entire plant is used in
folk medicine for fertility regulation and as antiasthmatic. The Guaraníes have
taken it as contraceptive. The aerial parts are used as vulnerary, contraceptive, hypo-
tensive, cardiotonic, antidiarrheal, decongestant, and odontalgic, for lumbago, kid-
ney disorders, gastric ulcers and gastritis, and articular pains. The decoction made
with the flowers is taken as anti-inflammatory and those from leaves as antiasth-
matic and antiseptic. The infusion of the leaves is considered as emmenagogue,
sialagogue, astringent, antispasmodic, and contraceptive. The roots are taken as
diuretic, contraceptive, and emmenagogue. In Argentina and South Brazil, the
decoction of leaves and stems is added to the beverage mate in cases of bloody
ulcers, hypertension, and articular pains, as depurative; as sialagogue, to treat
asthma; and as antitumoral (Bandoni 1976; Toursarkissian 1980; Martínez-Crovetto
1987; Arenas 1997; Hurrell and Bazzano 2003; Alonso 2004; Scheffer et al. 2005;
Barboza et al. 2009; Alonso and Desmarchelier 2015).

Chemical data  The leaves have terpenes, phenols, phytosterols, and alkaloids
(Figs. 5.17 and 5.18). The dried leaves contain the terpenoids α-amyrin, cangorosin
A and B; the triterpenoids maytefolins A, B, C, uvaol-3-caffeate, erythrodiol, betu-
lin, betulin-3-caffeate, moradiol, erythrodiol-3-caffeate, 20α-hydroxymaytenin,
22β-hydroxymaytenin, maytenin, celastrol, and pristimerin; the tannins afzelechin,
epiafzelechin, catechin, epicatechin, gallocatechin and epigallocatechin, 4O-methyl-
(epi)-catechin, and 4’O-methyl (epi) catechin; the flavones quercetin and kaemp-
ferol; and the maytansinoid alkaloids maitansine, maytanprine, and maytanbutin.
Also, it was found friedelin, friedelan-3-ol, α-tocopherol, simiarenol, lupeol, lupe-
none, β-sitosterol, stigmasterol, campesterol, ergosterol, brassicasterol, squalene,
hexadecanoic acid, T1 and T2 tocopherols, cangorin F, G, H, I, J, phytol, vitamin E,
dodecanoic acid, and geranyl acetate, and Fe, K, Mg, S, Na, and Ca. The dried root
bark contains the sesquiterpenes cangorin A, B, C, D, E, F, G, H, I, and J and the
triterpenes friedoolean-24-al-3-en-3-ol-2-on-29-oic acid (cangoronine), friedool-­
ean-­1-en-29-ol-3-one (ilicifoline), maytenoic acid, D:B friedoolean-5-en 3β,
29-diol, D:A friedoolean-29-ol-3-one, pristimerin, salasperimic acid, isopristimerin,
isotingenone, and 6-oxotingenol. The roots contain the triterpenoids milicifolines
AD (Marini-Bettòlo 1981; Ahmed et al. 1981; Itokawa et al. 1991; Silva and Récio
1992; Shirota et al. 1994; Cordeiro et al. 1999; Niero et al. 2001; Buffa Filho et al.
2002; Alonso 2004; Ohsaki et  al. 2004; Soares et  al. 2004; Pereira et  al. 2005;
Gutiérrez et al. 2007; Barboza et al. 2009; Leite et al. 2010; Santos et al. 2012).

Biological activities  The dried leaves extracts have barbiturate potentiation, anti-
ulcer, antimicrobial, antitumoral, antioxidant, antinociceptive, anti-inflammatory,
and relaxant activities. The root bark has activity as antioxidant, myeloperoxidase
inhibitor, and as cytotoxic against V79, KB, L1210, VJ300, KU 1920, and P 388
cells (Pereira et al. 1992; Jorge et al. 2004; Barboza et al. 2009; Leite et al. 2010;
Alonso and Desmarchelier 2015). The antiulcerogenic and antigastritis effects were
attributed to the terpene friedelin, to the tri- and tetraglycosid flavonoid derivatives,
and to the combined action of different phytocomplexes (Souza-Formigoni et  al.
1991; Cunha 2003).
5.11  Maytenus ilicifolia Mart. ex Reissek (Celesteraceae) 123

CH3
O
CH3

CH3
O
CH3 CH3
O
O H3C CH3
CH3 O
O CH3

CH3

CH3 CH3

HO HO
CH3 O

α-amyrin cangorosin B

CH2
H 3C CH3
H 3C CH3
OH
CH3 CH3 CH3
HO
OH
CH3 CH3
CH3

O O
CH3
H3C CH3 CH3

maytefolin A maytefolin C

Fig. 5.17  Some terpenoids from Maytenus ilicifolia Mart. ex Reissek leaves

The leaf infusion inhibited gastric secretion induced by histamine and acidity and
reduced dyspepsia symptoms (Carlini et al. 1988; Baggio et al. 2007). Also, the oral
administration of spray-drying powders maintained antiulcer activity in Wistar male
rats (Martins et al. 2003). The lyophilized aqueous extract from leaves inhibited the
acid secretion in isolated frog gastric mucosa (Ferreira et al. 2004). The arabinoga-
lactan isolated from M. ilicifolia leaves showed a dose-related potent gastro-­
protective activity against gastric lesions induced by ethanol after oral administration
(10–100 mg kg−1 body weight) to female Wistar rats and female Swiss mice (Baggio
et al. 2012).
The ethanolic extract from dried leaves (intraperitoneally, 200 mg kg−1 day−1 for
20 days and oral administration, 300 mg kg−1 day−1 for 30 days) did not arrest sper-
matogenesis in Swiss mice, although minor ultrastructural alterations were observed
(Montanari et al. 1998). Antitumoral and antioxidant effects were attributed to qui-
nonemethide triterpenes, 22β-hydroxymaytenin, and maytenin (Buffa Filho et  al.
2004). Patients with advanced stage of neoplasia were treated with 150 μg kg−1
124 5  Mesopotamia-Paranaense Forest

Fig. 5.18  Maytansinoid alkaloids from Maytenus ilicifolia Mart. ex Reissek

day−1 and 450 μg kg−1 day−1 of triterpenes extracted from Maytenus sp. enriched
in maiteina, by intravenous route. Patients with carcinoma epidermoid of the pillars
of the amygdala or tonsil, the base of the tongue, and the larynx were experimented
an injury reduction of around 40–60% as well as the disappearance of bleeding.
Subjective improvements were verified with respect to pain, asthenia, and anorexia.
It was not observed a positive response for carcinomas of uterus and ovary, osteo-
sarcoma, and chondrosarcoma (Santos-Oliveira et al. 2009).
References 125

Toxicity  There are contradictory reports about the M. ilicifolia interference with
embryo-fetal development. On one side, it was reported that M. ilicifolia interferes
with uterine receptivity to the embryo for its estrogenic activity in pregnant mice that
produced an uterotropic effect (Montanari and Bevilacqua 2002). On the other side,
it was reported that M. ilicifolia hydroacetonic extract (70:30) resulted nontoxic to
Wistar pregnant rats apparently not interfering with pregnancy. Furthermore, fertil-
ity of female rats was not altered (Oliveira et al. 1991; Cunha et al. 2014).

In vitro cultures  M. ilicifolia was micropropagated from leaves in MS medium

(Murashige and Skoog 1962)  supplemented with BAP 0.4  mg  l−1 and NAA
0.2  mg  l−1. Calli were obtained from leaf explants from micropropagated plants
placed on MS medium supplemented with Kin 1.0  mg  l−1 and 2,4-D 1.0  mg  l−1
(Buffa Filho et al. 2004).
Legal status: The species in not included in FNA. However, it was included in
the Primary Health Care Program from the province of MIS in 2003. In Brazil it
is included in the Brazilian Pharmacopeia as phytotherapic (Alonso and
Desmarchelier 2015).

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Chapter 6
Puna: A Surrealistic Landscape
in the Argentinean Highlands

6.1  Introduction: The Puna

The Puna is a geographical region located in the northwest of Argentina (the prov-
inces of JUJ, SAL, CAT, TUC, LRI, and SDE). It is an extensive highland, or high
mountain plateau, typical of the central area of the cordillera de los Andes (Fig. 6.1).
The limits of the Argentine Puna range from the Cordillera de San Buenaventura
(North of the San Francisco mountain pass, in the province of CAT) to the South, to
the extreme North of the province of JUJ, and from the bordering cordillera with
Chile, by the West, to the mountains of the Eastern Cordillera of the provinces of
CAT, SAL, and JUJ, by the East.
The lagoons in there form closed basins without drainage that were the origin of
extensive salt pans such as de Salinas grandes in the provinces of JUJ and SAL
(Fig.  6.2). The hills, of great height and rounded shape (volcanic morphology),
exceed 5000–6000 meters in height with volcanoes such as Pissis (6882  m),
Llullaillaco (6739  m), Ojos del Salado (6864  m), Bonete Chico (6759  m), Tres
Cruces (6749 m), and Walter Penck (6658 m).
At altitudes higher than 4300–4500 m.a.s.l. predominates the geological and bio-
climatic conditions of the high Andes eco-region. The climate is cold and dry, with
a large daily temperature range. The annual average temperature is less than 8 °C;
in summer the temperature can reach 30 °C, in winter less than −20 °C. The annual
average rain is 100–200 mm.
The fauna is well characterized; the domestic species are the llama (Lama glama):
the wild species are puma (Puma concolor), the guanaco (Lama guanicoe), and
numerous endemics species such as the vicuña (Vicugna vicugna), huemul del Norte
or taruca (Hippocamelus antisensis), Andean cat (Leopardus jacobitus), and real
skunk (Conepatus chinga rex). Among the rodents are chinchillas (Chinchilla chin-
chilla). The great diversity of birds is associated to high-altitude wetlands (salt flats
and lagoons); there are flamingos such as the parina James (Phoenicoparrus jamesi)
and big parina (Ph. andinus), ducks such as the guayata (Chloephaga melanoptera)

© Springer Nature Switzerland AG 2019 137

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_6
138 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

Fig. 6.1  Puna, section in the province of Jujuy, Argentina. Personal collection P G Alvarez

and puna duck (Spatula puna), gallareta cornuda (Fulica cristata), tero serrano
(Vanellus resplendens), Andean becasina (Gallinago andina), and the plover from
the Puna (Charadrius alticola). Among the terrestrial birds are the suri (Rhea pen-
nata, ñandú petiso), partridges such as quiula puneña (Tinamotis pentlandii), and
small species such as Geositta punensis and Muscisaxicola juniensis.
Flora is substantially affected by the climatic conditions. The lack of rain defines
a vegetal carpet of bushes and herbs, and in some areas soils without vegetation. The
dominant flora is the shrub steppe with dispersed bushes. Local edaphic microcli-
mates determine the existence of small sectors with different physiognomies.
6.1 Introduction: The Puna 139

Fig. 6.2  Salinas grandes

(a) and llama (Lama lama)
in the province of Jujuy
(b), Argentina. Personal
collection PG Alvarez

Soils  are stony or saline, scarcely covered by vegetation, such as Maihueniopsis

glomerata, very susceptible to erosion. In the Northern and Eastern areas, where the
annual rainfall exceeds 400  mm, predominate the tolas (Baccharis incarum).
Towards the South and West, in the high Andean steppe dominate the chijua
(Baccharis boliviensis), tolilla (Fabiana densa), añaguilla (Adesmia tucumanen-
sis), rica-rica (Acantholippia hastulata), suriyanta (Nardophyllum armatum),
airampu (Opuntia soehrendsii), and canguía or cailla (Tetraglochin cistatum)
among other species. Open groves of queñoa (Polylepis tomentella) grow on slopes
and ravines between 3800 and 4300 m height.
The cordillera de los Andes has two parallel cords, Western and Eastern, sepa-
rated by ravines and valleys. The Western cord shows higher heights (Chañi 6200 m).
The Eastern cord is of less height (Cerro Negro de Zucho around 5000  m). The
ravines, elongated and narrow valleys, are the communication routes of this area.
The main economic activities in the Puna are livestock (herds of sheep and lla-
mas raised for wool and meat production), subsistence agriculture, and mining
(gold, copper, lithium). The Puna reunites the optimal environmental conditions to
develop solar energy with minimal impact on the ecosystem.
140 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

6.2  Acacia caven (Molina) Molina var. caven (Fabaceae)

Synonymy  Acacia adenopa Hook & Arn., Acacia cavenia Colla, hom. illeg.,
Acacia cavenia (Molina) Hook & Arn, Acacia farnesiana (L.) Wild., Acacia farne-
siana (L.) Willd. var. heterocarpa, F. cavenia, Mimosa cavenia Molina, Mimosa
caven Molina, Vachellia farnesiana (L.) Wight & Arn. f. brachypoda, Vachellia
farnesiana (L.) Wight & Arn. f. cavenia.

Vernacular name  Espinillo, espinillo negro, aromo criollo, tusca, aromo, aromo
criollo, churqui, aromita, aromito, cavén, espino.

General information  A. caven grows in Argentina, south of Brazil, Bolivia, Chile,

Paraguay, and Uruguay. In Argentina, it could be found in the provinces of BAI,
CAT, CHA, COR, COS, ERI, FOR, JUJ, LPA, LRI, MEN, MIS, SAL, SDE, SFE,
SJU, SLU, and TUC. It develops in flat areas from forests and in mountain range
areas (serranías), in the hillocks with high drainage, and also in previously flooded
areas, being removed when the strong humidity returns. Among other applications,
fruits, seeds, and pods are used to make high protein content flours, roasted seeds to
make a coffee-like beverage, and flowers to make a tea. It is also used for firewood,
to make utility poles, and handicrafts; it is considered a melliferous species and also
of use in perfumery (Ortiz 1966; Aronson and Ovalle 1989; Carrere 1990; Benedetti
2012). Additionally, it is used to feed cattle.

Botanical characteristics  A. caven is a deciduous tree, 2–6 m height, with a short

trunk and a wide tree top (Fig. 6.3). It has extended lenticellate and glabrescent or
pubescent branches, dark brown bark, with deep longitudinal cracks. Stipules are
whitish, with thin and acute 5.0–30.0 mm length light gray pubescent thorns in the
base, dark and glabrescent in the apex. Leaves are bipinnately compound with
3–15 pinnate pairs (1.0–2.5 length), numerous opposed sessile 1–4 mm length leaf-
lets with incospicuous veins and entire margins, glabrous or with scarce hairs on
the margins, usually with one petiolar gland. The inflorescence is a densely flow-
ered globose head with orange-yellowish 1 cm diameter flowers. Flowers are her-
maphrodite, actinomorphic, gamopetalous, pentamers, sessile, with pubescent
peduncles. The sparkling blackish or green-yellowish fruit is an indehiscent
legume, with 3–5 mm seeds, 25–30 per fruit, surrounded by a spongy substance.
Fruit size and shape is highly variable. Blooming is before foliation in spring, and
fructification is in summer (Spegazzini 1924; Cialdella 1994; Cialdella 1996;
Lahitte et al. 1999; Marzocca 1997; Ebinger et al. 2000; Pometti et al. 2007).

Ethnobotanical information  The root is considered purgative: the bark, deter-

gent, cicatrizing, and antidiarrheal. The dried leaf and stems are used as antiseptic,
to treat hypertension, diabetes, skin mycosis, and irritations, and as cicatrizing. The
decoction of the leaf is taken as antidiarrheal and diuretic. Flowers are sedative.
The  combination of seeds and leaf is astringent, antiseptic, anti-asthmatic, and
6.2  Acacia caven (Molina) Molina var. caven (Fabaceae) 141

Fig. 6.3  Acacia caven (Molina) Molina var. caven (Fabaceae) tree (a), fruits and leaves (b), A.
caven parasited by Ligaria cuneifolia (c). MA Alvarez personal collection

a­ ntitussive. Some specimens were found with cyanogenic activity (Aronson and
Nash 1989; Ebinger et al. 2000; Martínez 2007; Barboza et al. 2009).

Chemical data  The dried leaf has the flavonols quercetin and quercitrin. Fresh
flowers contain p-anisaldehyde, (E,E)-farnesyl acetate, eugenol, benzyl alcohol,
methyl salicylate, and cuminyl alcohol (4-propan-2-ylphenyl methanol) (Fig. 6.4)
(Lamarque et al. 1998; Barboza et al. 2009).

Biological activities  The ethanolic extract from leaves and stems has cytotoxic,
antimicrobial, and antifungal activity (Quiroga et al. 2004). The infusion of leaves
showed an arterial contractile effect in a dose-dependent manner in pre-anesthetized
rats, which was prevented by the α-adrenergic antagonist phenoxybenzamine
(Adrados et  al. 1997; Alonso and Desmarchelier 2015). The dried seeds are
β-glucuronidase inhibitors (Barboza et al. 2009). The cortex, rich in tannins, is used
as astringent and to cure blows and wounds (Ortiz 1966). The seeds are considered
digestive and stimulants (Ortiz 1966). A macerate made with fresh flowers and alco-
hol is used as sedative and in cardialgias. The infusion made with the fruits is anti-
hemorroidal (Carrere 1990; Benedetti 2012).
142 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

O H

H3C O CH3

O CH3 CH3 CH3

O.CH3 OH

farnesyl acetate p-anysaldehide cuminyl alcohol

O
OH
HO

OH

methyl salicylate eugenol benzyl alcohol

Fig. 6.4  Chemicals from Acacia caven (Molina) Molina var. caven

In vitro culture  Micropropagation was attained from axenic seedlings in

Broadleaved Tree Medium with IBA (0.49 and 1.2 μM) and BAP (0.44, 1.10, and
2.22 μM). Calli differentiated and developed longer roots with the addition of 9 and
10% sucrose without plant growth regulators (Abedini et al. 2000). Somatic embryo-
genesis was also achieved (Sharry et al. 2010).

6.3  Chenopodium ambrosioides L. (Chenopodiaceae)

Synonymy  Dysphania ambrosioides (L.) Mosyakin & Clemants, Ambrina ambro-

sioides (L.) Spach., Ambrina parvula Phil., Ambrina. spathulata Moq., Atriplex
ambrosioides (L.) Crantz, Blitum ambrosioides (L.) Crantz, Chenopodium anthel-
minticum L., C. integrifolium Boros., C. spathulatum Sieber, Chenopodium suffru-
ticosum Willd.

Vernacular name  Paico, paico macho, hierba hedionda, té de los jesuitas, pichim
tirao, paico hembra, yerba de Santa María, ambrosía, apasote, epazote, té de México.

General information  C. ambrosioides is a native species from temperate America,

although today it is found in almost all the temperate regions of the world (Lorenzi
and Matos 2002). The plant is considered by the World Health Organization as one of
the most used among traditional medicines in the world (Lorenzi and Matos 2002). It
grows in the edges of cultivation lands and borders of gardens and parks. It adapts well
to clay, sandy, xerophytic, and sub-xerophytic soils and could be found at altitudes
between 0 and 2760 m.a.s.l. (Alban 1984). It has a wide d­ istribution in the country,
6.3  Chenopodium ambrosioides L. (Chenopodiaceae) 143

growing in the provinces of BAI, CHA, OR, COS, DFE, ERI, FOR, JUJ, LPA, LRI,
MEN, MIS, RNE, SAL, SDE, SFE, SJU, SLU, and TUC (Barboza et al. 2009).

Botanical characteristics  C. ambrosioides is an annual, perennial aromatic herb,

40–100 cm height, pubescent. Stem is branched with erect or prostrate branches on
their lower section (Fig.  6.5). Leaves are ovate-lanceolate, serrated-toothed, pin-
nately nerved, 3–10 cm length, 1–5 cm width, with petiole. The inflorescence is a
spike, with numerous green flowers in a pyramidal panicle that bloom in summer.
The calix has 5 membranoceous or herbaceous sepals free up to the half, 5 stamens,
2–5 long stigmas. The fruit is a utricle; the shining black seeds are elongated (Gupta
1995; Giusti 1997; Tolaba 2006).

Ethnomedical information  Decoctions and infusions of inflorescences, leaves,

and roots of C. ambrosioides have been used for centuries as traditional anthelmin-
tic and dietary condiments against intestinal worms (Hernández 1571; Ratera and
Ratera 1980; Martínez Crovetto 1981a, b; Kliks 1985; Carballo et al. 2005). Also,
those preparations are taken as emmenagogue, digestive, diuretic, antispasmodic,
hypotensive, and diaphoretic (Bandoni et al. 2009; Barboza et al. 2009; González
et al. 2009; Da Silva et al. 2014; Navone et al. 2014; Neiva et al. 2014; Alonso and
Desmarchelier 2015; Flores and Pellegrini 2015; Moya and Escudero 2015). The
infusion made with the aerial parts is considered antitussive, antiarthritic, anti-
pyretic, and hemostatic. Also, it is used for stomachache, for asthma and tumors,
and as appetitive (Gómez Castellanos 2008; Barboza et al. 2009). In Northern Brazil
the leaves are blended with milk and taken for treating flu (Morais et  al. 2005).
Leaves are used in cases of muscle pain and bone ­fractures (Santayana et al. 2005;

Fig. 6.5  Chenopodium ambrosioides L. leaves and inflorescences. MA Alvarez personal

collection
144 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

Garcia et al. 2010). Ascaridol and commercial preparations of the essential oil are
prescribed for the treatment of intestinal worms (Patel 2017).

Chemical data  The aerial parts have (-)(1R’,4S’)-1,4-dihydroxy-p-menth-2-ene,

(-)(1R’, 2S’, 3S’, 4S’)-1,2,3,4-tetrahydroxy-p-menthane. The EOs from the aerial
parts have aritasone, limonene, ascaridol, cis-isascaridol, α-terpinene, camphor,
trans-ascaridol, p–cymene, pinocarvone, α-and β-pinene, geraniol, α- and
γ-terpineol, thymol, β-caryophyllene, γ-gurjunene, myrcene, phellandrene, safrole,
n-docosane, n-hentriacontane, n-heptacosane, n-octacosane, p-cymol, spinasterol,
terpinyl-acetate, terpinyl-salicylate, and triacontyl-alcohol. Also, stigmasterol,
β-sitosterol, octadecanoic acid, scopoletin and 1-piperoylpiperidine T kaempferol,
isorhamnetin, quercetin, kaempferol 3-rhamnoside-4′-xyloside, and kaempferol
3-rhamnoside-7- xyloside (Jain et al. 1990; Shah and Khan 2017). The entire plant
has saponins, trimethylamine, citric acid, salicylic acid, tartaric acid, succinic acid,
butyric acid, ferulic acid, malic acid, methyl salicylate, urease, and vainillic acid
(Fig. 6.6). The fruits have quercetin, kaempferol rhamnoside, anethole, and santo-
nin. The roots,  contain heterosides, and betaine (Kiuchi et  al. 2002; MacDonald
et al. 2004; Barboza et al. 2009; Bakker et al. 2011; Neiva et al. 2011; Barros et al.
2013; Moya and Escudero 2015; Patel 2017; Shah 2014; Shameem et al. 2019).

Fig. 6.6  Chemicals found in Chenopodium ambrosioides L. essential oil

6.3  Chenopodium ambrosioides L. (Chenopodiaceae) 145

Biological activities  The EO of the aerial parts has antioxidant (Kumar et al. 2007;
Barros et  al. 2013), antiparasitic (Avila-Blanco et  al. 2014; Moya and Escudero
2015; Patel 2017), antimalarial (Kiuchi et al. 2002), anti-leishmania (França et al.
1996; Monzote et al. 2007, 2011, 2014, 2018), anti-Trypanosoma cruzi (Rojas et al.
2010), cardiotonic, hypotensive, antialatoxigenic (Kumar et al. 2007), and spasmo-
genic activity (da Silva et al. 2014). Also, it has fumigant (Chu et al. 2011; Zhu et al.
2012) and repellent activity against head lice.
The aqueous extracts of bark and fruits showed a high antioxidant potential
determined by the ABTS assay, whereas petroleum ether bark extracts showed the
maximal % DPPH (Ajaib et  al. 2016). The infusion of aerial parts gave higher
DPPH scavenging activity, β-carotene bleaching, and thiobarbituric acid reactive
substances (TBARS) inhibition than the methanolic extract (Barros et al. 2013).
The aerial parts have carcinogenic, giardicidal (Neiva et al. 2011), antiulcerous,
antimalarial (Pollack et al. 1990), anti-leishmania (Monzote et al. 2007), and anti-
bacterial activity (MacDonald et al. 2004; Sá et al. 2016; Jesus et al. 2018). The
crude hydroalcoholic extract had plasmodicidal potential since it inhibited the para-
site growth in  vitro. In vivo it decreased parasitemia in infected BALB/c mice
(Nunes Cysne et al. 2016).
The leaves have antimicrobial, analgesic, and anti-inflammatory activity, and the
combination of leaves with flower has an antispasmodic activity.
The antiparasitic activity included activity against Ancylostoma duodenale,
Trichuris tricura, Ascaris lumbricoides, and Giardia lambia (Giove 1996; Neiva
et al. 2014). The EO and the ethanol extract from leaves were useful for controlling
canine Ancylostoma spp. (Moraes Monteiro et al. 2017), gastrointestinal parasites
of fighting cocks (Gallus domesticus) (Álvarez et  al. 2011; Cazorla and Morales
2013), young cattle (Clavijo-López et al. 2016), and the cattle tick Rhipicephalus
(Boophilus) microplus (Cunha dos Santos et al. 2013).
Intraperitoneal and oral treatment with the EO (30  mg  kg−1) had better anti-­
leishmania effect than the treatment with the reference drug, amphotericin B
(1 mg kg−1) (Monzote et al. 2007).
The bark macerated in petroleum ether was active against pathogens such as
S. aureus, P. aeruginosa, Brevibacillus agri, Trichophyton mentagrophytes, and C.
albicans with MICs ranging from 0.25 to 0.80 μg ml−1. In the case of M. tuberculo-
sis MIC was 0.5 μg ml−1 (Lall and Meyer 1999; Mabona et al. 2013; Jesus et al.
2018). Against Staphylococcus aureus the minimal zone of inhibition was
0.009 ± 0.02 mm at 0.7 μg ml−1.
The n-hexane, ethyl acetate, dichloromethane, as well as n-butanol fractions
showed moderate to significant activities especially against B. subtilus, K. pneu-
moniae, and S. epidermidis (Shah 2014). C. ambrosioides also has antifungal prop-
erties (Fenner et  al. 2006; Jardim et  al. 2010). The maximal antifungal zone of
inhibition against Aspergillus niger (16  ±  1.5  mm) was attained with methanol
extracts of fruits (Ajaib et al. 2016).
The in vitro activity from EOs of the aerial parts against various Candida spp.
showed that all the microorganisms were sensitive in a concentration-dependent
way, being C. albicans the most sensitive ATCC 2091 (MIC: 2 μg ml−1). However,
146 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

in induced vaginal candidiasis rat model (0.1, 1.0, 10% weight volume−1), the
response was not dose-dependent, and rats recovered after 12  days of treatment
(Chekem et al. 2010).
In osteoarthritis experimentally produced to Wistar rats, the crude hydroalco-
holic extract of leaves produced a reduction of synovial inflammation and pain with
ascaridol probably acting as antagonist of the N-methyl-D-aspartate (NMDA)
receptor (Calado et al. 2015). The ethanolic extract of aerial parts exhibited anti-­
inflammatory effects in different inflammation animal models with extended effects
both to vascular and cellular events (inhibition of myeloperoxidase, decrease of
adenosine-deaminase, reduction of nitric oxide levels, inhibition of TNF-α in pleu-
ral fluid after carrageenan injection) of the inflammatory process (Trivellato Grassi
et al. 2013). A novel use of a gel made with lyophilized aqueous extract as a bone
graft substitute for fracture osseointegration  was reported. The treatment, which
was tested in rabbits, stimulates bone alkaline phosphatase activity during early
fracture healing in a time-dependent manner promoting early bone formation
(Pinheiro Neto et al. 2017).
The methanolic extract from lyophilized plant material revealed higher antioxi-
dant activity and antitumor effect against colon (HCT-15), cervical (HeLa), and
hepatocellular carcinoma (HepG2) cell lines (Barros et al. 2013; Ruffa et al. 2002).
The intraperitoneal administration of hydroalcoholic extract (5  mg  kg−1) from
leaves produced Ehrlich tumor inhibition in Swiss mice (Nascimento et al. 2006).
The hydrodistilled essential oils from leaves had antiproliferative activity against
A549 and MCF-7 lines (Shameem et  al. 2019) and cytotoxic activity
(IC50 = 1.0 μg ml−1) against Burkitt’s lymphoma (RAJI) cells. The dichlorometh-
ane fraction (IC50 34.0 μg ml−1) and ethanol extract (IC50 47.0 μg ml−1) were active
against myeloid leukemia (K562) cells (Degenhardt et al. 2016).
Ascaridol, which is present in essential oils from seeds and leaves, is consid-
ered to be responsible of the anthelmintic, antiplasmodial, insecticide, pain-reliev-
ing, sedative, and antifungal activities (Smillie and Pessoa 1924; Johnson and
Croteau 1984; MacDonald et al. 2004; Cafferata et al. 2005; Potawale et al. 2008;
Patel 2017). Ascaridol was also active on tumor cell lines such as human promy-
elocytic leukemia cells (HL60), human leukemia cell lines (CCRF-CEM), and
breast cancer cell line (MDA-MB-231) (Efferth et al. 2002; Patel 2017). As for the
anti-­inflammatory activity, it was attributed to ascaridol and p-cymene (Degenhardt
et al. 2016). There are numerous studies related to its application as biopesticide.
The aqueous extract (2 and 20%, 24 h of treatment) was effective to reduce the
population of the phytonematode Pratylenchus brachyurus; however, the concen-
tration resulted phytotoxic to the parasited soybean plants (Mello et  al. 2006).
Alcoholic extracts (5%) of leaves reduced the number of Tuta absoluta
(Lepidoptera: Trichogrammatideae) after 24 h of spraying tomato plants (Barbosa
et  al. 2011). The effect was attributed to flavonoids and terpenoids (Cruz et  al.
2007). Also, a 1.0 and 2.0% (P/P) extract had insecticide effect on Sitophilus zea-
mais Mots. (Coleoptera: Curculionidae) (Silva et al. 2005). When the EO (0.1–
0.5%) and aqueous extracts (5.0%) were incorporated into the diet of the cabbage
pest Copitarsia decolorata Guenée 1852 (Lepidoptera: Noctuidae) larvae, the
6.3  Chenopodium ambrosioides L. (Chenopodiaceae) 147

essential oils reduced larval weight (33%), increased the larval period (19–20%),
and reduced fecundity (88%) and fertility (93%). Also, 0.5% essential oil reduced
the mean survival time (53%). Aqueous extracts only reduced fecundity (70%)
and fertility (75%) (Barbosa et al. 2011; Vázquez-Covarrubias et al. 2015). The
EO (0.36 μl ml−1) resulted in 100% repellency and mortality of Callosobruchus
chinensis L. and C. maculatus F. on stored pigeon pea seeds (LD50 2.8  μl and
2.5 μl, respectively) (Pandey et al. 2014). The 5% ethanolic extracts eliminated
54% of C. maculatus (Fabricus) adult and reduced oviposition rate (72%)
(Kemabonta and Okogbue 2002); the 10% aqueous extract reduced oviposition
(98%) of Plutella xylostella (L) (Lepidoptera: Plutellidae) (Medeiros et al. 2005).
The insecticide activity from EOs varied according to the P. xylostella develop-
mental stage and the treatment method (Wei et  al. 2015). It also eliminated the
second instar larvae of Anticarsia gemmatalis (Hübner) (Barbieri and Fiuza 2004).
Similarly, EOs from leaves influenced the behavior of Diabrotica speciosa
(Coleoptera: Chrysomelidae) reducing the feeding activity causing its death (de
Andrade Santiago et  al. 2014). The EOs (16.75% in oil dispersion) sprayed on
leaves of ornamental crops (Viola x wittrockiana var. Lubega F1 Mix and Hebe
‘Purple Pixie’) showed a slow rate but significant reduction of the aphids Myzus
persicae and Aphis gossypii (Smith et al. 2018).
As for mosquito, it was effective against Culex pipiens larvae (EC50 = 0.750 ppm),
C. pipiens (EC50 3.097 ppm) (Harraz et al. 2014), and Anopheles gambiae Giles
(Diptera: Culicidae) larvae (a vector of malaria), with a 100% of mortality (at 200
and 300 ppm) and a 100% of repellent effect (200 ppm) against adult mosquitoes
(Bigoga et al. 2013).
C. ambrosioides infusion and ethanolic extract from aerial parts prevented micro-
bial development and lipid oxidation of raw food (for its flavonoids and citric acid
content), particularly raw ground pork maintained at 4 °C. As that could increase
shelf-life of raw ground meat it was proposed as a natural antioxidant for lipids and
myoglobin (Villalobos-Delgado et al. 2017).
It was reported an allelopathic activity of the EO from leaves against the weed
Avena fatua L. (Sangeeta et al. 2017). Moreover, crude aqueous extract from aerial
parts (1 g 100 ml−1 roots and 1.5 g 100 ml−1 leaves) inhibited hypocotyl growth of
Amaranthus hypochondriacus. Methanolic extracts from roots and leaves (0.552–
0.509 μl per dish) inhibited hypocotyl growth (55%) and germination of A. hypo-
chondriacus. Also, the EO (0.15 ml l−1) was detrimental to plant growth (root dying,
inhibition of root growth, smallest and greenest leaves) of Phaseolus acutifolius
after 12 h of treatment. After 30 d of treatment with the lowest concentration, bio-
mass was 30% less than the control while biomass loss was more than 50% at higher
concentrations (Jiménez Osornio et al. 1996).
Toxicity  The extracts from aerial parts showed activity against Artemia salina Lech
(Sousa et al. 2012). The infusion was safer than the EO in healthy individuals; how-
ever, it was presumed that it could potentiate hepatic and renal disorders (MacDonald
et al. 2004; da Silva et al. 2014). The subchronic treatment of Swiss mice with the
hydroalcoholic extract (5 mg kg−1) and of NIH Swiss mice with the infusion (32, 64
and 134 mg ml−1) was not lethal and did not induce toxic alterations at therapeutic
148 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

doses (Pereira et al. 2010; Moreno Mendoza et al. 2013). The EO at high doses was
toxic (Montoya-Cabrera et al. 1996; da Silva et al. 2014). The in vitro exposure of
human lymphocytes to C. ambrosioides extracts increased the frequency of chromo-
somal aberrations and reduced the mitotic index (Gadano et al. 2002, 2007). Signs
of toxicity were evident only in the animals treated by intraperitoneal route (Monzote
et al. 2007). Ascaridol, which constitutes more than 50% of the EO, produced hypo-
thermia and decreased locomotor activity in mice and was lethal at high doses (Patel
2017). In Egyptian toad (Bufo regularis), the EO administrated for 3 months pro-
duced hepatocellular carcinomas (el-Mofty et al. 1992).
Carvacrol, caryophyllene oxide, and ascaridol inhibited the mitochondrial elec-
tron transport chain. Carvacrol and caryophyllene oxide affected mitochondrial
electron transport by directly inhibiting the electron-transferring complex I, whereas
the effects of ascaridol depended strongly on the availability of redox active Fe2+
(Monzote et al. 2009).
As for genotoxicity, when human lymphocyte cell cultures where treated with
different decoction concentrations, the chromosomal aberrations and sister chroma-
tid exchange increased, and the mitotic index decreased (Gadano et al. 2002).
Legal status  The EO was considered officinal by the FNA sixth edition. According
to the National Institute of Drugs (INAME-ANMAT), it is a toxic species (Flores
and Pellegrini 2015). The EO of C. ambrosioides is included in the United States
National Formulary and in the British Pharmacopoeia.

6.4  A
 nemia tomentosa (Savigny) Sw. var. anthriscifolia
(Schrad.) Mickel (Anemiaceae)

Synonymy  Anemia fulva auct. non (Cav.) Sw., Anemia anthriscifolia Schrad.,
Anemia ferruginea Kunth var. anthriscifolia, Hemianemia anthriscifolia (Schrad.)
C. F. Reed.

Vernacular name  Doradilla.

General information  The family Anemiaceae comprises about 100 species of

monophyletic ferns included in only one genus, Anemia. It is an aromatic fern with
a wide distribution, from the Atlantic coast of Brazil to Bolivia and Argentina. In
Argentina it could be found in the provinces of BAI, CAT, CHA, COR, COS, FOR,
JUJ, MIS, SAL, SDE, SFE, and TUC.

Botanical characteristics  A. tomentosa is a perennial species up to 70 cm long

with creeping horizontal rhizomes, 3–7 mm diameter with 1–7 mm orange long
hairs. It has frond dimorphism; the fertile frond is longer than the sterile one and
has pilose stipes brown at the base. Fronds are deltoid to ovo-deltoid blades,
bipinnate-­pinnatifid, with 8–13 pairs of chartaceous o subcoriaceous pinna and
6.4  Anemia tomentosa (Savigny) Sw. var. anthriscifolia (Schrad.) Mickel (Anemiaceae) 149

multicellular trichomes, thin tomentose rachis, and free venation. Sporangia are
distributed in two rows on the last segments, with trilete spores, ornamented exo-
sporium, and thin perisporium (Mickel 1962; de la Sota and Mickel 1968; Martínez
et al. 2003; Di Pasquo et al. 2016).

Ethnobotanical information  It is used to improve blood circulation (Martínez

Crovetto 1981a, b). The leaf is used as digestive, antitussive, antigripal, and expec-
torant (Martínez 2005; Barboza et al. 2009; Scarpa and Cassá 2015; Trillo 2016).

Chemical data  Aerial parts contain EOs with spathulenol, caryophyllene oxide,
α-bisabolol, α-bisaboloxide, 14-hydroxy-9-epi-(E) caryophyllene, neral, geranial,
α-pinene, camphene, 6-methyl-5-hepten-2-one,1,8-cineole, pinocarveol, isoafrica-
nol, and triquinane sesquiterpenes such as epi-presilphiperfolan-1-ol and
silphiperfol-­6-ene (Figs. 6.7 and 6.8) (Santos et al. 2003; Juliani et al. 2004; Santos
et al. 2006; Barboza et al. 2009; Pinto et al. 2009; Joseph-Nathan et al. 2010).

Biological activities  The antimicrobial activity against Mycobacterium tuberculo-

sis H37Rv (ATCC-27294) of the EO (200 μg ml−1) (MIC 100 μg ml−1) was mainly
attributed to triquinane sesquiterpenes (Pinto et al. 2009). The EO of the aerial parts
also exhibited activity against Bacillus cereus (18 mm inhibition zone) (Demo et al.
2005). The repellent activity against Aedes aegypti at a 90% essential oil concentra-
tion was attributed to monoterpenoids and sesquiterpenes (Gillij et al. 2008).

In vitro culture  Spore-derived gametophytes developed into sporophytes in MS

(Murashige and Skoog 1962) culture media and ½ MS culture media without plant
growth regulators. The in  vitro sporophytes showed an inverted volatile profile
regarding the wild-grown plants. In wild-grown plants, sesquiterpenes represented
97.5% of the volatiles produced, whereas in in vitro plants, the monoterpene profile
prevailed (69.8–89.8%) over sesquiterpenes (9.4–28.7%). The main monoterpene
identified in those cultures was α-pinene followed by trans-pinocarveol,
­pinocarvone, and myrtenyl acetate. The trinquinane sesquiterpenes found were
silphiperfol-­6-­ene followed by α-guaiene, β-barbatene, and 9-epipresilphiperfolan-
1-ol. The plant growth regulator IAA was not adequate to promote biomass produc-
tion while jasmonic acid promoted gametophyte and sporophyte development.

Fig. 6.7  Some triquinane sesquiterpenes found in Anemia tomentosa (Savigny) Sw. var. anthrisci-
folia (Schrad.) Mickel
150 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

Fig. 6.8  Chemical compounds found in Anemia tomentosa (Savigny) Sw. var. anthriscifolia
(Schrad.) Mickel essential oils

None of the plant growth regulators were able to reproduce the monoterpene/ses-
quiterpene balance of wild-grown plants a phenomenon that it was found related to
the different irradiance levels of in vitro cultures, to the necessity of differentiated
plant tissues at a specific moment, or to the development phase of the plant (Pinto
et al. 2013; Velasco-Castilho et al. 2018).

6.5  Plantago australis Lam. subsp. Australis (Plantaginaceae)

Vernacular name  llantén, siete varas, lengua de vaca, llantén blanco.

Synonymy  Plantago macropus Pilg., Plantago denudata Pilg., Plantago hirtella

Kunth var. platensis, Plantago candollei Rapin, Plantago macrostachys Decne.,
Plantago myosuros Lam. var. latifolia, Plantago accrescens Pilg., Plantago kurtzii
Pilg., Plantago refracta Pilg., Plantago stuckertii Pilg., Plantago macrostachys
6.5  Plantago australis Lam. subsp. Australis (Plantaginaceae) 151

Decne. var. angustifolia, Plantago macrostachys Decne. var. denudata, Plantago

macrostachys Decne. var. platensis, Plantago macrostachys Decne. var. stuckertii,
Plantago macrostachys Decne. var. catamarcensis, Plantago macrostachys Decne.
var. typica, Plantago macrostachys Decne. var. brachypus, Plantago stuckertii Pilg.
var. catamarcensis, Plantago hirtella Kunth var. glabrescens, Plantago macro-
stachys Decne. f. brachypus, Plantago australis Lam. ssp. macrostachys (Decne.)
Rahn, Plantago australis Lam. ssp. angustifolia (Pilg.) Rahn, Plantago bicallosa
Decne. var. angustifolia, Plantago macrostachys Decne. var. accrescens, Plantago
gigantea Decne., Plantago macrostachys Decne. var. gigantea, Plantago durvillei
Fisch. & C.A. Mey.

General information  Plantago australis is a popular plant widely spread in Latin

America. In Argentina it grows in the provinces of BAI, CAT, CHU, COR, COS,
ERI, JUJ, LPA, LRI, MEN, NEU, RNE, SAL, SCR, SFE, SJU, SLU, and TUC.

Botanical characteristics  P. australis is a perennial herb with an ephemeral main

root that is replaced by numerous secondary roots. Leaves are simple, alternate or
whorled, rarely opposite, elliptic-lanceolate, without stipules. The blade margin is
entire or toothed. Primary veins are prominent to the hipophile; the petiole is long.
Flowers are actinomorphe, small, anemophilous, generally in spikes, rarely soli-
tary, in the axil of each bract. Perianth has four sepals, a gamopetal corolla, with
four imbedding lobules. Androecium has four stamens, with versatile introrse
anthers. Ovary is superior, bicarpelar, uni- or bilocular with uni-to-pluri-ovules;
style is short with a long filiform stigma. Fruit is a pyxidium (pyxis). Seeds, 3–4
for capsule, are oblong and albuminous (Rahn 1996; Bayón et al. 2000; Moroni
et al. 2018).

Ethnomedical information  P. australis is used in folk medicine as pectoral,

anti-­catarrhal, emollient, analgesic, anti-inflammatory, for postpartum, bladder
and kidney diseases, for cystitis, wounds, and vaginal discharge, to healing
wounds and to treat contaminated (thick or bad) blood. The root is used in case of
infections, hernia, and prolapse. Leaves are taken as astringent, vulnerary, ablu-
ent, to facilitate birth, and the seeds, as laxative. It is also referred its use to treat
flu and common cold by Polish immigrants from the province of MIS
(Toursarkissian 1980; Bayón et  al. 2000; Barboza et  al. 2009; Gheno-Heredia
et al. 2011; Kujawska and Pieroni 2015).

Chemical data  It has aucubin (iridoid monoterpene), verbascoside, isoverbasco-

side, salidroside, plantamajoside, baicalein, hispidulin, ursolic acid, and oleanolic
acid (Ronsted et al. 2000; Barboza et al. 2009; Flores et al. 2016a, b). Verbascoside
or acetoside is a phenolic glycoside compound with four chemical groups: caffeic
acid, 4,5-hydroxyphenylethanol linked to β-(D)-glucopyranoside, rhamnose, and
glucose (Korkina et  al. 2007), that is the main active component from extracts
(Fig. 6.9).
152 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

Fig. 6.9  Chemical compounds from Plantago australis Lam. subsp. Australis (Plantaginaceae)

Biological activities  The antioxidant, anti-inflammatory, healing, and anti-­

nociceptive properties were attributed to verbascoside (Lee et al. 2007; Hausmann
et al. 2007; Vertuani et al. 2011; Yoou et al. 2015; de Moura Sperotto et al. 2018). The
dried fruit, dried root, leaf, and fruits have antiulcer and anti-inflammatory activity
(Barboza et al. 2009). In vitro treatment of keratinocyte cells HaCaT cells (scratch
test) with several concentrations of the hydroalcoholic extract (standardized on
­verbascoside) or verbascoside for 24 h promoted migration of keratinocytes at all
6.5  Plantago australis Lam. subsp. Australis (Plantaginaceae) 153

concentrations tested. On the other hand, in vivo tests administrating different con-
centrations of the hydroalcoholic extract to male Wistar rats and with a LPS inflam-
mation model in N9 microglial cells showed wound healing promotion. The extract
decreased viability at 10 times higher concentrations (1000 μg kg−1) and promotes a
faster wound healing than the verbascoside. On the other hand, both the extract and
verbascoside reduced the oxidative parameters (SOD, CAT in N9 cells LPS-activated)
at the concentrations tested (de Moura Sperotto et al. 2018). Previously, the positive
cicatrizing effect of the leaf extract on induced wounds in mouse (Mus musculus)
was attributed to flavonoids and tannins (Asto-Guamán et al. 2017) or to prostaglan-
din synthesis inhibition (Palmeiro et al. 2002).
The analgesic and anti-inflammatory activity of the hydroalcoholic extract (70%)
from leaves, roots, and fruits from P. australis growing in Southern Brazil were
tested on carrageenan-induced rat hind paw edema (250, 500, and 1000 mg kg−1)
and 1.2% acetic acid (10 mg kg−1)- induced writhing. The oral administration inhib-
ited both inflammation and writhes significantly (Palmeiro et al. 2003). The crude
ethanolic extract (500, 1000 mg kg−1) obtained from leaves was tested against etha-
nol-, indomethacin-, and cold restrain-induced stress ulcers showing a reduction of
the lesion and ulcer indexes produced by ethanol and an increase of mucous at the
highest concentration tested in cold restrain-induced stress ulcer. In this case the
responsible mechanism proposed was an interference with histamine and leukotri-
ene C4 deleterious effect on the gastric mucosa (Bürger et al. 2002).
The in vivo anti-inflammatory activity on oral lesions of a pharmaceutical formu-
lation (solution or cream) containing an ethanolic extract (10%) of P. australis
showed that the topical use reduced signs and symptoms of recurrent aphthous sto-
matitis, lichen planus, and actinic cheilitis (Flores et al. 2016a). Also, the effect on
the growth of Candida albicans on resilient denture liners was verified (Pereira-­
Cenci et al. 2014).
Toxicity studies  In vitro cytotoxicity of different concentrations of leaf crude
extract was reported. The hydroethanolic extract on in vitro models showed toxic-
ity in the Salmonella/microsome assay. No cytotoxicity nor mutagenicity was
detected at the concentrations tested; however, when the toxicity was studied in
V79 cells by the dimethyl thiazolyl diphenyl tetrazolium salt test (MTT) and neu-
tral red uptake (NRU) assay, both showed cytotoxicity at the highest concentra-
tions (1000  μg  ml−1 for the hydroalcoholic extract and 50  mg  ml−1 for the
verbascoside). As for in vivo assays, acute oral toxicity and subchronic tests made
on healthy female and male Wistar rats did not detect clinical signs of toxicity.
Behavior, body weight, histopathological analysis, and biochemical parameters
did not show significant differences respect to the controls (Henn et  al. 2019).
Similar results were obtained with P. australis aqueous extract; however, in this
last case, a significant increase of aspartate aminotransferase, alanine aminotrans-
ferase, and creatinine serum levels was found suggesting a hepatic function altera-
tion (Palmeiro et al. 2003). On the other hand, when the MTT was tested on a 3 T3
mouse fibroblast cell line with serial dilutions (10−1, 10−2, 10−3, 10−4, and 10−5) of
the 70% ethanol extract, the stronger cytotoxic effect was obtained as the concen-
tration increased (Flores et al. 2016b).
154 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

As it was proposed the use of the extract in phytocosmetics, phototoxicity was

also tested. The neutral red uptake (NRU) test in 3T3 did not show signs of verbas-
coside phototoxicity at the concentrations tested (1.0, 2.5, 5.0, 10, 25, 50, and
100 μg ml−1) (Henn et al. 2019).
As for genotoxicity and mutagenicity, neither the hydroethanolic extract nor the
verbascoside generated DNA-strand breaks or alterations in micronucleus, bud fre-
quency, or polychromatic erythrocytes/normochromic erythrocytes relationship at
the tested concentrations (Henn et al. 2019).

6.6  Alternanthera pungens Kunth (Amaranthaceae)

Synonymy  Achyranthes lorentzii (Uline) Standl., Achyranthes mucronata Lam.,

nom. illeg., Achyranthes radicans Cav., nom. illeg., Achyranthes repens L.,
Alternanthera achyrantha (L.) R. Br. ex Sweet, nom. illeg., Alternanthera achy-
rantha (L.) R. Br. ex Sweet var. leiantha, Alternanthera echinata Sm., Alternanthera
lorentzii Uline, Alternanthera pungens Kunth var. leiantha, Alternanthera pun-
gens Kunth f. pauciflora, Alternanthera repens (L.) Link, comb. superfl., Celosia
echinata Willd. ex Schult., Desmochaeta sordida Bunbury, Guilleminea procum-
bens Rojas Acosta, Illecebrum achyranthum L., nom. illeg., Illecebrum pungens
(Kunth) Spreng., Pityranthus crassifolius Mart., Pupalia sordida (Bunbury) Moq.,
Telanthera pungens (Kunth) Moq., Achyranthes leiantha (Seub.) Standl.,
Paronychia achyrantha (L.) Desf., (Amorin and Rosow 1988).

Vernacular name  Yerba de pollo, guambarusa, raíz colorada, lagunilla, hierba del
pollo, llapá reliñ, achawál kachú (pasto del pollo), ierwé achawál (hierba del pollo)
in the Araucanian-Pampa language; taasó, taa-só in toba language; bashé um-­
patpatelét (espinita pegadita a la tierra) in vilela language; caá-pé, quisca-yuyu,
torito, sanguinaria, colchón de perro, yerba de carretero (Cuba), adorna jardín
(Puerto Rico), caroca, carauca, sangradera (Venezuela), yerba del moro hembra
(Perú), khakiweed, creeping chaffweed, spingflower alternanthera (English).

General information  Native to South America, A. pungens is characterized by its

pungent easily shedding tepals that become harder at maturity. It grows in tropical,
subtropical, semiarid, and warmer temperate environments, and also in disturbed
sites, bare areas, roadsides, parks, lawns, waste areas, watercourses, turf grasses,
orchards, and occasionally in native pastures and grasslands. It was naturalized in
Australia, northern Africa, Asia (China, Bhutan, Myanmar, Thailand, and Papua
New Guinea), and Pacific islands (Hawaii and New Caledonia) (Iamonico and
Sánchez del Pino 2016; Peña Rivera and Ferrer-Gallego 2016). In Argentina it usu-
ally grows as a dense mat of prickly vegetation in the provinces of BAI, CHA, COS,
ERI, FOR, JUJ, SAL, SDE, SFE, SJU, and TUC (Carrizo and Isasmendi 1998;
Barboza et al. 2009). It has nutritional value (Raju et al. 2004; Hundiwale Jogendra
et al. 2012). Flowering is in summer and autumn (Arenas 1981). It propagates by
seed and vegetatively.
6.6  Alternanthera pungens Kunth (Amaranthaceae) 155

Botanical characteristics  A. pungens is a small, perennial, creeping, pubescent

herbaceous plant with zigzag stems up to 60 cm long that springs from a tough and
thickened central crown. Stem is hirsute (abundant little short and soft hairs in the
nodes). Leaves are shortly petiolate, opposite, slightly asymmetric, egg-shaped or
elliptic with a rounded tip, and an entire or slightly waved margin. Leaves are usu-
ally glabrous in the main vain, with few hairs on the veins of the underside, which
are prominent in the hipophile. Greenish or greenish-yellow flowers are clustered as
axillary spikes (8–12 mm long), lanceolate or ovate oblong bracts, with 5 tepals, the
two external larger, with 5 stamens, a globular ovary with stigma. The fruit is a
utricle, shiny orange or yellowish lenticular seeds usually remained hidden inside
the old, prickly, flower parts (Filippa and Espinar 1993). The seeds are contained
inside a burr that is transported attached to animals, clothing, and other objects (e.g.,
vehicle tires). They may also be dispersed by water, in contaminated agricultural
products (e.g., fodder and pasture seed). The stem fragments can be spread by
machinery, livestock, or cultivation (Owa et al. 2016).

Ethnomedical information  A. pungens is used in case of measles, against flu,

headache, and abdominal pain. A spoonful of the decoction, made with the inflores-
cences, is taken as vermifuge before breakfast once a day (Hundiwale Jogendra
et  al. 2012). Root and leaves have activity as diuretic, depurative, digestive, and
hepatic. The infusion made with leaves and stems is active in cases of viral diseases,
hepatitis, epidemic parotitis, hemorrhagic fever, and influenza. There are also refer-
ences about its properties to treat gastric, liver, and intestinal problems (Domínguez
1928; Saggese 1959; Soraru and Bandoni 1978; Ratera and Ratera 1980;
Toursarkissian 1980; Hieronymus 1882; Amorín 1988; Boelcke 1989; Roig 1992).
In Argentina, infusions and decoctions made with roots, leaves, and flowering tips
are commonly taken to cure empacho and indigestion. The preparations must be
handled with care when administered to children because the diuretic activity could
cause dehydration. It has depurative mildly laxative and diuretic properties, regular-
izing stomach and intestinal functions (Orfila 1971; Martínez Crovetto 1981a, b;
Amorín 1988). It is also used in Ayurveda medicine (Kalpana et al. 2018).

Chemical Data  The chemical compounds found in the entire plant are choline,
oleanolic acid, and β-spinaestrol (de Ruiz et al. 1991; Rondina and Coussio 1969,
1981; Dogra et al. 1977) and in the flowers, azulene, α-curcumene, bornyl acetate,
α-borneol, camphene, camphor, 1,8-cineol, and α-terpineol (Fig. 6.10). In the dried
fruit, it was identified oleanolic acid and rutin (de Ruiz et al. 1991; Barboza et al.
2009). As for the carotenoids, it was found violaxanthin, lutein, zeaxanthin, and
β-carotene (Raju et al. 2007).

Biological activities  The dried aerial parts showed antitumor, antiviral (HIV and
anti-Hantaan) activity, and reverse transcriptase inhibition (Barboza et al. 2009).
It was also cited that the leaf extracts have immunological reactivity (Gayathri
et al. 2001).
156 6  Puna: A Surrealistic Landscape in the Argentinean Highlands

H3C CH3

OH
CH3 CH3
OH O
+ CH3
N
H3C
H3C CH3

choline oleanolic acid azulene

CH3
H3C

H3C
CH3
O

O HO

α-curcumene bornyl acetate borneol

camphene camphor

Fig. 6.10  Chemical compounds found in Althernanthera pungens Kunth (Amaranthaceae)

The antimicrobial activity from different crude extracts was analyzed against
several bacteria and fungi. The crude plant extracts (200, 100, 50, and 25 mg ml−1)
from aerial parts showed antibacterial activity against Klebsiella pneumoniae
MTCC NO 109 (ethanolic extract), Mycobacterium smegmatis MTCC NO 992
(acetone and ethanolic extract), Bacillus subtilis MTCC NO 2057 (aqueous extract),
and Chromobacterium violaceum MTCC NO 2656 (acetone extract) with inhibition
zones >10 mm. Antifungal activity was shown only against Aspergillus fumigatus
MTCC NO 3002 (acetone and aqueous extracts) (Jakhar and Dahiya 2017).
The ethanolic extract exhibited highest antioxidant activity (DPPH assay) (IC50
100.79  mg  ml−1) than the acetone (IC50 203.56  mg  ml−1), aqueous (IC50
324.43 mg ml−1), and petroleum ether extracts (IC50 931.63 mg ml−1), which cor-
relates with the total phenolic and flavonoid content of each extract (Jakhar and
Dahiya 2017).
References 157

The administration of a tea made with dried leaves in boiled water (1 g 40 ml−1)
to patients with HIV each 2 days during 12 months decreased the plasmatic levels
of oxidative stress biomarkers (AOPP and MDA) and increased those of T CD4
and CD8 lymphocytes without showing signs of hepatic or renal toxicity (Djohan
et al. 2009).
The ethanolic extract from the entire plant (50–300 mg kg−1) reduced glycemia
and improved serum lipid profile (lowered cholesterol and triglyceride, restored
HDL levels) in Wistar rats with alloxan-induced diabetes after 21 days of treatment,
being 50 mg kg−1 the most effective dose (Owa et al. 2016).
The ethanolic extract (2.8%) from aerial parts of flowering plants produced sig-
nificant increases of urinary volumetric excretion and sodium urinary excretion in
Wistar rats (100 mg kg−1 p.o.) as did furosemide (Calderón et al. 1997).
Methanolic extracts of the whole plants tested on Swiss albino mice by the tail
immersion method showed analgesic activity, showing the chloroform fraction the
most potent analgesia. The chloroform fraction was rich in terpenoids, steroids,
flavonoids, and phenolic compounds (Kalpana et al. 2018).
The aqueous, methanol, and n-hexane extracts (10, 20, 30 mg ml−1) were tested
against Sitophilus oryzae, Callosobuchus chinensis, and Tribolium castaneum. The
hexane (30 mg ml−1) and methanol (10 mg ml−1) extracts resulted effective against
S. oryzae (100% mortality after 2 and 7 days, respectively). The aqueous (30 mg ml−1)
and hexane (10 mg ml−1) extracts were effective against C. chinensis (100% mortal-
ity after 6 and 8 days of treatment, respectively). The aqueous (30 mg ml−1) and the
hexane (10 mg ml−1) extracts were effective against T. castaneum (100% mortality
after 10 and 15 days (LD50 < 10 mg ml−1after 24 hours) (Kazmi et al. 2017).
Toxicity  A. pungens burrs can contaminate lucerne hay and other stock feeds and
could cause digestive disturbances and skin ailment in cattle. Horses that graze on
areas containing large amounts of this species have developed a form of staggers.
The burrs can also contribute to vegetable fault in wool (Weeds of Australia 2016).

Legal status  This species is inscribed in the FNA sixth edition.

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Chapter 7
Yungas

7.1  Introduction: Yungas, the Green Face of the Andes

The Yungas are mountain jungles of Northwest Argentina that have just been
included by UNESCO in the World Network of Biosphere Reserves (2002). They
have different characteristics according to the altitude, with a dense and humid for-
est vegetation and high biodiversity in the lower regions and predominantly shrubs
and grasslands in the highlands (Fig. 7.1). The climate is warm and humid to subhu-
mid. The conditions of temperature and humidity vary according to the altitude, lati-
tude, and exposure of the slopes. The mountain ranges form a barrier that condenses
the humid winds from the South Atlantic anticyclone creating the conditions for a
thick wooded development. Rains are mainly during summer with annual precipita-
tion oscillating between 900 and 1300 mm. During the colder months, the charac-
teristic mists from the cloudy jungles compensate the lack of rain. It has a
well-organized fluvial network, and the soils have abundant organic matter, with
frequent local landslides.
The fauna has typical species such as Amazona tucumana (loro alisero), Antigone
vipio (white-naped crane), Cinclus schulzi (mirlo de agua), Eriocnemis glaucopoi-
des (picaflor frente azul), Psittacidae spp. (macaws), Pyrrhura molinae (chiripepé
de la Yunga), and Synallaxis scutata (pijuí anaranjado). Among the mammals there
are the Anoura caudifer (snout bat), Cavia tschudii (cuis serrano), Dasyprocta
punctata (reddish agouti), Hippocamelus antisensis (taruca), and Scirus ignites (red
squirrel). Among the amphibians, there is the marsupial frog (Gastrotheca gracilis),
which is at risk of extinction and was included by the ONG Global Wildlife in the
initiative The Search for Lost Species. There are also Leopardos wiedii (gato sal-
vaje), Mazama americana (corzuela roja), Panthera onca (yaguareté), Puma con-
color concolor, Pecari tajacu, and Tapirus terrestres (tapir), among others.
The flora varies according to the altitude, and four regions could be distinguished:
Pedemontane jungle, hot and humid with predominance of epiphytes, bryophytes,

© Springer Nature Switzerland AG 2019 167

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_7
168 7 Yungas

Fig. 7.1  Yungas, province

of Jujuy, Argentina.
(Personal collection PG
Alvarez)

ferns, and arboreal species such as Calycophyllum multiflorum Griseb.,

Chrysophyllum marginatum (Hook. & Arn.) Radlk., Enterolobium contortisiliquum
(pacará), Handroanthus impetiginosus, Jacaranda mimosifolia, Parapiptadenia
excelsa (cebil), and Tipuana tipu, among others; Montana jungle, temperate-warm
and humid, where Blepharocalyx gigantea (horco-molle), Cinnamomum porphy-
rium (Griseb.) Kosterm., Enterolobium contortisiliquum, Juglans australis (nogal
criollo), Parapiptadenia excelsa, myrtles, and cedars, among others, predominate;
Montana forest, temperate and humid with frequent winter frosts, where alder,
Podocarpus parlatorei (pino del cerro), and walnut predominate; and temperate-­
cold and subhumid high-altitude meadows, with patches of montane forest alternat-
ing shrubland and, at a higher altitude, pure herbaceous communities (De la Sota
1972; Cabrera 1976; Morales et al. 1995; Brown et al. 2001; Grau 2006; Coll-­Aráoz
et al. 2016).
Among the species with medical use are Acacia macracantha Humb. & Bonpl.
Ex Willd (Fabaceae), Achyrocline alata (Kunth) DC (Asteraceae), Achyrocline flac-
cida (Weinm) DC (Asterace), Ageratum conyzoides L. (Asteraceae), Aloysia polys-
tachya (Griseb.) Mold. (Lamiaceae), Amaranthus quitensis Kunth. (Amaranthaceae),
Anredera cordifolia (Ten.) Steenis (Basellaceae), Clinopodium gilliesii (Benth.)
Kunze (Lamiaceae), Equisetum giganteum L. (Equisetaceae), Buddleja brasiliensis
7.2  L epidium didymum L. (Brassicaceae) 169

Jacq. Ex Spreng. (Buddlejaceae), Fagara coco (Gill.) Engler (Rutaceae),

Erythroxylum coca Lam. var. coca (Erythroxylaceae), Lepechinia vesiculosa
(Benth.) Epling. (Lamiaceae), Lepidium didymus (L.) Brassicaceae, Maytenus cuez-
zoi Legname (Celastraceae), Myroxylon peruiferum L. f. (Fabaceae), Pavonia
sepium St. Hil. (Malvaceae), Petiveria alliaceae L. (Phytolaccaceae), Pluchea sag-
ittalis (Lam.) Cabrera, (Asteraceae), Sambucus nigra subsp. peruviana (Kunth)
R.  Bolli (Adoxaceae), Satureja boliviana (Benth.) Briquet (Lamiaceae), Satureja
parvifolia (Phil.) Epling (Lamiaceae), Schinus molle L. (Anacardiaceae),
Senecio crepidifolius DC. (Asteraceae), Smallanthus macroscyphus (Baker) A. Grau
ex A.A.  Sáenz (Asteraceae), Smilax campestris Griseb. (Smilacaceae), Solanum
sisymbriifolium Lam.var. sisymbriifolium (Solanaceae), Tabebuia impetiginosa
(Mart. ex DC.) Standl. (Bignoniaceae), Tagetes campanulata Griseb. (Asteraceae),
Tillandsia usneoides (L.) L. (Bromeliaceae), Trichocereus arborícola Kimnach
(Cactaceae), Urera baccifera (L.) Gaudich. (Urticaceae), and Xanthium ambrosioi-
des  Hook. & Arn. (Asteraceae).

7.2  Lepidium didymum L. (Brassicaceae)

Synonymy  Coronopus leptocarpus Boelcke, Coronopus leptocarpus Boelcke var.

microcarpus, Coronopus didymus (L.) Sm., Coronopus didymus L. var. procum-
bens, Coronopus didymus L. var. macrocarpus, Senebiera didyma (L.) Pers.,
Senebiera pinnatifica DC., Lepicochlea americana Rojas Acosta.

Vernacular name  Mastuerzo, quimpe, quimpi, calachi, yerba del ciervo, mas-
truço, erva-de-santa-maria, mentruz, mentruz-rasteiro (Portuguese), swinecress
(English).

General information  L. didymium is a stinky weed that spontaneously grows in

parks, gardens, orchards, and pasture, preferring shady places. It is also considered
an edible species, being added to salads or to beverages. In Argentina it could be
found in the provinces of BAI, CAT, CHA, COR, COS, DFE, ERI, FOR, JUJ, LPA,
LRI, MEN, MIS, SAL, SCR, SDE, SFE, SJU, SLU, TDF, and TUC. It is also used
to feed cattle, mainly sheep. Blooming is in spring (Amorin and Rosow 1988;
Lorenzi and Matos 2002; Alonso and Desmarchelier 2015).

Botanical characteristics   L. didymum is an annual or biannual herb whose height

ranges between 10 and 20 cm. It has a procumbent stem with leafy, glabrous, or
pubescent branches. Leaves are basal and bipinnate with whole or barely dentate
lobes. The white flowers have glabrous sepals on the back side. Nectaries are
tongue-shaped. The fruit is an indehiscent, reticulate, bi-globose silicle, with
a rough surface and a seed in each compartment. Pedicels are cylindrical with scarce
antrorse hairs (Prina 1995; Ferreira Jorge et al. 1986).
170 7 Yungas

Ethnobotanical information  The tisane made with the whole plant is used as an
expectorant and digestive and to treat intermittent fever. The juice is used as vulner-
ary; to treat hemorrhoids, gangrene, as antitussive, antimalarial, antiscorbutic, for
hernia and prolapse, menstrual diseases, neonatal and pediatrics care, in cases of
gingivitis. It also cited its use in pregnancy and to facilitate birth and in cancerous
processes (Sorarú and Bandoni 1978; Ratera and Ratera 1980; Toursarkissian 1980;
Scarpa 2004; Barboza et al. 2009). The tisane made with the aerial parts is taken as
antiscorbutic, vulnerary, anti-inflammatory, antiseptic, hemostatic, astringent for
humoral and blood diseases and to prevent pyorrhea. Decoction of flowers is used
in cases of vaginal fungus (Barboza et al. 2009).

Chemical data  In the dried entire plant, there were identified benzyl cyanide, glu-
cotrapaeolin (benzyl isothiocyanate glycoside), chrysoeriol, chrysoeriol-6”(OAc)-
4’β-D-­glucoside, cymaroside, rhamnetin, β-sitosterol, sinapine, 1,8-dihydroxy
anthraquinone, and myristic acid. The EO from aerial parts contains benzyl cyanide,
benzylmercaptan, benzene, acetonitrile, benzyl disulfide, benzyltrisulfide, and
hydrocarbons of high molecular weight (Fig. 7.2). In the dried seed, there are ben-
zylnitrile and glucosinolates (Haque 1989; Ruiz et al. 1989, 1993; Prabhakar et al.
2002; Mantena et al. 2005; Barboza et al. 2009; Moismann Busnardo et al. 2010).

Biological activities  The aqueous extract of the whole plant (200–400 mg kg−1)

was reported as effective in cases of allergies, inflammation, wound healing, hepa-
totoxicity, and hyperglycemia (Mantena et al. 2005). The aqueous extract was more
active as anti-inflammatory and wound healer than the ethanolic extract (Prabhakar
et al. 2002). The hydroalcoholic extract (200, 400, 600 mg kg−1) from the whole
plant showed a dose-dependent decrease of paw inflammation (decrease of leuko-
cyte migration, inhibition of neutrophil influx, and exudation) on the model of pleu-
risy and paw edema induced by carrageenan in male Swiss mice. Similar results
were obtained on edema induced by other phlogogens (substance P, bradykinin, and
histamine) (Moismann Busnardo et al. 2010).
The aqueous extract (1 drop during 7 days) was effective to accelerate skin scars
of Wistar rats, showing an increase of fibroblasts and collagen fibers (Nitz et  al.
2006). The aqueous and ethanolic extracts (200 mg kg−1) showed similar activity
(Prabhakar et al. 2002).
The crude methanol extract had antimicrobial activity against Staphylococcus
aureus ATCC 25923, Staphylococcus epidermidis ATCC 12228, Bacillus subtilis
ATCC 6633 (Gram- positive), and Klebsiella pneumonia ATCC 13883 (Gram-­
negative). Against the  fungi Aspergillus niger ATCC 16404, Aspergillus flavus
ATCC 32611, and Alternaria solani ATCC 11078, the inhibitory activity was lower
(Uddin et al. 2014). On the other hand, the methanolic extract (10 mg ml−1) did not
show in vitro antiviral activity against herpes simplex virus type 1 and 2, bovine
viral diarrhea virus type 1, influenza virus type A, and human immunodeficiency
virus type 1 (HIV-1) (Ruffa et al. 2004). The biological activities were attributed to
the antioxidant activity of flavonoids and tannins (Mantena et al. 2005; Prabhakar
et al. 2006; Moismann Busnardo et al. 2010).
7.3  Clinopodium gilliesii (Benth.) Kunze (Lamiaceae) 171

OH O OH O

OH

HO O O O

OH
OH
N O
OH

benzyl cyanide chrysoeriol rhamnetin

+
N

O
O O

OH OH

sinapine myristic acid

Fig. 7.2  Chemical compounds from Lepidium didymum L.

Toxicity  The hydroethanolic extract (0.1–100 mg ml−1) showed moderate toxicity

against Eisenia andrei (DL50  =  3.498  mg  ml−1) and Artemia salina
(CE50 = 773.69 ppm) (Ponce et al. 2007).

7.3  Clinopodium gilliesii (Benth.) Kunze (Lamiaceae)

Synonymy  Satureja parvifolia (Phil.) Epling, Micromeria eugenioides (Griseb.)

Hieron., Xenopoma eugenioides Griseb., Bystropogon minutus Briq., Satureja euge-
nioides Griseb., Satureja oligantha Briq., Micromeria gilliesii Benth., Oreosphacus
parvifolia Phil., Satureja gilliesii (Benth.) Briq.

Vernacular name  Muña-muña, salvilori, muña, hillencoma, mulla-mulla, yerba

del pájaro, hierba del amor.
172 7 Yungas

General information  C. gilliesii grows in the mountainous regions, between 1200

and 4000 m of altitude, of the provinces of CAT, COR, JUJ, LRI, MEN, SAL, SJU,
and TUC. The aerial parts are used as a condiment and are also added to mate infu-
sion. The branches are used as fuel (Orfila and Farina 1997; Brodkiewicz et  al.
2017).

Botanical characteristics  C. gilliesii is an aromatic undershrub up to 2  m high,

with stiff, erect branches; quadrangular reddish-brown pubescent stems; and large
hairs, eventually glabrescent. Leaves are imbricate and subsessile or with petioles up
to 1  mm long; they are oblong or lanceolate, obtuse to rounded at the apex, and
broadly cuneate by the base, with an entire or slightly serrate margin. Both leaf sur-
faces are shortly pubescent, the lower surface paler and dotted with yellowish glands.
The inflorescence is a cyme, with 1–3 flowers. Flowers are solitary or pauciflorus,
shortly pedicelled. The shortly tubular-campanulate calyx is slightly pubescent, with
shortly curly hairs, teethed, ribbed, and glandular on the exterior, with a basal ring of
hairs on the interior. The white corolla is externally hairy, and anthers are subsessile.
The fruit is a trigonous, oblong-ovoid-shaped, glabrous nutlet (Wood 2011).

Ethnobotanical information  The aerial parts have antioxidant, stomachic, laxa-

tive, emmenagogue, and aphrodisiac activity. It is used in cases of hernia and pro-
lapse, menstrual diseases, postpartum pains, menopause, and against soroche and
headache (Ruthsatz 1974; Barboza et al. 2009; Dadé et al. 2009; Simoni and Perea
2016; Bektas and Cilkiz 2016).

Chemical data  The leaf  EOs contain pulegone, cis-dihydrocarvone, p-cymene,

1,8-cineol (eucalyptol), E-isocitral, carvacrol, carvacryl acetate, carvone, cis-­
dihydrocarvone, piperitone oxide, piperitenone, piperitenone oxide, lippione, dihy-
drolippione, linalool, isopulegol, α- and β-pinene, limonene, pulegone, sabinene,
menthol, menthone, neoisomenthol, and α-terpinene (Fig. 7.3) (Zygaldo et al. 1993;
Muschietti et al. 1996; Viturro et al. 2000; van Baren et al. 2006; Luna et al. 2008;
Barboza et al. 2009; Vázquez et al. 2016; Reynoso et al. 2018). The composition of
the EOs varies according to the chemotype, characteristics of soil, altitude, and
weather (Viturro et al. 2000).

Biological activities  The decoction yielded the highest concentration of phenolic

compounds and consequently the highest antioxidant activity, than the hydrolate
and the ethanolic extract (Cabana et al. 2013). The EOs obtained by hydrodistilla-
tion from the aerial parts have antioxidant activity, being pulegone (74.5%) and
piperitenone oxide the most abundant chemicals (Dadé et al. 2009; Cabana et al.
2013; Barbieri et al. 2015). The anti-inflammatory activity from the aqueous and
ethanolic extracts was tested in male Wistar rats through the carrageenan-induced
edema paw and cotton pellet-induced granuloma formation models. Rats pretreated
with the ethanol extract (500 mg kg−1) and the topical application (100 μl ml−1) of
the EO produced a significant reduction of edema after 2 h of treatment. Both treat-
ments reduced transudative and granuloma weights (Reynoso et al. 2018).
7.3  Clinopodium gilliesii (Benth.) Kunze (Lamiaceae) 173

Fig. 7.3  Chemical compounds from Clinopodium gilliesii (Benth.) Kunze

The hexane, dichloromethane, and methanol extracts were active against

Escherichia coli and Trichophyton mentagrophytes (MIC 1 mg ml−1) (Feresin et al.
2001). The total flavonoid extract (methanol 50% and 80%) from aerial parts tested
against Gram-negative rods (Escherichia coli ATCC 25922, Pseudomonas aerugi-
nosa ATCC27853, Klebsiella pneumoneae, Salmonella enteriditis, and Shigella
sp.), Gram-positive cocci (Staphylococcus aureus ATCC 6538, Staphylococcus, and
Streptococcus β-hemoliticus A Group), and Gram-positive spore-forming rods
(Bacillus subtilis BSGC A1) inhibited bacterial growth, except in the case of S.
epidermidis, Klebsiella, and Salmonella (Hernández et  al. 2000). Methanol and
dichloromethane extracts tested against Staphylococcus aureus F7 methicillin-­
resistant and P. aeruginosa ATCC 6738 growing as biofilms were inhibited by both
extracts (Brodkiewicz et al. 2017). The EO (24% w/w) obtained by hydrodistillation
of aerial parts (leaves and flowers) (0.25–2.00 mg ml−1) was active against Yersinia
enterocolítica-PI, Salmonella enteritidis-MI, Escherichia coli ATCC 25922, E. coli
174 7 Yungas

isolated from clinical strains, and Salmonella sp-LM. Moderate activity was found
against Staphylococcus aureus methicillin-sensitive and Staphylococcus aureus LM
(Luna et al. 2008). A reversible inhibition of spore germination was attained between
0.1% and 1.0% EO concentration for the fungi Alternaria solani (Ell. Et Mart.),
Sclerotium cepivorum Berk., and Colletotrichum coccodes (Wallr.) Hughes. At
1.0% EO concentration, mycelial growth of fungi was totally inhibited, but no effect
on mycelia was shown at 0.01% (Zygaldo and Grosso 1995).
The EOs have insect repellent, antifungal, and antibacterial activities (Zygaldo
et al. 1995). The in vitro effect of the EO (50% and 100%) and the corresponding
hydrolate (residue of hydrodistillation) showed activity against Streptococcus
mutans (ATCC 25175). After 72 h of exposition, the inhibition of growth was simi-
lar to that obtained with chlorhexidine glycolate (0.12%) with an increasing effect
with time (Rivadeneira-Cajas and Álvarez-Velasco 2015).
Insect repellant activity against Triatoma infestans (Klug) was also reported
(Lima et al. 2011). EOs obtained from leaves resulted active against permethrin-­
resistant head lice (50% knock down, 32.06 min after fumigation) being the most
effective EO from plant chemotypes with more piperitone content (Toloza et  al.
2010).
Methanol and water extracts of leaves showed significant activity against
Plasmodium falciparum K1 (IC50 3.0 and 8.5 μg ml−1, respectively) that was attrib-
uted to the content on eriodyctiol (IC 17.2 μg ml−1), luteolin (IC 6.4 μg ml−1), urso-
lic (IC 4.9 μg ml−1), and oleanolic acid (IC 9.3 μg ml−1) (Debenedetti et al. 2002).
That extracts also have activity against P. falciparum 3D7 strain and Trypanosoma
brucei rhodesiense (van Baren et al. 2006).
The aqueous extracts (100 mg ml−1) showed allelopathic activity (67% inhibition
of growth) on wheat (Mongelli et al. 1997).
Toxicity  The aqueous extract (100, 1000, and 2000 μg ml−1) was cytotoxic against
Artemia salina (Mongelli et al. 1995). Cytotoxicity of eriodictyol on mammalian
KB cell was relatively low (IC50 174.2 μg ml−1) (van Baren et al. 2006). The admin-
istration of a single dose (of 4000 and 8000 mg kg−1) of ethanolic extract of aerial
parts of C. gilliesii on shaved skin of male rats did not show death or toxic symp-
toms after 24–48 h of treatment (Reynoso et al. 2018).

7.4  Schinus molle L. Var. Molle (Anacardiaceae)

Synonymy  Schinus molle var. argentifolius Marchand, Schinus molle var. hyungan
angustifolium Sessé & Moc., Schinus biluminosus Salisb., Schinus molle var. huigan
(Molina) Marchand (Alonso and Desmarchelier 2015).

Common name  Aguaribay, árbol de la pimienta, curanguay, chichita sauce, molle,

pimentero, chichita péndula, falso pimentero, anacahuita, pirul, aroeira (Portuguese),
Califorina pepper tree (Eng.).
7.4  Schinus molle L. Var. Molle (Anacardiaceae) 175

General information  S. molle is an evergreen tree, very resistant to high tempera-

tures and drought, and native from Central and South America that is also world-
wide found, mainly in arid and saline soils. In Argentina, S. molle grows in the
provinces of BAI, COR, COS, ERI, JUJ, MIS, SAL, and SFE. Leaves are used as
dye, and its reddish fruits are edible having a taste similar to pepper; they are also
used to produce fermented beverages. It is also considered as an ornamental tree
(Goldstein and Coleman 2004; Albán-Castillo et al. 2018). It was proposed as an
indicator of heavy metal contamination (Al, As, Co, Cu, Cd, Cr, Ni, Pb, Ti, V, and
Zn) (Alcalá-Jáuregui et al. 2015).

Botanical characteristics  S. molle is a tree, 3–15 m height, with dark brown and
intensely fissured bark. The leaves are imparipinnate, persistent, alternate, and aro-
matic, with a winged rachis and 20–40 leaflets. Leaflets are linear-lanceolate and
entire or toothed by the margins. The fruit is a drupe, 4–6 mm in diameter, pink to
reddish, and bright (Rodríguez et al. 2005; Machado et al. 2018).

Ethnobotanical information  The decoction of the entire plant is used as anti-

pyretic, anti-inflammatory, emmenagogue, antifungal, antibacterial, antirheumatic,
and anti-conjunctivitis, to treat skin affections, ulcer, urinary tract diseases, and
respiratory diseases. The infusion made with the bark is taken as purgative and
stimulant. The decoction of the leaves is considered as analgesic, antitussive, anti-
depressant, anti-inflammatory, and hypotensive, for neonatal and pediatrics care and
for hernia and prolapse. Leaves and fruits are taken as emmenagogue, anti-­
inflammatory, and antiseptic. The fruit is used as diuretic, antirheumatic, and anti-­
inflammatory for amenorrhea and dysmenorrhea. The EO is prescribed as
antibacterial, antifungal, and cytotoxic, and for venereal diseases. The infusion
made with the oleoresin is considered antirheumatic, wound healing, and purgative
(Bernhard et al. 1983; Yueqin et al. 2003; Goldstein and Coleman 2004; Díaz et al.
2008; Barboza et al. 2009; Guerra-Boone et al. 2013; López et al. 2014; Martins
et al. 2014; Maema et al. 2016; Pellegrini et al. 2017).

Chemical data  S. molle contains the pentacyclic triterpenoid α-amyrin. The

EO  from leaves contains croweacin, limonene, phellandrene, carvacrol,
­cismen-2-en-­1-ol, trans-piperitol, and laccase. In the leaves there are isoquercitrin
6”-O-phydroxybenzoate, 2”-O-α-l-rhamnopyranosyl-hyperin-6“-O-gallate, gallic
acid, methyl gallate, chlorogenic acid, 2”-O-α-l-rhamnopyranosyl-hyperin, quer-
cetin 3-O-β-d-neohesperidoside, miquelianin, quercetin, rutin, quercetin 3-O-β-d
galacturonopyranoside, isoquercitrin, hyperin, isoquercitrin 6“-gallate, hyperin
6”-O-gallate, and (+) catechin. In the EO from fruits, there are α-bergamont-trans,
α-cadinol, γ-cadinene, bourbonene, α- and β-phellandrene, β-spathulene,
D-limonene, silvestrene, α-pinene, β-pinene, carvacrol, myrcene, camphene,
o-­ethylphenol, p-cymene, p-cymol, γ-cubebene, α- and γ-muurolene (Fig. 7.4). In
the fruits there are the triterpenoid isomasticadienonic, isomasticadienonalic, mas-
ticadienonic, and epi-isomasticadienolic acids and the flavonoid chamaejasmin
176 7 Yungas

OH

HO O
OH

O
O
OH OO OH
O
O OH
HO
O
OH OH

α-amyrin croweacin miquelianin

HO

α-cadinol γ-cadinene α-muurolene γ-muurolene

OH

OH O

OH
O
OH O

OH
O
HO O
O
OH

chamaejasmin isso-masticadienoic acid bourbonene

Fig. 7.4  Chemical compounds from Schinus molle L. var. molle

(Fig. 7.4) (Díaz et al. 2008; Barboza et al. 2009; Deveci et al. 2010; González et al.
2011; Simionatto et  al. 2011; de Mendonça Rocha et  al. 2012; Caballero et  al.
2014; Doleski Muhd et al. 2015; Dos Santos et al. 2015; Malca-García et al. 2017;
Machado et al. 2018).

Biological activities  Light, high temperature, and high pressure produced a

decrease of the biological activity of the EOs due to degradation or volatilization
(Beyki et al. 2014; de Oliveira et al. 2014).
The hexane extract from leaves showed antidepressant-like activity in the tail
suspension test in mice, a predictive model of depression. The extract (600–
1000 mg kg−1) significantly reduced the locomotor activity of mice in the open-field
test. A similar effect was produced by the flavonoid rutin (0.3, 1.0, and 3.0 mg kg−1)
isolated from the ethanolic extract (Machado et al. 2007; Machado et al. 2008).
7.4  Schinus molle L. Var. Molle (Anacardiaceae) 177

An ointment containing the EOs (3.5, 2.0, or 1.75% final concentration) and
vaseline showed wound healing activity in cattle and Balb C53 mice, being 2% the
most effective EO concentration (Alba González et al. 2009).
The aqueous extract of fruits showed inhibitory effect on the angiotensin-­
converting enzyme in dogs and rabbits (Olaffson et al. 1997).
The hexane extract and the EO from fruits had activity against Streptococcus
pneumoniae (MIC 62.5  mg  ml−1), attributed to γ-cadinene (Molina-Salinas et  al.
2007; Pérez-López et al. 2011). The hexane extract was also effective against H.
influenzae and Mycobacterium tuberculosis (Molina-Salinas et al. 2007). The meth-
anolic extract was active against Proteus spp., Streptococcus spp., Escherichia coli,
and Entereobacter spp. (Matela et  al. 2018). The EO and the hexane extract
(30 μg ml−1) have antimicrobial activity against E. coli 0157:H7 and Bacillus cereus
(Deveci et  al. 2010). The ethanolic extract of fruits was effective against
Staphylococcus aureus (Cruz-Carrillo et al. 2010), S. aureus ATCC 25923, and E.
coli ATCC25922 and has a weak activity against P. aeruginosa ATCC 27853 (de
Mendonça Rocha et al. 2012). The ethanol extract (25 μg ml−1) was active against
Candida albicans ATCC 10231 (inhibition halo ≥20 mm) that colonize dental pros-
thesis, but at a lesser extent than fluconazole (inhibition halo ≥31 mm) at the same
concentration (Saravia-León and Guillinta-Vallejos 2012). The EO from leaves
showed a low activity against C. albicans (MIC 2.5 mg ml−1), but it was effective
against Fusarium solani, Aspergillus flavus, A. japonicus, A. niger, A. oryzae, A.
fumigatus, A. parasiticus ATCC 16992, Fusarium moniliforme ATCC 52539,
Alternaria alternata (Fries), and Penicillium italicum Wehmer (Dikshit et al. 1986;
Alanís-Garza et al. 2007; Rhouma et al. 2009; Martins et al. 2014; López-Meneses
et al. 2015).
The EOs are used as food preservative for their antioxidant and antimicrobial
activity (Guala et al. 2016), for example, against Salmonella (103 CFU g−1) inocu-
lated in minced beef meat (Hayouni et al. 2008). Chitosan nanoparticles loaded with
S. mole leaf extract (0.05% v/v) were effective against Aspergillus parasiticus
spores, reducing the viability of fungal spores as the concentration of the chitosan:
S. molle bionanocomposites raised (Luque-Alcaraz et al. 2016).
S. molle ethanol extracts and rutin extracted from leaves showed anti-lymphoma
potential on induced lymphoma mice models (Male CD-1 nu/nu nude), cytotoxic
activity on U-937 cells, and strong lethality on brine shrimp (BS) larvae. The anti-
cancer property was connected with the presence of α-phellandrene and rutin
(Calzada et  al. 2018). It also showed antitumor and cytotoxic activity (IC50
50 ± 7 μg ml−1) against the human hepatocellular carcinoma cell line, Hep G, and
cytotoxicity on breast carcinoma and leukemic cell lines (Ruffa et al. 2002; Díaz
et al. 2008).
The aqueous extract had a discrete inhibitory activity of trypsin and elastase that
was postulated as partially responsible of the insecticidal activity (Macció et  al.
2014). The crude hexane extract from fruits was effective against Cydia pomonella
(Linneaeus) larvae, a phytopathogen of fruit trees, with 60% mortality at 5 g kg−1
concentration; from the surviving adults, a great proportion showed malformations
(Chirino et  al. 2001). The hexane extract from leaves was active against oriental
178 7 Yungas

adult cockroach (Blatta orientalis L.) with a 83.33% repellency at a concentration

of 0.075 mg l−1 (Deveci et al. 2010). The EO from leaves (0.78% w/v) was active
against bed bugs (Cimex lectularis), reaching a 100% of efficiency after a 24-hr
treatment (250 μg 125 cm−2); that activity was attributed to γ-pinene, muurolol, and
limonene content (Machado et al. 2018). The aqueous and ethanol extracts had lar-
vicidal and pupicidal activity against immature Culex quinquefasciatus with an
increased mortality as exposure time and concentration rise (100% mortality of
third instar after 72 h exposure at 100 mg l−1) (Girmay et al. 2014). Also, the aque-
ous and methanolic extract (4%) of leaves was active against the adult Bophilus
decoloratus and Rhipicephalus pulchellus cattle ticks (Feyera and Abdisa 2016).
Activity of the n-hexane extract against eggs and adults of Ctenocephalides felis
felis, a flea that infests dogs and cats in Brazil, was in vitro demonstrated. Efficacy
was 100% at 800 μg cm−2 (LD50 524.80 μg cm−2 after 48 h). EOs from fruits and
leaves showed 100% efficacy against adult fleas at 800 μg cm−2 (LD5 353.95 μg cm−2)
and at 50 μg cm−2 (LD50 12.02 μg cm−2), but not against flea eggs (Batistia et al.
2016). The EOs from seeds produced 77–100% mortality of Drosophila melano-
gaster Meigen (Diptera: Drosophilidae) and had a repellent effect on Tribolium
confusum Jacquelin du Val (Coleoptera: Tenebrionidae) (Steimbauer 1995).
In the case of Trypanosoma cruzi, the methanolic extract (150 μg ml−1) resulted
effective against epimastigotes (IC50 16.31 μg ml−1) after 96 h of treatment (Molina-­
Garza et  al. 2014); the crude hydro-alcoholic extract (250  μg  ml−1) produced a
higher inhibition (IC50 50 μg ml−1) (Pizzolatti et al. 2002). The differences were
attributed to variation in bioactive concentration, geographical origin, etc. of the
extracts. The extract from leaves was repellent and insecticidal against Triatoma
infestans (vinchuca, winchuka) a blood-sucking bug that is the main vector of the
Chagas-Mazza disease (Ferrero et  al. 2006). An acaricidal effect against varroa
(Varroa destructor) parasitizing hive of bees and against tick (Rhipicephalus micro-
plus) was also reported (Torres et al. 2012; Guala et al. 2014; Guala et al. 2016).
On the other hand, a slight activity of the aqueous extract (50% and 100% w/v)
from leaves on Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae) showed 16%
mortality (Guevara et al. 2018). On the contrary, the antifeeding effect of the aque-
ous extract (2.5, 3.0, 4.3, and 5.6% w/v) on the leaf beetle Xanthogaleruca luteola
(Muller) (Coleoptera: Chrysomelidae) produced a 98.32% antifeeding effect. The
ethanolic extract produced more larval mortality (LC50 1.28% w/v at day 7) than
the aqueous extract at the same concentration (Huerta et al. 2010; Chiffelle et al.
2013). An insecticidal effect was also reported against Sitophilus zeamais
Motschulsky (Coleoptera: Curculinoideae) (Silva et  al. 2005; Benzi et  al. 2009;
Awoke et al. 2014; Arias et al. 2017) and Sitophilus oryzae L. (Benzi et al. 2009).
Crude leaf methanolic extract had antihelmintic activity against Haemonchus con-
tortus, a parasite of small ruminants (Zenebe et al. 2017). Leaf EO has allelopathic
activity on Allium cepa and Lactuca sativa (Simionatto et al. 2011).
Toxicity  The EO was reported not to be toxic for humans (Carrasco 1998; Guba
2008) nor carcinogenic for animals (Guba 2008). The EO was highly toxic (LC50
47 𝜇g ml−1) to Artemia salina (Martins et al. 2014). Also, it was cytotoxic to breast
7.5  Smilax campestris Griseb. (Smilacaceae) 179

carcinoma, leukemia cell line K562 (0.89 μg ml−1), and NCI-ADR/RES (ovarian

expressing the resistance phenotype for adryamycin) (3.40 μg ml−1), attributed to
caryophyllene oxide (Díaz et  al. 2008). Immunotoxicity to human lymphocytes
(LD50 30.07 μg ml−1) and human macrophages (42.07 μg ml−1) was also reported.
A reduction in IL-10 production at low concentrations was correlated with the
increase in TNF-ǖFC; production (Apollo Duarte et al. 2018).

7.5  Smilax campestris Griseb. (Smilacaceae)

Synonymy  Smilax campestris Griseb. var. rubiginosa, Smilax rubiginosa Griseb.,

Smilax campestris Griseb. var. marginulata, Smilax marginulata Mart. ex Griseb.,
Smilax montana Griseb., Smilax scalaris Griseb., Smilax viminea Griseb.

Vernacular name  Zarzaparrilla, zarzaparrilla negra, zarzaparrilla blanca, japi-

canga, japecanga (Portuguese).

General information  S. campestris grows in temperate thickets and grasslands

near rivers and streams, in mountains, and fields at high altitude (1800 m.a.s.l.) in
the provinces of BAI, CHA, COR, COS, ERI, FOR, JUJ, MIS, SAL, SDE, SFE, and
TUC. The phenotypic plasticity of leaves and rhizomes is due to ecological factors
(Andreata 1997). Blooming is from July to September. A tonic, bitter, and digestive
beverage is prepared from the leaves and aerial stems (Rugna et al. 2003, 2011). It
contains steroid saponins which are used as precursors in the semi-synthesis of
cortisone.

Botanical characteristics  S. campestris is a dioecious climbing plant about 2–4 m

high with lignified rhizome and tuberose hickening. The underground stem runs
horizontally near the ground surface (10–15 cm depth). The woody, brown stem,
approximately 5–10 cm length, has swollen nodes that can produce negative geotro-
pic stems and adventitious roots. Leaves are variable both in shape and in size
(ovate, oblong-elliptic, elliptical, oblong, rarely cordate) with obtuse, acute, or
emarginated coriaceous apex. Basal acrodrome venation has three major midribs,
with spiniform emergences in midrib extensions. Inflorescences are umbels with
30–45 flowers, terete peduncle, and 2–20-mm-long globose receptacle. Perigone is
purple-red, with elliptic tepals and papillae at the apex. The globular ovary contains
hemotrope ovules, 1 per locule. The fruit is a black-violet globose berry with purple
pulp, and subglobose seeds, wrapped by a layer of hyaline, elastic tissue (Guaglianone
and Gattuso 2006; Martins et al. 2010; Martins et al. 2013; Colares et al. 2017).

Ethnobotanical information  The roots are considered as antisyphilitic, diapho-

retic, and diuretic, for infections, hernia and prolapse, and menstrual diseases.
Rhizomes are considered as antisyphilitic, diuretic, sudorific, aperitive, refrigerant,
anti-gout, hypolipemic, and laxative. The aerial parts are used as anti-inflammatory,
180 7 Yungas

antidiarrheal, and for healing wounds. The leaves are considered as antisyphilitic
and antirheumatic. Finally, flowers are used to treat vaginal fungus. It is also pre-
scribed to treat impotence, rheumatism, gout, and skin conditions; it is considered
diuretic and diaphoretic. The roots and the basal parts of the stems are used as
diuretics and laxative for infants (Mandrile and Bongiorno de Pfirter 1991;
Toursarkissian 1980; Rodrigues and Carvalho 2001; Barboza et  al. 2009; Silva
et al. 2013).

Chemical data  Leaves contain kaempferol, kaempferol-3-O-rutinoside, quercetin,

quercetin-3-O-glucoside, quercetin-3-O-rhamnogalactoglucoside, isorhamnetin,
isorhamnetin-3-O-rutinoside, and rutin. In rhizomes there are quercetin-3-­glucoside,
3-rutinoside, procyanidin, and propelargonidin (Fig. 7.5) (Rugna et al. 2002a, 2004,
2008, 2013; Barboza et al. 2009; Morais et al. 2014; Salas-Coronado et al. 2017).

Biological activity  The antioxidant activity from rhizomes, which was partly
attributed to the proanthocyanidin content, follows a circannual pattern. The highest
antioxidant activity is reached in summer (February) with two more peaks during
pre-flowering and fructification (Rugna et al. 2002b, 2003). The ethanol and butanol
fractions showed a strong antioxidant activity.

Fig. 7.5  Chemical compounds from Smilax campestris Griseb

7.6  Smallanthus macroscyphus (Baker) A. Grau ex A.A. Sáenz (Asteraceae) 181

The methanolic extracts (80%) from rhizome showed antifungal activity against
Candida albicans and Aspergillus niger (Battista et al. 2007). The ethanol extract
and its fractions were active against C. albicans, C. glabrata, C. krusei, C. parapsi-
losis, C. tropicalis, and Cryptococcus gattii. The ethyl acetate fraction exhibited the
highest activity against C. krusei, (MIC 62.5 mg ml−1) and was also active against
the C. gattii (MIC 1000 mg ml−1). The hexane fraction was active against C. gla-
brata (MIC 125 mg ml−1) and C. krusei (MIC 500 mg ml−1). The ethanol extract was
active against C. krusei, C. parapsilosis and C. gattii (MIC 1000 mg ml−1), and C.
albicans, C. glabrata, and C. tropicalis (MIC 2000 mg ml−1). The dichloromethane
fraction displayed activity against C. krusei and C. tropicalis (MIC 2000 mg ml−1).
Based on those results, a new therapeutic strategy could be implemented specifi-
cally against C. krusei, which presents intrinsic resistance to fluconazole, and
against C. gattii, which is implicated in HIV-associated infections (Morais et  al.
2014). The ethanolic extract of leaves showed inhibitory activity against
Staphylococcus epidermidis (MIC 1000 mg ml−1) and a worthy antioxidant activity
(IC50 13.1 ± 0.1 mg ml−1) (Ferreira de Lima 2016).
In vitro culture  In vitro cultures were initiated from seeds in MS solid culture
medium without plant growth regulators. Explants transferred to MS medium with
the addition of different plant growth regulator relationships (BAP, IAA) were
tested to regenerate the whole plant. A methanolic extract (50%) of leaves from
regenerated plantles rendered kaempferol, quercetin, and isorhamnetin. The highest
total phenolics content was obtained in MS medium with BAP (2 mg ml−1) and IAA
(0.4 mg ml−1) (Rugna et al. 2010).

Toxicity  Mutagenic activity on Salmonella/microsome assay was reported. S.

campestris infusions had direct mutagenic activity on TA98 strain (Fernández de
Sá Ferreira and Vargas 1999).

7.6  S
 mallanthus macroscyphus (Baker) A. Grau ex
A.A. Sáenz (Asteraceae)

Synonymy  Polymnia macroscypha Baker, Smallanthus riograndensis

C.A. Mondin, Smallanthus araucariophilus C.A. Mondin.

Vernacular name  Yacón del campo, wild yacon.

General information  S. macroscyphus belongs to the yacon group whose main

representative is S. sonchifolius. Smallanthus species show a clear preference for
disturbed habitats, such as riverbanks, landslides, and roadsides. The distribution is
affected by precipitation seasonality (Grau and Rea 1997; Vitali and Katinas 2015).
This specie is an invader of abandoned sugar cane fields as well as of the free space
between land plots (Serra-Barcellona et al. 2014). In Argentina, S. macroscyphus
182 7 Yungas

grows in the provinces of JUJ, SAL, and TUC. It has been consumed as a fruit since
pre-Columbian times. Lately it was revalued as a functional or nutraceutical food
(Grau and Rea 1997; Aybar et al. 2001). Under modern nutritional and agronomic
requirements, some traits like carbohydrate high polymerization degree and short
growth cycle may be very useful in breeding programs (Grau and Rea 1997; Coll
Aráoz et al. 2008, 2014). Blooming is from December to June.

Botanical characteristics  S. macroscyphus is a perennial herbaceous plant, 1–3 m

tall, with a caulescent rhizome or rhizophore that gives rise to tuberose fusiform
adventitious roots and aerial caulescent branches. The stem is pubescent at the apex
and almost glabrous by maturity with opposite decussate opposite simple, green-­
yellowish leaves. The leaf blade is membranous, pubescent, of triangular symmetry,
and deltoid ovate, with an acute apex and margins irregularly scalloped. Terminal
inflorescences are capitula clusters arranged in corymbiform or paniculiform cymes,
with yellow tubular corolla and ovate, subtriangular anthers (Coll Aráoz et al. 2008;
Vitali et al. 2015).

Ethnobotanical information  The roots are used to treat urinary and hepatic disor-
ders; it is also used as hypoglycemic, antitumoral, antimicrobial, antifungal, and
anti-inflammatory (Hilgert 2000; Cabrera et al. 2007; Barboza et al. 2009; Serra-­
Barcellona et al. 2012; Vitali 2017).

Chemical data  Kaurenoic acid methyl ester is the predominant diterpene in the root
cortex; also there are fructooligosaccharides with high polymerization degree. In the
aerial parts, there are melampolides, melampolide-type sesquiterpene lactones
(STLs), (6R∗,7S∗,8S∗,9S∗)-8-hydroxy-9-methacryloxy-10-­carbomethoxymelampo-
1(10),4,11(13)-trien-6,12-olide, (6R∗,7S∗,8S∗, 9S∗)-8-Hydroxy-9-
angeloxy-10-carbomethoxymelampo-1(10), 4,11(13)-trien-­ 6,12-olide, and
( 4 S ∗ , 5 S ∗ , 6 R ∗ , 7 R ∗ , 8 S ∗ , 9 S ∗ ) - 4 , 5 - e p o x y - 8 - h y d r o x y - 9 - a n g e l o x y - 1 0 -­
carbomethoxymelampo-­1(10),11(13)-dien-6,12-olide (de Pedro et al. 2003; Barboza
et al. 2009; Coll Aráoz et al. 2010, 2014). The decoction of leaves contains caffeic
acid, 3-caffeoylquinic acid (ac. chlorogenic), three dicaffeoyl quinic acids (3,4-dicaf-
feoylquinic acid, 3,5-dicaffeoylquinic acid, and 4,5-dicaffeoylquinic acid), and the
lactone polymatin A (Fig. 7.6) (Serra-Barcellona et al. 2016). The sesquiterpene lac-
tone content varies according to the latitude. In lower latitudes the sesquiterpene lac-
tone profile is more complex containing mainly uvedaline, fluctuanine, and enhydrin.
In higher latitudes there are mainly polymatin A and a few other lactones as isopoly-
matin A and sonchifolin (Coll-Aráoz et al. 2016).

Biological activities  The sesquiterpene lactones are produced as a mechanism of

defense in glandular trichomes of the foliar surface. They accumulate underneath
the cuticle and are released when the glandular trichomes are mechanically dis-
rupted. The biological activity of those sesquiterpene lactones is deterrent to herbi-
vores, toxic to pathogens like fungi and bacteria, and allelopathic (Coll-Aráoz et al.
2016).
7.6  Smallanthus macroscyphus (Baker) A. Grau ex A.A. Sáenz (Asteraceae) 183

Fig. 7.6  Chemical compounds from Smallanthus macroscyphus (Baker) A. Grau ex A.A. Sáenz

The aqueous extract of dried S. macroscyphus leaves has high hypoglycemic

effect on rats. Glucose tolerance tests on Wistar rats showed that both the leaf decoc-
tion and polymatin A significantly diminished glycemia. Daily administration of the
decoction (70  mg polymatin A 100  ml−1, dose 8  ml  kg−1) or a water solution of
polymatin A (dose 7 ml kg−1) during 30 days produced a strong decrease of glyce-
mia in diabetic rats. On the other hand, the aqueous extracts (10% decoction,
140 ml kg−1) of dried S. macroscyphus leaves produce a hypoglycemic effect in a
dose-dependent manner on adult male Wistar rats with induced diabetes. Polymatin
A has an effective inhibition of postprandial blood glucose peak and a hypoglyce-
mic activity in diabetic animals. Probably, the glucose-lowering effect of the decoc-
tion could be mediated by an improvement in the glycemic control mechanisms,
including insulin secretion and/or extrapancreatic pathways without risk of hypo-
glycemia, which is one of the most undesirable side effects of diabetes treatments.
Moreover, the treated animals reduced urinary glucose excretion. Polymatin A was
identified as the major hypoglycemic principle of S. macroscyphus (Cabrera et al.
2007; Coll Aráoz et al. 2008; Serra-Barcellona et al. 2014).
Toxicity  The 10% decoction and pure polymatin A have a safety margin in rats at
the doses that led to the hypoglycemic effect. The aqueous extract of leaves orally
administered is nontoxic, at least up to the maximum level assayed
(280 mg kg−1 day−1). Similarly, polymatin A has no toxic effect even at the highest
dose tested (28 mg kg−1 day−1) (Serra-Barcellona et al. 2016).
184 7 Yungas

The lethal dose 50 (LD50) of the 10% decoction of leaves and of polymatin A
(above 14.0 and 2.8 g kg−1 b.w., respectively) is significantly higher than the effec-
tive hypoglycemic doses.

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Chapter 8
Chaco and Espinal

8.1  Introduction

8.1.1  Chaco

The Chaco region extends through the province of FOR, CHA, eastern SAL, JUJ,
TUC, CAT and LRI; all of SDE; north of SLU, COR and SFE Fe; and northwest of
COR (Cabrera 1971). It covers plains and mountains of low elevation with mostly
subtropical climate, warm and humid summers, and temperate and dry winters
(Fig 8.1a). Rains are concentrated in summer, and occasionally a wave of polar cold
is perceived. As for the fauna, large emblematic animals such as the Panthera onca
(yaguareté) and the Priodontes maximus (tatú carreta, giant armadillo) are in critical
danger of extinction. The Myrmecophaga tridactyla (giant anteater) and the Tapirus
terrestris (tapir) are also among the typical fauna of the eco-region, as well as mon-
keys, coatis, foxes, and weasels.
In the Dry Chaco, the characteristic vegetation is the xerophilous forest, with
trees that become lower and thinner towards the Arid Chaco. The deciduous xeroph-
ilous forest predominates, having also palmares, savannahs, and halophilic steppes.
The highest areas have xeric forests (quebrachales) of red quebracho and white
quebracho, with mistol, itin, yuchán, pitch, several cacti, duraznillo, and bushes of
the Acacia (tusca, teatín) and Capparis (sacha-quince, sacha-watermelon) genus.
In some low areas, salinity and restrictions in drainage condition the floristic com-
position, giving rise to communities of palo santo (Bulnesia sarmientoi, the palos-
antales), carob trees (Prosopis spp.), and chañar (Geoffroea decorticans) and, in the
salt flats, to vegetation with a predominance of halophytic species. The forests alter-
nate with pastures (“Pampas”) of aibé (Stipa paposa), associated with old clogged
channels. There are also horco-quebracho, visco, churqui (espinillo), molle, molle
de coco, and coconut. A particular formation of the Córdoba and Puntana mountain
ranges is the palmares of carandilla (Trithrinax spp.). Above these forests within the
altitudinal gradient are graminoid steppes, which form biogeographic islands with

© Springer Nature Switzerland AG 2019 193

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_8
194 8  Chaco and Espinal

Fig. 8.1  Chaco (a) and

Espinal (b) eco-regions
from Argentina. MA
Alvarez personal collection

features of the Puna. Tabaquillo (Polylepis australis) groves appear in these tall
grasslands.
In the Humid Chaco, the high forests of the albardones are mainly composed of
quebracho colorado chaqueño (Schinopsis balansae), quebracho blanco
(Aspidosperma quebracho-blanco), guayaibí (Patagonula americana), urunday
(Astronium balansae), lapacho (Tabebuia sp.), guayacán (Handroanthus chrysan-
thus), viraró (Ruprechtia salicifolia), palo amarillo (Phyllostylon rhamnoides, ibirá-­
catú), and mistol (Ziziphus mistol). There are dense or open forests, bordering
estuaries and ravines, mainly composed of carob (C. siliqua), tatané (Chloroleucon
tenuiflorum), guaraniná (Sideroxylon obtusifolium), churqui (A. caven, espinillo del
Espinal), and Prosopis sp.; grasslands, which are composed of wire grass as domi-
nant species, accompanied by many other herbaceous and often tree-like elements
(forming savannahs) of low-forest species; cañadas, constituted by pure scrublands
(pajonales) or by palm tree forests (palmares), with white palm (Trithrinax camp-
estris or caranday); and estuaries, without vegetation cover and with diverse aquatic
species, such as pirí, peguajó (Thalia geniculata), totoras (Schoenoplectus califor-
nicus), cucharero (Echinodorus grandiflorus), and camalote (Eichhornia crassipes),
among others.
8.2  Adiantum raddianum C. Presl (Pteridaceae) 195

8.1.2  Espinal

The Espinal extends in the form of an irregular arch around the LPA province
through the center of SFE and COR, a large part of SLU, the center of LPA to the
south of BAI. Towards the south it extends along the ravines of the Paraná river and
through the banks of conchilla and dead dunes of the northeast of BAI, and towards
the West it penetrates through the depressions and along the rivers until finally
reaching the area of Sierra de los Padres (Cabrera 1971). This is a region of plains,
low mountains areas, and dunes (Fig 8.1b). The climate is, due to the extension of
the region, extremely variable, warm and humid in the north, and temperate and dry
with marked water deficits, towards the west and south.
Many animals from the Pampa, affected by hunting and habitat transformation,
are more frequent or are only associated with the Espinal forests. The most emblem-
atic case is the pampas deer, since two of its last populations are located in the
grasslands with calden (Prosopis caldenia), in San Luis, and in the pastures with
talas (Celtis spinosa), in the northeast of Buenos Aires. Other examples are the
wildcat, the gray fox, and the rhea. Much of the Espinal is located in regions of
high-agricultural and urban development, which is why Espinal has been greatly
reduced during the last decades. The creation of more protected natural areas and a
sustainable management of livestock are the challenges the Espinal is currently fac-
ing. The Espinal can be divided into three subregions, namely: Espinal del
Ñandubay, the Mesopotamian sector, with a humid climate; the Espinal de los
Algarrobos, the central, subhumid sector of transition between the Pampa and the
Chaco; finally, Espinal del Caldén, semiarid sector, transition between the Pampa
and the Monte. In Espinal predominates the xerophilous forest vegetation  that  is
quite similar to the Chaco region but more impoverished. There are also palm
groves, grass savannahs, and steppes, and predominant trees are Prosopis spp. In the
northeast of the Espinal del Ñandubay, algarrobos (Prosopis sp.), molle (Schinus
molle), and espinillo (A. caven) forests predominate, with other species, among
which the palm trees yatay (Butia yatay) and carandilla, which are associated with
the forest or palm trees. In the center of the eco-region, the dominant Prosopis spe-
cies are the white and black algarrobo (carob) that are accompanied by A. caven, G.
decorticans, and C. spinosa. The herbaceous plants are mostly pampeanas species,
with predominance of grasses characteristic of temperate grasslands and, in part,
chaqueñas.

8.2  Adiantum raddianum C. Presl (Pteridaceae)

Synonymy  Adiantum cuneatum Langsd. & Fisch., hom. illeg., Adiantum aemulum
T. Moore, Adiantum colpodes T. Moore, Adiantum moorei Baker, Adiantum ama-
bile T. Moore, hom. illeg.
196 8  Chaco and Espinal

Vernacular name  Culantrillo, culandrillo, curatú, tipichá, avença (Portuguese),

maidenhair ferns (English).

General information  A. raddianum is a widely distributed species in Brazil, Peru,

Uruguay, and Argentina. In Argentina it is located in the provinces of BAI, CAT,
CHA, COR, COS, ERI, JUJ, LRI, MIS, SAL, SFE, SLU, and TUC. It has tolerance
to different light conditions, and it usually grows isolated or forming bushes in the
riverbanks, rock walls, waterfalls, slope forests, swamps, and ravines. It is cultivated
as ornamental (Fujino et al. 1983; Silva et al. 2007; de la Sota et al. 2009; Arana and
Bianco 2011; Arana et al. 2013).

Botanical characteristics  A. raddianum is a fern with a terrestrial or saxicolous

perennial sporophyte, creeping suberect rhizomes without clathrate scales, exception-
ally with hairs. Petiole is shiny brown, and deltoid pinna is glabrous, several times
divided, with scales or wax and free to reticulated veins (Fig. 8.2). They have polysac-
charide mucilage in the epidermis that would function as protection from solar radia-
tion. The sporangium (reproductive structure) is located alongside and at the end of
the veins forming marginal-clustered sores or covering the entire surface of the blade
or segment, without indusium or with a false glabrous kidney-shaped indusium, or
protected by the modified leaf margin. Spores are trilete, with or without perisporium
(Arana and Bianco 2011; Santos-Silva et al. 2011; Winter et al. 2011).

Ethnobotanical information  In folk medicine, leaves are considered abortive,

antitussive, and emmenagogue, with the ability to dissolve the mucus for respiratory
disorders, as diaphoretic, emollient, diuretic, and expectorant, for treating pain,
coughs, urinary disorders, alopecia, and menstrual difficulties (Martínez Crovetto
1981; De Feo 1992; Bresciani et al. 2003; Silva et al. 2007; Barboza et al. 2009).

Chemical Data  The compounds identified in the spores are the alkaloids spermi-
dine and N(1)-N(10)-caffeoyl-feruloyl spermidine (Fig. 8.3).
The whole plant contains the flavonoids, querciturone, kaempferol, astragalin, the
terpenoids filicene (filic-3-ene), filicenal, adiantol (adian-5-en-3α-ol) and isoadian-
tone, secofilicanes, hydroxyhopane, neohopene, trisnorhopane, 17 αH-trisnorhopan-­
21-one, glaucanol B acetate, 21-hydroxy-30-norhopan-22-one, isoadiantol B,
fern-9(11)-en-25-ol, fernane, fern-7-en-25-ol, 25-norfern-7-en-10b-yl formate,
7α,8α-epoxyfernan-25-ol, 7β,25-epoxyfern-8-ene, 7β,25-epoxyfern-9(11)-en-8α-ol
(Fig. 8.4) (Shiojima et al. 1997; Bresciani et al. 2003; Barboza et al. 2009; Guerra
Santos et al. 2010; Lai and Lim 2011; Pan et al. 2011; Thomas 2014).
Biological activity  The main biological activities reported were antioxidant, anti-
nociceptive, and antimicrobial. The antioxidant activity was attributed to its content
in phenolic compounds and particularly flavonoids (Lai and Lim 2011; Shrivastava
et al. 2014).
The hexane fraction obtained after a methanolic extraction from raw material, as
well as pure filicene and filicenal, showed potent analgesic activity in two models of
8.2  Adiantum raddianum C. Presl (Pteridaceae) 197

Fig. 8.2  Adiantum raddianum C.  Presl (Pteridaceae). Buenos Aires Botanical Garden. M A
Alvarez personal collection

OH

OH
H
H2N N N

O
O
O

H2N NH3
N
H HO

spermidine N(1)-N(10)-caffeoyl-feruloyl spermidine

Fig. 8.3  Chemical compounds found in Adiantum raddianum C. Presl spores

pain in mice (writhing test and formalin-induced pain) (Bresciani et al. 2003; Pan
et al. 2011). Filicene had antinociceptive effect in male Swiss mice challenged with
acetic acid-, capsaicin-, and glutamate-induced nociception tests and hot-plate and
open-field tests, each test involving different systems. Apparently, the mechanism of
action involves interactions with the cholinergic, dopaminergic, glutamatergic,
GABAergic, and tachykinergic systems. Filicene had higher antinociceptive activ-
ity of inflammatory pain (acetic acid-induced abdominal constrictions) in the ani-
mal model and was less effective in models of acute pain (glutamate and capsaicin
tests). It was not found interacting with the opioid, serotoninergic, or adrenergic
systems (de Souza et al. 2009).
198 8  Chaco and Espinal

Fig. 8.4  Some chemical compounds found in Adiantum raddianum C. Presl

For testing the antimicrobial activity, successive extractions with petroleum

ether, acetone, methanol, and water were made from air-dried fronds with sori. A
sample of each extract (0.46%, 2.8%, 5.1%, and 0.70%, respectively) was tested for
its antibacterial activity against bacteria involved in skin infections such as
Staphylococcus aureus subsp. aureus (MTCC 96), Escherichia coli (MTCC 443),
Pseudomonas aeruginosa (MTCC 741), Klebsiella pneumoniae subsp. pneumoniae
(MTCC-109), and Serratia marcescens (MTCC 6164). Antibacterial activity was
not found in the aqueous and petroleum ether extracts. Acetone extract showed
moderate level of inhibition towards S. aureus and a lower level of inhibition
towards E. coli. The methanol extract was effective against P. aeruginosa and S.
marcescens and S. aureus (MIC 50 mg ml−1, MBC 25 mg ml−1) (Thomas 2014).
Moreover, silver nanoparticles (AgNPs) were elaborated with aqueous extracts
(10 ppm) from A. raddianum fronds. The antibacterial activity of those particles
(1 nmol ml−1) was tested against S. aureus, Proteus spp., K. pneumoniae, and E. coli
resulting in significant inhibition of both P. aeruginosa (inhibition zone 10  mm)
and K. pneumoniae (inhibition zone 8 mm), a moderate activity against S. aureus,
and no activity against Proteus spp. The potential of A. raddianum aqueous extracts
for developing novel eco-friendly biocidal formulations was hypothesized (Dev
et al. 2018).
Toxicity  Toxicity analysis is not available (Rastogi et al. 2018).
8.3  Aloysia citriodora Palau (Verbenaceae) 199

8.3  Aloysia citriodora Palau (Verbenaceae)

Synonymy  Aloysia citriodora Ortega ex Pers., hom. illeg.; Aloysia sleumeri

Moldenke; Aloysia triphylla (L’Hér.) Britton, hom. illeg.; Lippia citriodora (Ortega
ex Pers.) Kunth, comb. illeg.; Lippia triphylla (L’Hér.) Kuntze; Verbena citriodora
Cav., hom. illeg.; Verbena triphylla L’Hér.; Zappania citriodora Lam., hom. illeg.;
Aloysia triphylla (L’Hér.) Britton f. serrulata.

Vernacular name  Chaquesayasaya, cedrón, verbena citronela, limón verbena,

haurippampacay, verbena olorosa, hierba de la princesa, yerba Luisa, vervein
odorante.

General information  A. citriodora is a herbal species distributed from the South

of the United States down to South America and cultivated worldwide. It grows in
light and well-drained soils and in temperate-warm climate resisting frost quite
well. Their leaves are very aromatic; hence, it is frequently used as a flavor essence
and odor corrector in food industry and pharmacy, as well as in perfumery and cos-
metics. It is also employed in the elaboration of beverages, sweets, cookies, des-
serts, and pastry. In Argentina A. citriodora grows in the provinces of CAT, JUJ,
LRI, and SAL (Rzedowski and Calderón de Rzedowski 2002; Bandoni et al. 2003;
Santos-Gomes et al. 2005).

Botanical characteristics  A. citriodora is a woody, profusely branched shrubby

plant, 3–3.5 m tall, with woody stem, and simple, thin, elongated, rough leaves with
a very characteristic sweet and sour lemon smell. Leaves are opposite, subsessile to
shortly petiolate, with an entire to serrated margin. The small flowers are white or
pale violet in the outside and violet in the inside, usually arranged in terminal spikes.
Mericarps were glabrous and rough on the outside (Muñoz López de Bustamante
1996; Rzedowski and Calderón de Rzedowski 2002; Gattuso et al. 2008; Múlgara
de Romero et al. 2012; Calzada-Sánchez et al. 2014).

Ethnobotanical information  There are reports about the traditional use of this
species dating back to the seventeenth century; it was considered as a medicinal
plant by the Inca culture. The infusion and decoction are traditionally used as diges-
tive, eupeptic, antispasmodic, mild sedative, expectorant, carminative, and tonic.
The infusion and decoction made with leaves are taken in cases of stomachache, as
cardiotonic, febrifuge, diuretic, antispasmodic, in odontalgia, and against hemor-
rhoids. The aerial parts are used as antiemetic, for heart problems and backache.
There are also references about its addition to the traditional beverage mate (Gupta
1995; Barboza et al. 2009; Elechosa et al. 2017; Bahramsoltani et al. 2018).

Chemical data  The leaf essential oil has neral, geranial, nerol, geraniol, bicyclo-
sesquiphellandrene, spathulenol, nerolidol, ß-bourbonene, trans-ß-caryophyllene,
caryophyllene-2,6-β-oxide, limonene, geranyl acetate, neryl acetate, trans
200 8  Chaco and Espinal

c­ alamenene, β-citronellene, β-pinene, neryl acetate, α-thujone, cis-carveol, carvone,

1,8-cineole, caryophyllene oxide, and linalool. The leaves have vervain, geniposidic
acid, geniposide, mussaenoside, luteolin 7-diglucuronide, salvigenin, eupatorin,
eupafolin, verbascoside (or acteoside), hispidulin, 6-hydroxyluteolin, luteolin
7-O-β-glucoside, luteolin, cismaritin, diosmetin, cirsiliol, chrysoeriol, apigenin,
pectolinarigenin, p-cymene, and ß-bourbonene. The flower oil has 1-octene, 1-octen-
3-ol, p-cymene, (Z)-β-ocimene, and trans-carveol (Fig. 8.5) (Lamaison and Petitjean
1993; Carnat et al. 1999; Barboza et al. 2009; Ricco et al. 2011). The content and
composition of the EOs differ according to the region and the phenological state of

Fig. 8.5  Chemical compounds from Aloysia citriodora Palau (Verbenaceae)

8.3  Aloysia citriodora Palau (Verbenaceae) 201

the plant (Molina et  al. 2003; Di Leo et  al. 2007; Oliva et  al. 2010; Ricco et  al.
2011). According to Di Leo et al. (2007), in Argentina A. citriodora leaves contain
more than 0.2% of oil (v/w of dried material). Four different chemotypes, thujones,
citronellal, carvone, or citral, were identified in northwestern Argentina depending
on the predominating component from the EO (Elechosa et al. 2017).

Biological activities  The biological activities reported for A. citriodora aerial parts
are giardicidal, analgesic, anti-inflammatory, antipyretic, sedative, and proconvul-
sant. The EO has antidiarrheal, nematicidal, protective, antimicrobial, antigeno-
toxic, and spasmolytic activities (Barboza et al. 2009).
The infusion made with crushed leaves had high phenylpropanoid, glucuroni-
dated flavonoid, and iridoid glycoside content that correlates with its high antioxi-
dant activity (Rodríguez Vaquero et  al. 2010). One of the chemical compounds
found in A. citriodora, verbascoside, was reported to have antioxidant, anti-­
inflammatory, and chemopreventive activities (Alipieva et  al. 2014). There are
numerous reports about the antimicrobial activity of A. citriodora EO and its main
components (Bagher Hashemi et al. 2018; Pérez Zamora et al. 2018). The acetone
extract from aerial parts has antimicrobial activity against Bacillus subtilis,
Staphylococcus aureus, Escherichia coli, Klebsiella pneumoniae, and Proteus vul-
garis (Kumar et  al. 2008), and the ethanolic extract (20%) of leaves was active
against Listeria monocytogenes ATCC 19117 (Bayoub et al. 2010).
The EO (900 mg ml−1) was active against S. aureus, S. epidermidis, B. cereus, E.
coli, K. pneumoniae, Proteus mirabilis, Enterobacter aerogenes, K. ozaenae,
Enterococcus sp., B. subtilis, and C. albicans (Oliva et al. 2010; Oliva et al. 2015). Other
assays have reported that the EO had inhibition values between 0.05 and >2 mg ml−1
against Gram-positive bacteria being not active against Gram-negative bacteria
(Sartoratto et al. 2004). However, it showed a moderate activity on 12 of the E. coli
serotypes tested with MIC values between 400 and 1000 μg ml−1 (Duarte et al. 2007).
On the other hand, A. citriodora inhibited in vitro replication of four dengue
virus serotypes (DENV-1, DENV-2, DENV-3, and DENV-4 isolated from Colombian
patients), attributed to the EO geranial and neral content (Ocazionez et al. 2010).
Treating Trypanosoma cruzi epimastigotes with A. citriodora EO at different con-
centrations (50, 100, 250, and 500 μg ml−1) resulted in the inhibition of parasite growth
(IC50 96.49 mg ml−1) (Rojas et al. 2010). In vivo analysis of anti-T. cruzi activity in
mice has shown that parasitemia, amastigote amount, inflammatory infiltration in the
heart, and kinase-MB plasma levels were significantly reduced at the 28th postinfec-
tion day (Rojas et al. 2012). At an EO dose of 250 mg kg−1, parasitemia diminished
(85.4%) in 5-day-old male BALB/c mice, while at a dose of 300 mg kg−1 of citral, the
parasitemia reduction is lower (43.7%). Additionally, the inflammatory infiltrates
diminished 51.7% at 300 mg kg−1 citral (Rojas Armas et al. 2015).
Extracts and verbascoside were tested on an insulin-resistant hypertrophic
3T3-L1-adipocyte model to study their effects on triglyceride accumulation, inflam-
mation, and oxidative stress, while a murine model of diet-induced obesity was
studied to assess the in vivo metabolic response. It was found a dose-dependent
decrease of triglyceride accumulation. The extract showed higher activity than the
202 8  Chaco and Espinal

verbascoside, probably do to the synergistic action of the EO components (Herranz-­

López et al. 2015).
Verbascoside reduced the severity of intestinal macroscopic and microscopic
lesions and decreased pro-inflammatory cytokine levels in various models of murine
colitis (Hausmann et al. 2007; Mazzon et al. 2009), colonic injury, and inflamma-
tion in mice (Nishitani et al. 2013). The infusion offered beneficial effects against
dextran sulfate sodium-induced colonic inflammation in rats, prevented epithelial
destruction, glandular dilation and edema, and reduced the severity of colonic alter-
ations (Lenoir et al. 2012). The bioavailability and intestinal absorption of polyphe-
nols derived from lemon verbena infusion (1268  μM verbascoside, 270  μM
isoverbascoside, 217 μM luteolin, 88 μM apigenin, 674 μM diosmetin) in healthy
and colitic rats were not altered by colonic inflammation; on the contrary, urinary
excretion of hydroxycinnamic acids derived from verbascoside was affected
(Felgines et al. 2014).
The spasmolytic action of the decoction (15% w/w) was tested on isolated rat
duodenum. The aqueous extract relaxed the intestinal smooth muscle and non-­
competitively inhibited the dose-response acetylcholine curve. Vitexin, but not iso-
vitexin, was reported to contribute to the effect (Ragone et al. 2007). The hexane
extract and citral were in vitro tested for their effect on uterine contractions, in vivo
anti-inflammatory properties, and gastric wound healing capabilities (3, 10, 30, and
56 μg ml−1). Both inhibited the contractions induced by prostaglandine-PGF2α and
had anti-inflammatory activity without any injury even at the highest dose (100–
800 mg kg−1). These results supported the popular use of A. citriodora infusion to
treat menstrual pain and primary dysmenorrhea (Ponce-Monter et al. 2010).
Preliminary analysis of the hydroalcoholic extract (600 mg kg−1) effect on the
tail suspension test in Swiss albino mice showed an antidepressive effect (Barrios
et al. 2016).
The anesthetic effect in silver catfish and the recovery times from anesthesia of
subadult and postlarvae white shrimp were also tested. The EO was effective in both
cases, inducing anesthesia at the concentrations of 150 and 300  μl  l−1 being the
shortest recovery time from anesthesia (10  min) in subadult shrimp reached at
300 μl l−1. In postlarvae shrimp, the shortest induction and recovery times (less than
10 min) were both seen at 300 μl l−1. The EO induced deep anesthesia without mor-
tality (Parodi et al. 2013).
The chemoprotective effect of leaf infusions (5%) against genetic damage
induced by acrylamide (50 mg kg−1) was tested by the alkaline version of the comet
assay technique (Singh et al. 1988) in BALB/c mice. The administration of the infu-
sion before acrylamide administration reduced the capacity of acrylamide to induce
genetic damage (Zamorano-Ponce et al. 2006). A similar model was used to analyze
the protection of the A. citriodora infusion (5%) from the genetic damage induced
by cisplatin (6 mg kg−1) in mouse bone marrow cells. A significant prevention of
cisplatin-induced genetic damage was observed in animals previously treated with
the infusion. The authors attributed those results to the high-superoxide and hydroxyl
radical scavenging activity related to the content of flavonoid and polyphenols
(Valentao et al. 2002; Zamorano-Ponce et al. 2004).
8.4  Bauhinia forficata subsp. pruinosa (Vogel) Fortunato & Wunderlin (Fabaceae) 203

A repellence test against Aedes aegypti was made by an arm-in-cage assay with
the EO (90%). Differences were seen depending on the geographic origin of A.
citriodora, and the EO from plants from Traslasierra showed more repellence with
a significantly longer repellency time (90 min) than those from Córdoba, probably
due to the highest limonene content in the former one (Gillij et al. 2008). Also, the
acute larvicidal toxicity of the EO against Culex quinquefasciatus Say (Diptera:
Culicidae) was reported. C. quinquefasciatus is a vector of lymphatic filariasis and
of diseases, such as West Nile and St. Louis encephalitis. Early 3rd larvae of mos-
quito were treated with EO (5–100 μl l−1) being LC50 65.6 μl l−1after 24 h. A com-
bination of A. citriodora and Satureja montana EOs (1:1) was highly effective
(LC50 18.3  μl  l−1, LD90 28.7  μl  l−1) against C. quinquefasciatus larvae (Benelli
et al. 2017). Activity from pulegone and citral against head lice (Pediculus humanus
capitis) was also demonstrated. Knockdown and mortality activities were 47–53%
and 42–55%, respectively (Gonzalez-Audino et al. 2011).
Toxicity  Cytotoxicity on RAW 264.7 ECACC (European collection of cell cul-
tures) mammal cells at a concentration of 4 × 105 cells ml−1 was evaluated with 20,
50, and 100 μg ml−1 EO. The CI50 was superior to 100 μg ml−1; therefore, the EO
was not considered as toxic (Rojas et al. 2010). Oral acute toxicity was tested in
albino female mice with a single dose of A. citriodora EO (50, 300 or 2000 mg kg−1).
No signs of toxicity or death were observed (Rojas Armas et al. 2015).

In vitro culture  Explants (pieces of leaves and nodal segments) were cultivated in
MS, ½ MS, and ¼ MS culture medium (Murashige and Skoog 1962) with sucrose
3%, 0.7% agar, and different concentrations of BAP and NAA. Calli appeared in all
media and plant growth regulators relationship after 14 days of culture. Rhizogenesis
appeared after 60 days in MS medium with 2 mg l−1 BAP and 1 mg l−1 NAA. Stems
were obtained from nodal segments in ½ MS medium without plant growth regula-
tors (Severin et al. 2005).

Legal status  A. citriodora figures in the FNA VI ed. (1978) and in the Argentina
Food Code (1969). It is included in the pharmacopoeias of France, Spain, Mexico,
and Europe (Bandoni 2000). It figures in the GRAS (generally regarded as safe) list,
and it was considered as a dietary supplement by the US Food and Drug
Administration (FDA) (Barboza et al. 2001).

8.4  B
 auhinia forficata subsp. pruinosa (Vogel) Fortunato
& Wunderlin (Fabaceae)

Synonymy  Bauhinia pruinosa Vogel, Bauhinia candicans Benth., Bauhinia forfi-

cata Link var. pruinosa, Bauhinia forficata Link var. candicans, Pauletia candicans
(Benth.) A. Schmitz, Pauletia pruinosa (Vogel) A. Schmitz, Bauhinia forficata auct.
non Link, Bauhinia forficata auct. non Hook. & Arn.
204 8  Chaco and Espinal

Vernacular name  Pezuña de vaca, pata de vaca, caobá, mahogani, pata de buey,
pata de chivo, pata de toro, falsa caoba, caoba falsa, caoba del país, palo buey, torop,
mororó do sertão (Portuguese).

General information  The natural habitat of this plant is South of Brazil, Uruguay,
and North of Argentina, in soils rich in organic matter, being originally from Asia.
B. forficata is a quite-resistant tree, highly tolerant to drought. Optimal growth is
attained in sunny temperate climate. It is cultivated as ornamental. In Argentina it
grows in the provinces of CAT, JUJ, LRI, MEN, SAL, SJU, and TUC. Blooming is
in summer.

Botanical characteristics  B. forficate is an annual tree, 4–8 m high, and trunk has a
fissured brown-grayish bark. The crown is globular and irregular, with hairy and
thorny pubescent branches. Leaves are alternate, light green, and divided in two
oblong lobes resembling the hoof of a cow. The white flowers are hermaphrodite,
solitary, or arranged in axillary clusters (Fig. 8.6). The fruit is a legume with several
flattened, oval, blackish seeds (Juárez de Varela 2001; Garibotti Lusa and Bona 2009).

Ethnobotanical information  In Argentinean folk medicine, B. forficata has sev-

eral uses. The leaf is considered as antidiabetic, astringent, diuretic, expectorant,
antinephritic, antitussive, antihemorroidal, abluent, and blood depurative. The flow-
ers are considered as antihemorroidal and digestive, and they are externally used
against dandruff (Barboza et al. 2009).

Chemical data  The dried leaf have 3,7-di-O-a-rhamnopyranosylquercetin, 3,7-di-­

O-a-rhamnopyranosyl kaempferol (kaempferitrin), astragalin, bauhinoside, guani-
dine, quercitrin, rhamnose, rutin, saponins, and trigonelline. Leaves and flowers have
sitosterol, campesterol, stigmasterol, cholesterol, stigmasta-3,5-dien-7-one, choline,
trigonelline, sitosterol 3-O-β-glucoside, sitosterol 3-O-β-D–xylopyranoside, and

Fig. 8.6  Bauhinia forficata subsp. pruinosa (Vogel) Fortunato & Wunderlin (Fabaceae). Flower
(a) personal collection M C Fernández and leaves (b) MA Alvarez personal collection
8.4  Bauhinia forficata subsp. pruinosa (Vogel) Fortunato & Wunderlin (Fabaceae) 205

Fig. 8.7  Chemical compounds of Bauhinia forficata subsp. pruinosa (Vogel) Fortunato &
Wunderlin (Fabaceae)

sitosterol 3-Oα-D-xyluronofuranoside (Fig.  8.7) (Barboza et  al. 2009; Castro dos
Anjos et al. 2013; Toloza-Zambrano et al. 2015).

Biological activities  The extracts from dried leaves have been used as hypoglyce-
mic, antioxidant, hypocholesterolemic, diuretic, antimicrobial, and antinematode
(Sertié et al. 1990; Silva et al. 2002, Menezes et al. 2004; Arigony 2005). Also, it is
referred a cardiovascular (tachycardia) and calmative effect. The antibacterial activ-
ity of stems and fruits containing seeds was reported. The aqueous, ethanolic, and
butanolic leaf extracts and kaempferitrin were effective against the oxidative dam-
age resulting from hyperglycemia (Jorge et al. 2004; Sousa et al. 2004) supporting
the main use of B. forficata tea in folk medicine to treat diabetes mellitus. The
mechanism of action was hypothesized to be related to the inhibition of insulin
catabolism, reabsorption of glucose in the kidney, and/or stimulating glucose uptake
by peripheral tissues and not by stimulating insulin secretion, nor by the inhibition
of glucose absorption at intestinal level. Kaempferol did not promote glycemia
reduction; hence, the authors concluded that the aglycone-bound sugars are essen-
tial for the antidiabetic activity (Cazarolli et al. 2008).
206 8  Chaco and Espinal

More recently, Ferreres et al. (2012) demonstrated that the glycemia reduction
promoted by B. forficata resulted from the inhibition of α-glycosidase, the enzyme
responsible for catalyzing the final process in the digestion of carbohydrates. Such
action was ascribed to the presence of the quercetin and kaempferol derivatives of
the species (Tadera et al. 2006; Silvestre Marques et al. 2013). The hypoglycemic
activity was tested in vitro and in vivo. The aqueous extract (80 g l−1) was adminis-
trated to normotensive (5, 10, 20, 40 mg kg−1; i.v.) and hypertensive-induced (200
and 400 mg kg−1) male Wistar rats (Goldblatt et al. 1934). The oral acute treatment
(400 mg kg−1) induced blood pressure reduction (19%) without modifying heart rate
(Castro dos Anjos et al. 2013). Studies made in euglycemic mice treated with the
aqueous extract (7 mg kg−1) were also successful (Martínez et al. 2009). The effect
of the infusion on oxidative stress and liver damage in streptozotocin (STZ)-induced
diabetic mice (313  mg  kg−1) was tested by the determination of the activities of
ǖFF;-aminolevulinate dehydratase (ǖFF;-ALA-D), superoxide dismutase (SOD),
and catalase (CAT) in the liver and NF-E2-related nuclear factor erythroid-2 (Nrf2),
NADPH quinone oxidoreductase 1 (NQO-1), and heat shock protein 70 (HSP70) in
the liver and in the pancreas. These enzymes related to high levels of oxygen reac-
tive species (ROS) are usually increased in diabetes. The treatment did not change
the increased liver/body weight ratio and AST, ALT, and ǖFF;-ALA-D enzyme level
characteristics of diabetic rats. Also, Nrf2, NQO-1, and HSP70 protein levels in the
liver were not altered. On the contrary, the treatment attenuated the reduction of
CAT and the increase of pancreatic NQO-1 (Fernandes Salgueiro et  al. 2016).
Similar results were obtained in alloxan-diabetic rats with the aqueous, ethanol, or
hexane extracts (200, 400 mg kg−1) (de Sousa Lino et al. 2004).
Diabetic patients have erythrocytes with increased lipid peroxidation. It was
studied the effect of B. forficata leaf tea on oxidative damage and antioxidant mark-
ers in an in vitro experimental model of hyperglycemia in human erythrocytes. A
significant DPPH scavenging activity, the reduction of the deoxyribose degradation
induced by the Fenton reaction, an increase of thiobarbituric acid reactive sub-
stances (that reflect the lipid peroxidation), and a decrease in nonprotein SH levels
in human erythrocytes incubated with high-glucose concentrations were found.
Those results were attributed to the high level of phenolics and flavonoids acting as
DPPH+ radical scavenger and as iron chelators (Salgueiro et al. 2013).
The effects of oral treatment with a decoction of B. forficata on characteristic
metabolic parameters of streptozotocin (STZ)-diabetic and nondiabetic rats was also
reported (Pepato et al. 2002). After 31 days of chronic treatment with B. forficata
(mean daily dose 35.29/7.8 ml−1/100 g−1 body weight), the diabetic group showed a
significant reduction in plasma glucose, urinary glucose, and urinary urea levels
without any alterations in body weight, water or food intake, urinary volume, serum
lipid, or hepatic glycogen either in nondiabetic or STZ-diabetic rats. Those results
suggested that the antidiabetic activity is the result of inhibition of gluconeogenesis
and a reduction of counterregulatory hormones similar to biguanides. It was also
seen a sharp reduction in epididymal and retroperitoneal adipose tissue in the STZ-
diabetic rats as compared to the nondiabetic rats (Pepato et al. 2002). The ethanolic
8.4  Bauhinia forficata subsp. pruinosa (Vogel) Fortunato & Wunderlin (Fabaceae) 207

extract (40 mg l−1) interfered with hepatic glycolysis, glycogenesis, and lactate pro-
duction from L-glutamine. In vivo, a dose of 300 mg l−1of ethanolic extract produced
a minimal increase in the liver glycogen content in STZ-diabetic rats, but it did not
restore them to control levels. The treatment did not change glycemia (Olivera
Gomes et al. 2016). A diuretic action was also observed, and no signs of toxicity
were found (Toloza-Zambrano et al. 2015).
The continuous intake of a decoction (50  g  l−1) by male Wistar rats (150  ml
decoction per 100 g feed) resulted in a modification of the exploratory condition and
a decrease in  locomotor activity without changes in the glycemic index (Gellert
Moranza et al. 2017).
Antioxidant, anticoagulant, and antifibrinogenolytic activities against Bothrops
jararacussu Lacerda, 1884 (Ophidia, Viperidae) venom and its isolated thrombin-­
like serine protease enzyme were induced by the aqueous extract from B. forficata
(Damasceno et al. 2004; Sousa et al. 2004; Oliveira et al. 2005). The aqueous extract
neutralized Bothrops jararacussu and Crotalus durissus terrificus snake venom-­
induced coagulation, possibly by inhibition of serine protease enzymes involved in
blood coagulation disorders. There is also a significant reduction in the C. durissus
terrificus venom-induced edema, and Tityus serrulatus scorpion venom electrolyte
level changes; however, it was not able to counteract the glycogenolysis, plasma
catecholamines increase, and serum insulin change produced by the venoms
(Vasconcelos et al. 2004).
The proteinase activity of the acetone precipitate of B. forficate leaves corre-
sponded to the protein baupain (Andrade et al. 2011).
The antiulcer properties of an infusion of fresh leaves (150 g l−1) was demon-
strated in three different models of gastric ulcer: 0.3 M HCl: 60% EtOH (Mizui and
Doteuchi 1983), indomethacin/bethanechol (30  mg  kg−1, 5  mg  kg−1) (Rainsford
1978), and absolute ethanol induction (Morimoto et al. 1991). Moreover, the previ-
ous administration of the aqueous extract (100 mg kg−1) resulted in an increase of
gastric mucosa, possibly associated with the extract flavonol content (Costa Mazzeo
et al. 2015).
The ethanolic extract (250  mg  ml−1) had antimicrobial activity against
Streptococcus mutants (ATCC 9927) and S. sanguis (ATCC 10557) from oral cavity
(Batista de Souza et  al. 2014). The hexane, di-chloromethane, and ethyl acetate
extracts from leaves, stems, and barks evaluated for their activity against fungi and
pathogenic yeasts were not effective against Candida albicans, C. tropicalis,
Saccharomyces cerevisiae, Cryptococcus neoformans, Aspergillus flavus, A. fumig-
atus, A. niger. In contrast, the ethyl acetate extract of the stems was moderately
active against Epidermophyton floccosum (MIC < 750 mg ml−1); the hexane extract
from the bark was active against Microsporum canis, Trichophyton mentagrophytes,
T. rubrum, and Epidermophyton floccosum (MIC < 1000 mg ml−1); and the dichlo-
romethane extract from bark was active against T. rubrum and E. floccosum (MIC
500 μg ml−1) (Silva & Cechinel-Filho 2002).
HeLa cell cycle progression was inhibited by a compound isolated from B. forfi-
cate (HY52 = C17H30O2N2) (Lima et al. 2006).
208 8  Chaco and Espinal

Seeds contain a lectin (BfL), a glycoprotein with 6.24% carbohydrate content,

that has anticoagulant and antiplatelet aggregation properties (Silva et al. 2012).
Toxicity  B. forficata aqueous extract (4.65 g l−1) was tested against cyclophospha-
mide, a mutagenic chemotherapeutic agent, using the model of Wistar rat bone mar-
row cells from Rattus norvegicus. It was not detected chromosomal alterations or
modification in the mitotic index concluding that the aqueous extract of B. forficata
is not cytotoxic but showed an effective antimutagenic activity endorsed to its total
phenolic and flavonoid content (Düsman et al. 2013). On the other hand, the oral
administration of a decoction to rats did not show any modification of the toxicity
markers amylase (pancreas toxicity), creatine kinase (muscle toxicity), lactate dehy-
drogenase (muscle and liver toxicity), bilirubin (liver and biliary toxicity), and
angiotensin-converting enzyme (renal microcirculation and kidney toxicity) (Pepato
et al. 2004).
Dry extracts obtained using colloidal silicon dioxide at 125 and 250  mg  ml−1
showed hepatotoxicity signs (AST and alkaline phosphatase ALP increase) in
hyperglycemic rats after a 35-day oral treatment. A formation of toxic derivatives
resulting from the interaction between colloidal silicon dioxide and compounds
from the vegetable extract or modifications of the colloidal silicon dioxide caused
by the high temperature of the drying process were presumed responsible of that
effect (Pepato et al. 2010). No significant quantities of the toxic metals Pb, Cr, and
Ni were detected in B. forficata coming from different producers (Franco et  al.
2011).
In vitro cultures  Axenic plants were obtained from disinfected seeds in MS
(Murashige and Skoog 1962) culture medium with 30 g l−1 sucrose, 2.3 g l−1 agar,
and 4.0 mg l−1 BAP. Calli were initiated from hypocotyl explants on ½ MS medium,
containing 30 g l−1 sucrose, 2.3 g l−1 agar and supplemented with 3.0 mg l−1 BAP
concentration. Shoot buds appeared in media with 2.0 and 4.0  mg  l−1 BAP after
60 days in culture (Mello et al. 2000). Cell suspension cultures were established in
MS liquid medium with 30 g l−1 sucrose and 17.7 μM BAP (Mello et al. 2001).

8.5  C
 aesalpinia gilliesii var. gilliesii (Wall. ex Hook.) D. Dietr.
(Fabaceae)

Synonymy  Erythrostemon gilliesii Hook Klotzsch var. gilliesii, Poinciana gilliesii

Wall. ex Hook, Caesalpinia macrantha.

Vernacular name  Lagaña de perro, algarrobillo, piscala, barbón, barba de viejo,

barba de chivo, mal de ojos, flor del indio, chosme, espiga de amor, mal de ojos,
yellow bird of paradise, bird of paradise bush (Eng.), desert bird of paradise (Eng.),
uccello del paradiso (Italian).
8.5  Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.) D. Dietr. (Fabaceae) 209

General information  C. gilliesii is a native species to Argentina that is worldwide

cultivated (Osman et al. 2008). It grows sub-spontaneously in the Mesopotamian
region, Uruguay, Chile, and Southwest of North America. It is endemic to the Chaco
and Monte’s phytogeographic formations, and its center of dispersion seems to be
the province of COR (Burkart 1936). It grows in dry, stony soils, plains, or hillsides,
between 50 and 2500 m.a.s.l. in the provinces of BAI, CAT, CHA, COR, COS, ERI,
LPA, LRI, MEN, NEU, RNE, SAL, SDE, SFE, SJU, SLU, and TUC (Cocucci et al.
1992; Barboza et al. 2009). The flowering period lasts 5 months, from September to
January, and it is variable among populations and individuals. Pods and seeds are
poisonous. Like many members of the legume family, C. gilliesii can fix its own
nitrogen from the soil because of specialized bacteria living in root nodules.

Botanical characteristics  C. gilliesii is a shrub with perfect and staminate flowers,

arranged in variable numbers in simple clusters. Leaves are bi-pinnately compound
with small bluish-green leaflets that give the foliage a fernlike look. Branches are
pubescent and glandular, with reddish hairs mixed with single hairs (Macaya 1999).
Flowers are cyclic, pentameric, zygomorphic, and hermaphrodite with imbricated
pre-flowering. They are grouped into glomeruli; calyx have five hairy glandular
sepals; the hairless, yellow corolla has five petals, 2–3  cm long; the androecium
commonly has 10 free red stamens with filaments 2–3 times longer than the petals.
Anthers have two thecae, and pollen is usually tricolporate; the unicarpelar, super-
imposed, linear, unilocular gynoecium has filiform ovules in a row in the placenta
(Fig. 8.8). The fruit is an oblong and flat dehiscent pod with membranous consis-
tency, 6–9 cm length, with transverse grooves between the seeds; in maturity the

Fig. 8.8  Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.) D. Dietr. (Fabaceae). Buenos Aires
Botanical Garden. MA Alvarez personal collection
210 8  Chaco and Espinal

fruit separated into two twisted valves. Ovoid chestnut-olive seeds measure 10 mm
in length (Steibel and Troiani 2000). Nectar is found on the inner surface of the
hypanto, without a containing structure, always accompanying the vascular bundles
of the stamens, being secreted mostly at the beginning of the anthesis (Jausoro and
Galetto 2000; Jausoro and Galetto 2001). Pollination is zoophilic (entomophile or
ornithophile) (Viglierchio et al. 2000).

Ethnobotanical information  The dried flower and leaf infusion are febrifuge,
purgative, and against cavities and toothache. Washings made with flowers are used
for eye conditions and decoction of the aerial part to treat earaches (Agüero 2017).
The dried seed is taken as anthelminthic and insecticidal (Viglierchio et al. 2000).

Chemical data  The dried flowers contain phenyl alanine and arabinose. The flowers
contain β-sitosterol-3-O-butyl, sterol β-sitosteryl-3-O-glucopyranoside (daucos-
terol), isorhamnetin and isorhamnetin-3-O-rhamnoside, 12,13,16-trihydroxy-14(Z)-
octadecenoic acid, luteolin-7,4’-dimethyl ether, genistein-­ 5-methyl ether,
luteolin-7-O-rhamnoside, isovanillic acid, and p-methoxy benzoic acid (Fig.  8.9).
The seeds contain cesalin, glutamic acid, pipecolic acid, and 3-hydroxyproline
(Fig. 8.9). The leaves contain gossypetin-3-O-β-D-robinobioside, quercetin-3-O-β-
D-glucoside, quercetin-3-O-β-D-galactoside, rutin, kaempferol-3-­O-β-D-rutinoside,
and luteolin-7-O- β-D-glucoside (Suarez et al. 1984; Viglierchio et al. 2000; Osman
et al. 2008, 2013; Barboza et al. 2009; Emmam et al. 2019)

Biological activities  There are reports about antitumor activity, respiratory stimu-
lant effect, and smooth muscle relaxant activity (Barboza et al. 2009).
As for the antioxidant activity, scavenging activity was seen in extracts from
leaves (IC50 95.3), flowers (IC50 38.7), and pods (IC50 44.7 μg ml−1). The dichlo-
romethane extract of C. gilliesii flowers showed antioxidant activity
(IC50  =  45.5  μg  ml−1) comparable to standard rutin (24  μg  ml−1). Flower ethyl
acetate extract had the highest radical scavenging activity (IC50 = 7.43 μg ml−1),
even higher than rutin (Osman et al. 2013). Isolated isorhamnetin and isorhamnetin-­
3-­O-rhamnoside possess moderate antioxidant activity with IC50 values of 370 and
372.5  μg  ml−1, respectively. On the other hand, β-sitosterol-3-O-butyl and
­daucosterol possess variable prooxidant activity against DPPH radicals. Such an
antioxidant activity was attributed to the presence of phenolic compounds as tannins
and flavonoids (Ulubelen et al. 1967; Osman et al. 2008, 2016).
The methanol extract counteracted the liver injury induced by carbon tetrachlo-
ride (CCl4) in mature male albino Wistar rats (reduction of ALT and AST and
increase of TAC) protecting the liver architecture at a dose of 300 mg kg−1. The oral
administration of the leaf methanol extract significantly diminished carrageenan-­
induced paw edema in rats (100 and 200 mg kg−1), exhibited substantial gastropro-
tective properties, and diminished ulcer number and ulcer severity in a dose-dependent
way. Those activities were attributed to the presence of antioxidant polyphenolics
and phytosterols (Osman et al. 2016; Emmam et al. 2019).
8.5  Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.) D. Dietr. (Fabaceae) 211

Fig. 8.9  Some chemical compounds from Caesalpinia gilliesii var. gilliesii (Wall. ex Hook.)
D. Dietr. (Fabaceae)

It was reported the green synthesis of metallic silver nanoparticles (AgNPs)

containing C. gilliesii leaf extracts. The AgNPs were synthesized by the reduction
of 10 ml of AgNO3 solution (1 mM) with different concentrations of C. gilliesii
hydroalcoholic extract (100 μl to 300 μl, stock solution 0.04 g extract 10 ml−1 sol-
vent). The antimicrobial activities of C. gilliesii extract and AgNPs were tested
against Gram-positive and Gram-negative bacterial pathogens and yeast.
Synthesized C. gilliesii extract-AgNPs showed highest activity against all tested
pathogens than the hydroalcoholic extract (Scott  1989, Emam et  al. 2017). The
cytotoxicity of AgNO3, C. gilliesii hydroalcoholic extract, and AgNPs was evalu-
ated against both normal skin fibroblast (BJ-1) and breast cancer cell line (MCF-7).
Cytotoxicity against MCF-7 was 4.2% for the hydroalcoholic extract and 96.5%
for the c­ ombination of extract with AgNPs. They also produced cytotoxicity against
BJ-1 (33.3% and 70.5%) at 100 μg ml−1, respectively. AgNO3 showed high cytotox-
icity ≥90% at 12.5 μg ml−1 (Emmam et al. 2017).
The aqueous extract from leaves, flowers, and pods (60 g 500 ml−1) was fraction-
ated using solvent-solvent extraction with n-hexane, methylene chloride, ethyl ace-
tate, and n-butanol and tested on Wistar male albino rats (1, 2, 3, 4, and 5 g kg−1) to
study the protective effect on CCl4-induced hepatotoxicity (Osman et  al. 2013).
Aspartate aminotransferase (AST), alanine aminotransferase (ALT), and alkaline
phosphatase (ALP) blood levels showed a significant protection of the hepatic cells.
Rats pretreated with leaves or flower extracts of C. gilliesii showed normal hepato-
cytes with minimum fatty changes, portal vain congestion, and mild inflammatory
212 8  Chaco and Espinal

cell infiltration around portal vein. Pretreatment with pod extract showed mild pro-
tection against the toxic effects of CCl4 on the liver cells.
Toxicity  Total extract of the leaves, flowers, and pods was tested for their cytotoxic
against different cancer cell lines by SRB (sulforhodamine B) cell viability assay
(Skehan and Storeng). The flower extract exhibited a potent inhibition of cell growth
in different cancer cell lines (MCF7, HepG2, HCT116, HeLa, and PC3 cells) with
IC50 values of 10 and 15.6 μg ml−1, respectively. Also the extracts from leaves and
pods were cytotoxic. The dichloromethane extract and the isolated steroidal com-
pounds were tested for their cytotoxicity against MCF7 and HepG2 cells. The most
active compound against HepG2 and MCF7 cells was β-sitosterol-3-O-butyl, and
IC50 was 13.1 and 14.4 μg ml−1, respectively. Doxorubicin showed IC50 values of
3–4  μg  ml−1 in the same cell lines. These findings indicate that β-sitosterol-3-O-
butyl and the other phytosterols are responsible for the cytotoxicity of the extracts
which would correlate with the known abortifacient, antimalarial, and anthelmintic
properties. Galactomannan isolated from the aqueous extract from seeds revealed a
concentration-dependent potency to induce cell death of HepG2 cells with IC50
value of 0.375 μg ml−1 (Abdel-Megeed et al. 2018).
Cesalin, an antitumor and antineoplastic protein isolated from C. gilliesii, inhib-
ited carcinosarcoma growth by binding to its plasma membrane (Elting and
Montgomery 1979).

8.6  Pilocarpus pennatifolius Lemaire (Rutaceae)

Synonymy  Pilocarpus selloanus Engl., Pilocarpus pennatifolius Lem. var. selloa-

nus, Pilocarpus selloanus Engl. var. gracilis, Pilocarpus selloanus Engl. f. brevi-
pedicellata, Pilocarpus pennatifolius Lem. var. genuinus, Pilocarpus pennatifolius
Lem. f. typicus, Pilocarpus pennatifolius Lem. f. gracilis, Pilocarpus pennatifolius
Lem. f. latifoliolatus, Pilocarpus pennatifolius Lem. f. brasiliensis, Pilocarpus pen-
natifolius Lem. f. intermedius, Pilocarpus pennatifolius Lem. f. paraguariensis.

Vernacular name  jaborandí, jaborandí del Paraguay, cutia, yaguarundí, camba,

canilla, yvyrá tai.

General information  P. pennatifolius grows in the provinces of CHA, COS, FOR,

and MIS.

Botanical characteristics  P. pennatifolius is an evergreen shrub about 6 m high

with alternate, imparipinnate leaves with ribbed petiole and rachis, 2–3 leaflet pairs,
narrowly elliptic or oblong, obtuse and emarginated at apex, a revolute margin.
Flowers are 5-merous, 9–11 cm diameter, numerous, disposed in long axillary and
terminal racemes, subsessile, pendulous, purple, and with a very small teethed
8.6  Pilocarpus pennatifolius Lemaire (Rutaceae) 213

calyx, and the petals are deciduous and reflexed when flowering. Stamens are pur-
ple, and ovary is superior, with carpels inserted on gynophores. The fruit is divided
into mericarps united at the base, truncate, and mucronate at the apex (Miranda
et al. 2000; Spegazzini et al. 2002).

Ethnobotanical information  The Mbyá native community referred the use of the
infusion made with the whole plant to treat diarrhea and parasitosis. The guaraníes
used it against vomiting and intestinal parasitosis and as diaphoretic. The infusion
of leaves, dried leaves, and extracts is used in popular medicine for the treatment of
asthma, diabetes, liver diseases, and in the form of cosmetic for hair treatment. The
dried leaves possess gastrostimulant, peristaltic, and secretagogue activities, suit-
able for the treatment of urinary, gastrointestinal, and pulmonary disorders. A not
specified part is referred with activity as odontalgic, abortive, and to treat ear disor-
ders. Also it is used as repellent, acaricide, and insecticide (Schultz 1997; Martínez
Crovetto 1981; Rodríguez Barbosa 1985; Müller 1997; Pochettino et  al. 2003;
Barboza et al. 2009; Ferreres et al. 2015)

Chemical data  The stem contains pilocarpine and isopilocarpine. The EOs from
stems contain 2-tridecanone, 2-pentadecanone, caryophyllene oxide, germacrene D,
E-β-ocimene, Z- β-ocimene, δ-cadinene, β-cubebene, γ-muurolene, selinene,
β-elemene, bicyclogermacrene, α- humulene, α-copaene, β-bourbonene,
β-caryophyllene, and spathulenol, among others (Fig.  8.10) (Lucio et  al. 2002;
Santos and Hrihorowitsch Moreno 2004; Barboza et al. 2009)

Fig. 8.10  Chemical compounds from Pilocarpus pennatifolius Lemaire (Rutaceae)

214 8  Chaco and Espinal

Biological activities  It is well known the activity of pilocarpine, an imidazole alka-

loid extracted from the leaves, as a direct cholinergic, stimulating the parasympa-
thetic system. It is used for the treatment of glaucoma and also for the stimulation
of sweat and lachrymal glands (Goodman and Gilman 2001; Valdez et al. 1993).
As for the study of its antimicrobial activity, leaves (2 kg) and stem bark (98 g)
(both dried and crushed) were extracted with methanol. Subsequently, the aqueous
residue was thoroughly extracted with ethyl ether rendering acidic and basic ether
fractions. The acid stem bark solution was successively extracted with ethyl ether
and ethyl acetate. Antibacterial and antifungal assays were performed with the
crude leaf methanol extract (CLME), crude bark methanolic extract (CBME), leaf
acidic ether fraction (LAEF), and leaf basic acetate fraction (LBAF) and with the
isolated compounds.
The fractions showed activity against the bacteria tested, the most sensitive was
Shigella flexneri (MIC 7.8 μg ml−1), being the CBME and LAEF fractions actives
also against Candida krusei (MIC 15.6  μg  ml−1 and 31.2  μg  ml−1, respectively).
Some of the isolated compounds were active against Shigella flexneri (MIC
3.12  μg  ml−1), Salmonella typhimurium (MIC 12.2  μg  ml−1), Enterococcus spp.
(MIC 25 μg ml−1, 50 μg μg ml−1), E. faecalis (MIC 1.56 μg ml−1), and Salmonella
enteritidis (MIC 1.56 μg ml−1) (do Carmo et al. 2018).
The leaves extract obtained by exhaustive maceration (200 mg ml−1) and its six
sequenced serial 1:2 dilutions (starting at 20,000 ppm) were tested for their in vitro
activity against the ectoparasites Ctenocephalides felis felis, Rhipicephalus san-
guineus, and R. microplus. No insecticidal activity was observed after 24 h of treat-
ment. However, insecticidal activity was observed from 1250 ppm after 48 hours. A
drop in tick’s weight was observed with concentrations between 625 and 10,000 ppm.
Hatching and reproductive efficiency of ectoparasites were also lower. The most
effective concentration was 10,000 ppm (59.06% efficiency) being the extract more
effective as an anti-tick than as an insecticide. As an anti-tick, it was more effective
against R. sanguineus than against R. microplus. The insecticidal activity was asso-
ciated to 2-pentadecanone (de Sousa Oliveira Batista et al. 2013).
A tea prepared with 3 g plant material per 500 ml distilled water (100 °C, 20 min),
a commercial sample (CS) (6-methoxy-quercetin-3-O-robinobioside), and an
authenticated sample (BGS) (hesperetin-7-O-rutinoside) were tested for their
in  vitro ability to inhibit acetyl-cholinesterase (AChE), butyl-cholinesterase
(BuChE), and α-glucosidase. Inhibition of α-glucosidase and cholinesterase was
dose-dependent in all cases. The two samples were similarly active against AChE
(IC25 872.02 and 1056.03 μg ml−1 for BGS and CS, respectively) and α-glucosidase
(IC50 604.34 and 573.48 μg ml−1 for BGS and CS, respectively). On the other hand,
a stronger capacity to inhibit BuChE was observed for BGS (IC50 1112.14 μg ml−1)
than for CS (IC50 1743.96 μg ml−1) (Ferreres et al. 2015).
In vitro culture  Calli and cell suspension cultures were established from leaves on
MS medium supplemented with 5 mg l−1 2,4-D and 1.0 mg l−1 kin. Both callus and
cell suspension cultures produce the alkaloid pilocarpine at a concentration of
1.01 μg g−1 and 0.045 μg g−1 at day 30 and 19 of culture, respectively (Andrade
Salles et al. 2004).
8.7  Jodina rhombifolia (Hook. & Arn.) Reissek (Cervantesiaceae) 215

8.7  J odina rhombifolia (Hook. & Arn.) Reissek

(Cervantesiaceae)

Synonymy  Celastrus rhombifolius Hook. & Arn., Ilex cuneifolia L. var. bonarien-
sis, Celastrus jodina Steud., Jodina ruscifolia Hook. & Arn., Jodina cuneifolia
Miers, Jodina bonariensis Kuntze.

Vernacular name  Quebracho flojo, peje, sombra de toro, quebrachillo.

General information  J. rhombifolia is a native species from Bolivia, Brazil,

Paraguay, Uruguay, north and center of Argentina. It is one of the eight species that
conform the forest of tala (C. spinosa) in the Talares district. Its soft wood is used
for construction, yokes construction, and axles of carts; its bark is used to clean
rawhide harnesses (Pérez Meroni et al. 2010). It is also used as an ornamental in
gardens and to form defensive fences (Cabrera and Zardini 1978; De la Peña and
Pensiero 2004; Dimitri 2004). It is also one of the adulterants of yerba mate (Lahitte
et al. 1999). Blooming is from January to July. Pollination is entomophile.

Botanical characteristics  J. rhombifolia is a tree of medium to small texture, 3–5

meters high, with a globose, open, and irregular crown. The bark is thick and
reddish-­brown. Leaves are alternate with a rhomboidal shape, hard, with spines on
the apices and lateral vertices, coriaceous, bright, dark green (Fig. 8.11). The ­flowers

Fig. 8.11  Jodina rhombifolia (Hook. & Arn.) Reissek (Cervantesiaceae). Buenos Aires Botanical
Garden. MA Alvarez personal collection
216 8  Chaco and Espinal

are bisexual, formed by 5 floral pieces arranged in axillary groups of about 7–8
­yellowish green little fragrant flowers. The fruit is a pseudodrupe, rough, brownish-­
green with a modified nectary disc (white fleshy layer). During the development of
the fruit, the persistent petals became redish. Seeds are subspherical, flattened, top
depressed, and chestnut clear of approximately 5 mm in diameter (Luna et al. 2017).

Ethnobotanical information  Leaf and bark preparations are used in folk medicine
as antidiarrheal, antidysenteric, anti-inflammatory, antitussive, abortive, and hypo-
tensive, for sore throats, kidney stones or pain, stomach indisposition, and articular
and muscular pain. The leaves are used against alcoholism, as digestive, and for
carcinoma. Tisanes made with leaves and branches are taken to treat stomach ail-
ments and colds. The decoction is used as astringent and antidysenteric; the leaves
are used to heal wounds and to cure nasal polyps. The infusion of leaves is drunk as
digestive, hepatic, antihidrotic, pectoral, antiasthmatic, and anti-alcoholic. The
decoction of the ingested bark is taken against dysentery (Ratera and Ratera 1980;
Toursarkissian 1980; Martínez Crovetto 1981; Adzet et  al. 1991; Filipov 1994;
Demaio et al. 2002; Davicino et al. 2007). Hieronymus (1882) attributed to the oil
extracted from fruits the property of healing buboes and venereal sores. In the
Buenos Aires-La Plata conurbation, J. rhombifolia leaves are marketed as having a
cholesterol-lowering effect (Hurrel et al. 2015).

Chemical data  The seed oil contains 9-octadec-ynoic acid (stearolic acid), trans-­
10-­heptadecen-8-ynoic hydroxytransl (-heptadecen-8-)-ynoic acid, trans-­10,16-­
heptadecadien-­ 8-ynoic acid, 7-hydroxytrans-10,16-heptadecadien-ynoic acid,
trans-ll-actadecen-yn oic acid (ximenynic acid), 8-hydroxytranslloctadecen-ynoic
acid, trans-ll, 17-octadecadien-9-ynoic acid, 8-hydroxytransll, 17-octadecadien-­9-
ynoic acid, 9-hydroxystearic acid, and 9,10-epoxystearic acid. The leaves contain
phenolic compounds, organic acids, tannins, flavonoids, steroids, gum, and muci-
lage. The aerial parts contain steroids, triterpenoids, alkaloids, coumarins, saponins,
vicenin-2, vitexin, orientin, and swertisin (Fig. 8.12) (Spitzer et al. 1994; Barboza
et al. 2009).

Biological activities  The activity of the aqueous (250–750 mg kg−1) and hydroal-

coholic extracts (250–750 mg kg−1) from the aerial parts were tested against HCl/
ethanol-induced gastric lesions modified model (Yesilada et  al. 1997). Female
Wistar rats, pretreated with the aqueous extract (10 mg kg−1), had gastric lesions in
all the cases. On the contrary, animals treated with 250 and 750  mg  kg−1 of the
hydroalcoholic extract had gastric ulcers in 75% and 62.5% of cases, respectively.
It was postulated that the antiulcer activity was related to the presence of the
C-glycosyl flavonoids vicenin-2, vitexin, isovitexin, orientin, and swertisin
(Montanha et al. 2009).
As for the diuretic activity of aerial parts, when methanolic extracts of leaves,
bark, phloem, and branches were administered to Wistar rats (125, 250, and
500 mg kg−1of weight) in normal saline solution, a diuretic activity was observed in
all cases. The more potent was the 250  mg  kg−1 dose of leaf methanolic extract
8.7  Jodina rhombifolia (Hook. & Arn.) Reissek (Cervantesiaceae) 217

Fig. 8.12 Chemical compounds found in Jodina rhombifolia (Hook. & Arn.) Reissek,
(Cervantesiaceae) aerial parts

(diuretic index = 2.01), followed by 500 mg kg−1 dose of leaf methanolic extract

(diuretic index = 1.98). The minimal diuretic activity was observed with the bark
methanolic extract at all concentrations tested (Teves et al. 2018).
The anti-alcoholic activity of J. rhombifolia lyophilized aqueous extract was
tested in male Wistar rats. Rats received the extract twice daily (1 ml 200 g−1), one
of the doses of extract (125, 250, and 500 mg kg−1 weight) for 10 consecutive days.
The daily ethanol intake fall was evidenced from the first day of treatment and per-
sisted throughout the entire treatment period. The daily water intake neither the
body weight gain was affected; however, daily food intake was higher in the group
of treated rats (Teves et al. 2015a). When the treatment (62.5, 125, 250 mg kg−1) was
applied to adolescent Wistar rats (postnatal day 29; 83–105 g of weight), it mark-
edly reduced ethanol voluntary intake on a dose-dependent base (Teves et al. 2015b).
Repeating administration of Jodina rhombifolia lyophilized extract markedly
reduced alcohol voluntary intake in rats in a self-administration model treated dur-
ing 10 consecutive days, without signs of tolerance or apparent side effects (Teves
et al. 2015a, b, Wang et al. 2016).
Toxicity  There are no trials that demonstrate toxicity (Alonso and Desmarchelier
2015).
218 8  Chaco and Espinal

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Chapter 9
Patagonia

9.1  Introduction

9.1.1  The Steppe and the Andine Forest

Patagonia Argentina is an extensive region (Fig.  9.1) located to the South of the
American continent that includes the provinces of the LPA, NEU, RNE, CHU, SCR,
TDF, Argentina, Antarctica, and South Atlantic Islands. The borders of the region
are to the North in East-West direction, the line that integrates the mouth of the
Colorado River into the Mar Argentino, the channels of Rio Grande and Rio
Barrancas that are the border between the provinces of NEU and MEN, until the
limit with the Chilean Patagonia and the mountainous formation of the Cordillera
de los Andes. Towards the South end, it is the Cape Horn in the Drake Passage that
is the boundary between the Atlantic and Pacific Oceans, and to the East is the Mar
Argentino in the South Atlantic Ocean.
Ecologically, Patagonia could be divided into the cordillera de los Andes sector,
which is more humid and has forests and a great variety of lakes of glacial origin,
and the Patagonia desert or Patagonian steppe with semiarid characteristics and
predominance of shrubs.
The highest peak in Patagonia is the Domuyo Volcano (4709 m.a.s.l), and the
lowest point is the Laguna del Carbón (108 m.b.s.l.). This area has numerous volca-
noes such as Catedral, Piltriquitrón, Chapelco, Caviahue, Bayo, and La Hoya.
Binational volcanoes (Argentina-Chile) are Lanin, Copahue, Pantojo, Tronador,
Fitz Roy, Cerro Torre, Murallón, and San Lorenzo. Some of the lakes of the region
are Nahuel Huapi, Puelo, Argentino, Strobel, Futalaufquen, Traful, and Mascardi,
and the important rivers are Colorado, Negro, Chubut, Deseado, Santa Cruz,
Gallegos, and Limay. There are also glaciers such as the Perito Moreno and
Vespignani.

© Springer Nature Switzerland AG 2019 227

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2_9
228 9 Patagonia

Fig. 9.1  Patagonia Argentina. (a) Ushuaia, Vespignani glacier viewpoint, (b) Patagonian steppe,
(c) Perito Moreno Glacier, (d) MA Alvarez personal collection

There are wide temperature amplitude with a difference between the North and
the South of the region. The cordillera de los Andes retains humidity from the
Pacific Ocean, beneficial for the Patagonian forest; on the other hand, the arid
Patagonian steppe is crossed by dry winds.
As for the economic activities, Patagonia has natural forest resources (Nothofagus
pumilio, cypresses, and Lomatia hirsuta) that have to be preserved. The agricultural
produce includes apples, pears, cherries, fodder, potatoes, peaches, and vegetables.
9.1 Introduction 229

Livestock includes breeding of bovine, sheep, and goat cattle. Mining includes
deposits of gold, silver, gas, oil, coal, lead, and uranium, among others. Sea prod-
ucts include silverside, anchovy, haddock, shrimp, hake, king crab, crustaceans,
rainbow trout, and Pacific salmon. Also, Patagonia produces hydroelectric power.
As for the fauna, the marine fauna includes southern right whale, orca, dolphins, sea
lions, and sea elephants. Among the birds, in the oceanic coast, there are the
Magallanes penguin, cormorants, seagulls, and Antarctic dove. In the Andean
Patagonia, the most witnessed birds are raptors such as the condor. The South area
has foxes, wild cats, Patagonian hares, armadillos, Pampean deer, huemul, puma,
guanaco, and ñandú, among others.
As for the flora, in the sea coast of the provinces of RNE and CHU, there are
grasslands, shrubs, and trees. The central region of Patagonia has steppe vegetation
with more sparse grassland, mounts of Larrea spp., Mulinum spinosum, and
Festuca gracillima. However, this region is going through a desertification process
as a consequence of livestock grazing (sheep and goats) and the effect of volcanic
ashes. Small oases, with Salix humboldtiana, and other species are located in the
riverbanks from the central plateau; in the mountain ranges and the Andean
Patagonia forest, there are coniferous, Nothofagus pumilio (lenga), N. antarctica
(ñires), and N. dombeyi (coihue), shrubs such as Berberis darwinii (michay) and B.
microphylla (calafate), ferns and mosses in muddy grounds, and fungus such as
Cyttaria hariotii (llao llao) growing in the coihue bark. Among the medicinal spe-
cies from Patagonia are Acantholippia seriphioides (A.  Gray) Moldenke
(Verbenaceae), Adesmia boronioides (Fabaceae), Adiantum chilense (Pteridaceae),
Apium panul (Bertero ex DC.) (Apiaceae), Araucaria araucana Molina K. Koch
(Araucariaceae), Arjona tuberosa (Santalaceae), Atriplex lampa (Moq.) D. Dietr.
(Chenopodiaceae), Baccharis crispa Spreng. (Asteraceae), Baccharis sagittalis
(Asteraceae), Berberis darwinii Hook. (Berberidaceae), Berberis heterophylla
(Berberidaceae), Buddleja globosa (Buddlejaceae), Colliguaja integerrima Gillies
& Hook. (Euphorbiaceae), Coriaria ruscifolia L. Deu (Coriariaceae), Chenopodium
ambrosioides (Chenopodiaceae), Cyttaria darwinii (Cyttariaceae), Equisetum
bogotense (Echicetaceae), Empetrum rubrum (Empetraceae), Ephedra ochreata
(Ephedraceae), Euphorbia serpens (Euphorbiaceae), Fabiana imbricata
(Solanaceae), Festuca argentina (Poaceae), Fragaria chiloensis (Rosaceae),
Fuchsia magellanica Lam. (Onagraceae), Grindelia chiloensis (Cornel.) Cabrera
(Asteraceae), Gunnera tinctoria (Gunneraceae), Larrea divaricata Cav.
(Zygophyllaceae), Lycium chilensis (Solanaceae), Maytenus boaria (Celastraceae),
Mulinum spinosum (Cav.) Pers. (Apiaceae), Nothofagus antarctica (G.  Forst.)
Oerst (Nothofagaceae), Nothofagus betuloides (Nothofagaceae), Oxalis
enneaphylla Cav. (Oxalidaceae), Plantago lanceolata L. (Plantaginaceae),
Prosopis denudans (Fabaceae), Ribes magellanicum (Grossulariaceae), Rubus
geoides Sm. (Rosaceae), Rumex crispus (Polygonaceae), Salix humboldtiana
(Salicaceae), Clinopodium darwinii (Benth.) Kuntze (Lamiaceae), Schinus
marchandii (Anacardiaceae), Senecio filaginoides (Asteraceae), Urtica dioica
(Urticaceae), Usnea barbata (Usnaceae), Valeriana carnosa (Valerianaceae), and
Viola magellanica (Violeaceae) (Díaz 2010; Toledo and Kutschker 2012; Molares
and Ladio 2015).
230 9 Patagonia

9.2  Adesmia boronioides Hook. f. (Fabaceae)

Synonymy  Patagonium boronioides (Hook. f.) Kuntze.

Vernacular name  Paramela, yagneu, lonckotrevo, té pampa, té silvestre, yerba car-

melita, and éter.

General information  A. boronioides is an aromatic and medicinal plant native to

Argentina and Chile. In Argentina, it is found in the provinces of CHU, NEU, RNE,
SCR, and TDF; it inhabits sites with little irrigation. It is widely valued by the origi-
nal Patagonian peoples that have used it both as medicinal and in rituals. Some
mapuche-tehuelche communities cultivated and protected the species. The aerial
parts have a large amount of resin and a sweet-woody licorice-spicy odor. The EO
has become an important economic resource for the fragrance industry. The leaves
are used as cattle sheep and horse fodder (Burkart 1967; Ulibarri and Burkart 2000;
González et al. 2016; Ladio and Molares 2017).

Botanical characteristics  A. boronioides is a medium-sized shrub reaching 1 m in

height, very branched, with glandular ascending branches, fragrant and very resin-
ous. Leaves are petiolate, with numerous elliptical leaflets (10–20). Leaflets have an
irregular margin (smooth-wavy). Scented flowers are arranged in terminal clusters.
The calyx is glandulous and pubescent, with tooth equal in length to the calyx tube.
The corolla is yellow, with a heart-shaped banner. The fruit is a legume with 3–5
strangulations (Peltz 1962; Dimitri 1974; Correa 1984; Nájera et  al. 2000; Vidal
2007).

Ethnobotanical information  The aerial organs, fresh or dried, are used as mastica-
tory and to prepare infusions against rheumatic pain, cold, alopecia, and gastrointes-
tinal and renal problems, to treat burns and bruises, and to relieve menstrual
discomforts (Montes and Peltz 1963; Martínez Crovetto 1980; Montes and
Wilkomirsky 1985; Silva et al. 2004; Backhouse et al. 2008). Also, it is considered
having bronchodilator properties and aphrodisiacs (Montes and Wilkormirsky 1985;
Nájera et al. 2000).

Chemical data  The EO have a high percentage of bisnorsesquiterpenes as esquel-

6-en-9-one and esquel-7-en-9-one. Also, it has α-copaen-11-ol, δ-cadinene, 10-epi-­
γ-eudesmol, 4α-hydroxydihydroagarofuran, 1-epicubenol, guaia-6,9-diene, and
furopelargones A and B (Fig. 9.2). The relative proportion of the main compounds
was directly related to the season in which the plant material was collected. Autumn
appeared as the optimal harvest time for industrial purposes of this species (González
et al. 2002, 2004; Barboza et al. 2009; Gastaldi et al. 2016).

Biological activities  The antioxidant activity (DPPH and BCB assays) was evalu-
ated in a 5% infusion (Backhouse et al. 2008; Gastaldi et al. 2016, 2018). The polar
9.2  Adesmia boronioides Hook. f. (Fabaceae) 231

O
HO
OH

α-copaen-11-ol 4α-hydroxydihydroagarofuran guaia-6,9-diene

HO

δ-cadinene 10-epi eudesmol esquel-6-en-9-one

O

O
O
O O

furopelargone A furopelargone B esquel-7-en-9-one

Fig. 9.2  Chemical compounds from Adesmia boronioides Hook. f.

fractions (methanol and aqueous extract, 0.5  ml 100  g−1) and EOs obtained by
hydrodistillation of A. boronioides aerial parts were in vitro tested at concentrations
of 15 and 50 μg ml−1 for their effects on cyclooxygenases (COX) and 5-­lypoxygenases
(LOX) pathways of eicosanoid generation such as thromboxane-B2 (TXB2),
­prostaglandin E2 (PGE2), and leukotriene B4 (LTB4) in stimulated rat peritoneal
leukocytes. The methanol extract caused a potent inhibition of LTB4 and TXB2
generation. The EO displayed strong inhibition of LTB4 but was less active on
TXB2 generation. The aqueous extract was inactive on LTB4 and with very low
activity on TXB2 generation. The effect on PGE2 production was poor, suggesting
that the site of action is on thromboxane synthetase rather than on COX (González
et al. 2003).
The lyophilized infusions (5 mg ml−1) showed an antiproliferative activity (MTT
assay) on the T84 (tumoral) and HTR-8/SVneo (non-tumoral) cell lines. The per-
centage of viable cells decreased with the increase in the concentration; however,
the highest antiproliferative effect was on T84 cells (Kogiannou et al. 2013; Gastaldi
et al. 2018).
Toxicity  A. salina bioassay showed toxic effects (LC50 5–17 mg ml−1); additional
studies have to be performed in order to define the safety and long-term effects of
the extracts (Gastaldi et al. 2018).
232 9 Patagonia

In vitro culture  Plants were micropropagated from sterile seeds, grown in MS

(Murashige and Skoog 1962)  medium supplemented with 2219  μM BAP and
0.053  μM NAA.  The micropropagated plants had 1  mg  g−1 polyphenolic and
2 mg g−1 FW anthocyanin content (Estomba et al. 2010).

9.3  Berberis microphylla G. Forest (Berberidaceae)

Synonymy  Berberis buxifolia Lam. var. antucoana, Berberis buxifolia Lam.,

Berberis heterophylla Juss. ex Poir., Berberis michay Job, Berberis parodii Job,
Berberis inermis Pers., Berberis cuneata DC., Berberis antucoana C.K. Schneid.,
Berberis barilochensis Job, Berberis buxifolia Lam. var. nuda, Berberis marginata
Gay, Berberis spinosissima (Reiche) Ahrendt, Berberis buxifolia Lam. var. spino-
sissima, Berberis buxifolia Lam. var. gracilior, Berberis buxifolia Lam. var. papil-
losa, Berberis dulcis Sweet, Berberis buxifolia Lam. var. antarctica, and Berberis
heterophylla Juss. ex Poir. var. pluriflora.

Vernacular name  Calafate, michay, box-leaved barberry, barberry, Magellan bar-

berry, and cheu-cheu.

General information  B. buxifolia is a native Patagonian species with a wide geo-

graphical distribution. In Argentina, it grows from the Patagonian woods to Tierra
del Fuego, in the provinces of CHU, NEU, RNE, SCR, and TDF (Bottini et al. 2000;
Bottini and Arena 2000). It also could be found in the Patagonian steppe and sub
Antarctic forests growing in the cordillera de los Andes, from MEN and NEU to
TDF (Rapoport et al. 1999; Roig 2001). The fruits were consumed by the onas from
TDF that also had used calafate wood to make arrows. The tehuelches prepared a
not fermented drink with the crushed and macerated fruits (Martínez Crovetto
1982). Recently, the fruits are having a high demand both to be consumed fresh and
for the elaboration of foodstuff (Arena and Cabana 2008). It is also used as orna-
mental, in parks and gardens (Bottini 2000).

Botanical characteristics  B. buxifolia is a perennial rhizomatous bush, highly

branched, with an axillary main central thorn and two low developed thorns at each
side. The leaves are simple, fasciculate, elliptic, coriaceous, and mucronate, with a
smooth margin. The adaxial side of the leaves is bright and the abaxial  one is
opaque. The orange-yellowish flowers are solitary or grouped in inflorescences,
umbels with 2–3 flowers. The perianth has six sepals and six petals in alternating
cycles of three (Arena et al. 2011). The calyx has leathery and yellow velvety petals
with two basal nectaries. Androecium has six retractable stamens, opposite to the
petals, with two dithecae-anthers with valvar dehiscence. Fruits are subglobose ber-
ries purplish blue when mature with 6–10 dark seeds (Fig. 9.3) (Orsi 1984; Landrum
1999; Arena and Curvetto 2008; Arena et al. 2011, 2013; Arena and Radice 2014).
Blooming is in November; the fruiting stage is between January and March (Tell
et al. 1997; Rapoport et al. 1999).
9.3  Berberis microphylla G. Forest (Berberidaceae) 233

Fig. 9.3  Berberis microphylla G.  Forest (Berberidaceae) branches with leaves and fruits. MA
Alvarez personal collection

Ethnobotanical information  This plant was used by native communities to treat

fever, inflammation, stomachache, diarrhea, urinary tract infection, throat infection,
gingivitis, and liver problems. Also, as an antiseptic and as a source of natural dyes
(Zin and Weiss 1998). The fruit was taken by the Kawésqar people as food; the
Aonikenk used the yellow scraping of the bark as tobacco for its hallucinogenic
effect; and the Mapuche community uses the fruit to treat colds and intermittent
fever (Domínguez et  al. 2012). Hieronymus (1882) attributed to B. buxifolia the
astringent and substitutive properties of quinine. The extracts are employed in folk
medicine mainly for their antibacterial and antidiarrheal properties. The decoction
made with leaves and stems is used for stomachache and other intestinal irritations.
The fruits are considered as astringent, digestive, and antidiarrheal, while the roots
as antidiarrheal, astringent, febrifuge, digestive, hepatic, anti-inflammatory, and
tonic. A non-specified part was referred as antipyretic, and for cold. The decoction
of dried bark is used for urinary and respiratory tract infections (Ratera and Ratera
1980; Conticello et al. 1997; Villinski et al. 2003; Caraballo and Caraballo 2004;
Alvarez et al. 2009; Barboza et al. 2009; Alonso and Desmarchelier 2015).

Chemical data  The dried entire plant has the alkaloids chillanamine, chiloenamine,
chiloenamine A, B, chiloenine, calafatamine, calafatine, calafatine 2α-N-oxide,
calafatine 2-β-N-oxide, curacutine, and talcamine. It also has resins, oil, starch, and
malic acid (Pomilio 1973; Podestá et al. 1987). The fruits have the anthocyanins
peonidin, malvidin, petunidin and delphinidin, peonidin-3-glucoside, malvidin-­3-­
234 9 Patagonia

rutinoside, malvidin-3-glucoside, petunidin-3-rutinoside, petunidin-3-glucoside,

peonidin-3-rutinoside-5-glucoside, delphinidin-3-rutinoside, delphinidin-3-­
glucoside, petunidin-3-rutinoside-5-glucoside, and petunidin-3-gentiobioside
(Pomilio 1973). The dried root has argemonine and berberine. The roots have ber-
berine, allocryptopine, calafatine, jatrorrhizine, palmatine, protopine, reticuline,
and thalifendine. The stems have berberine, allocryptopine, isocorydine, jatrorrhi-
zine, protopine, scoulerine, and thalifendine. Leaves have berberine and tetrahy-
droberberine (Fig. 9.4) (Fajardo et al. 1979; Podestá et al. 1987; Manosalva et al.
2014).

Fig. 9.4  Chemical compounds from Berberis microphylla G. Forest (Berberidaceae)

9.3  Berberis microphylla G. Forest (Berberidaceae) 235

Biological activities  Berberine has been defined as a potential drug with relevant
pharmacological properties (Singh et al. 2010; Furrianca et al. 2015). The antimi-
crobial activity was attributed to the alkaloids berberine, jatrorrhizine, and tetrahy-
droberberine (Iwasa et al. 1996
Manosalva et al. 2014). B. buxifolia fruit appeared to possess excellent antioxi-
dant activity as it ripens, which may contribute to its functional value, acquiring
relevance and applicability for nutraceutical purposes (Arena et al. 2013). The co-­
administration of B. buxifolia aqueous extract in combination with chloramphenicol
was proposed to contribute to the detoxification of oxidized metabolites originated
by antibiotic treatment (Albrecht et al. 2009).
When the methanolic extracts of leaves, stems, and roots were tested against
Escherichia coli (ATCC 25922), Salmonella typhimurium (ATCC 13311), Listeria
monocytogenes (ATCC 13932), Enterobacter aerogenes (ATCC 13084),
Staphylococcus aureus (ATCC 25923), Bacillus cereus (ATCC 11778),
Staphylococcus epidermidis (ATCC 12228), and Bacillus subtilis (ATCC 6633), the
antibacterial activity was only observed against Gram-positive bacteria (Manosalva
et al. 2016). The antibacterial activity of the extract differs according to the plant
tissue and the bacteria, which would be related to their different alkaloid composi-
tions, which seems to act synergically in stems and roots. On the contrary, the lower
activity found in leaf extracts could be related to the presence of the inactive antimi-
crobial tetrahydroberberine.
A synergistic effect was found against B. cereus, B. subtilis, and S. epidermidis
with the association of the alkaloid extract with ampicillin (AMP). A similar effect
was found with the associations of the stem alkaloid extract and AMP against S.
aureus and S. epidermidis and root alkaloid extract and AMP against B. subtilis.
Also, a synergistic effect was shown between the leaf alkaloid extract and cephalo-
thin (CFL) against S. aureus, B. cereus, and S. epidermidis, the stem alkaloid extract
and CFL against S. aureus and B. subtilis, and the root alkaloid extract with CFL
against B. cereus and S. epidermidis (Manosalva et al. 2016). Berberine, which is
the principal component of the alkaloid extracts, has been shown to in vitro enhance
the antibacterial activity of selected antibiotics against coagulase-negative
Staphylococcus strains (Wojtyczka et al. 2014). On the other hand, berberine dem-
onstrated in vitro activity against Staphylococcus epidermidis, Streptococcus spp.,
Corynebacterium diphtheriae, Trichomonas vaginalis, Neisseria gonorrhoeae, N.
meningitidis, Treponema pallidum, Entamoeba histolytica, and Leishmania don-
ovani (Kowalewski et al. 1972).
Berberine is highly effective to treat human diarrhea, with a potent activity
against Escherichia coli enterotoxins, as well as Salmonella typhi, Shigella dysente-
riae, Giardia lamblia, Salmonella paratyphi B, Klebsiella pneumoniae, and Vibrio
cholerae (Rabbani et al. 1987). However, the aqueous extract from leaves and cortex
did not present antimicrobial action against Staphylococcus aureus, E. coli,
Aspergillus niger (Anesini and Perez 1993), and Salmonella typhi (Pérez and
Anesini 1994).
236 9 Patagonia

It was reported the use of berberine against Candida albicans, to fight trachoma,
an ocular infection produced by Chlamydia trachomatis (Pizzorno and Murray
1985), the in vitro inhibition of influenza A and B virus replication (Lesnau 1990),
and the inhibition of the α-amylase enzyme, essential for the production of aflatox-
ins by the human pathogenic fungus Aspergillus flavus (Tintu et al. 2012).
Berberine has hypoglycemic effect; early studies arise from 1986; since then,
berberine was introduced as an alternative medicine for the treatment of diabetes in
Asian countries (Yin et al. 2012). The alkaloid jatrorrhizine had also hypoglycemic
activity, causing a pronounced glycemia decrease both in normal and hyperglyce-
mic mice mediated by an improvement of aerobic glycolysis (Yan et  al. 2005).
However, the response was lower than that obtained with berberine at the same dose
(Fu et al. 2005). Palmatine also lowered glycemia in normal rats (Patel and Mishra
2011; Sangeetha et al. 2013).
In vitro culture  Rooted shoots (89%) were obtained on 1/2 MS medium and 1.25
pM IBA for 7 days, followed by IBA-free medium until day 28 (Arena et al. 2000).
Callus was induced in1/2 MS medium with 2-inositol phosphate N6-(2-isopentenyl)
adenine (2iP) as plant growth regulator. Cell suspension cultures and in vitro shoot
cultures capable of producing berberine were obtained by a two-stage culture, using
in the first step picloram (PIC) and thidiazuron (TDZ) as plant growth regulators for
producing biomass and in the second step by transferring the cultures to MS medium
containing BAP (4.4 mM) as plant growth regulator. Berberine yields were signifi-
cantly lower in cell suspension cultures (102 mg g−1 DW) and shoots (200 mg g−1
DW) than in plants growing in the field (416 mg g−1 DW). Furthermore, berberine
was secreted into the medium which is advantageous for conducting biotechnologi-
cal processes in bioreactors. Extracts from in vitro cultures have demonstrated anti-
microbial activity against Escherichia coli and Staphylococcus aureus RN 4222 and
KLE820 (Alvarez et al. 2009).

Legal Status
The Argentine Food Code has incorporated berberis as a native fruit of the Andean
zone. It is authorized the use of berberis in foodstuffs such as sweets, jams, liqueurs,
ice cream, and confectionery  (ANMAT Resolutions 22/2006 and 409/2006, Res
SPReI 93/2008, and SAGPyA 356/2008). The United States Department of
Agriculture (USDA) has approved its entry into the USA.

9.4  Buddleja globosa Hope (Scrophulariaceae)

Synonymy  Buddleja capitata Jacq., Buddleja connata Ruiz & Pav., and Buddleja
globifera Duhamel.

Vernacular name  Matico, pañil, panguil, matica, palquín, matico, pallín, and
ballín.
9.4  Buddleja globosa Hope (Scrophulariaceae) 237

General information  B. globosa grows in dry and moist forests, from sea level up
to 2000 m.a.s.l. Blooming is from November to February and fructification from
December to February. It grows in the provinces of CHU, NEU, and RNE.  The
flowers are used to prepare fermented cold drinks and the fruits to make foodstuff.
Traditionally, it is grown in home gardens for domestic use (Norman and Ariza
Espinar 1995; Vogel et al. 2010).

Botanical characteristics  B. globosa is a dioecious bush, 1–5 m high, with tomen-

tose branches. Leaves are subsessile, with conspicuous stipulation, lanceolate or
elliptic, subcoriaceous to membranoceous, glabrescent, and decurrent to the base.
Flowers are clustered in pairs in pedunculated heads. The tubular tomentose calix is
tubular, with acute lobes. The intense yellow to orange corolla is glabrous on the
outside (Fig. 9.5). Stamens are sessile; ovary is tomentose in the upper half with a
claviform style. The fruit is an ellipsoidal capsule, tomentose and glandular, with
septicidal dehiscence and rounded valves (Norman and Ariza Espinar 1995).

Ethnobotanical information  The infusion made with leaves has a pleasant taste
and is consumed by the Mapuche to treat ulcers and wounds. The infusion made
with dried aerial parts is taken as digestive and hepatic, the decoction to treat urinary
and respiratory tract infections. The infusion made with dried leaves is used as uro-
genital antiseptic; to treat chronic dysentery; for stomach ulcers; as hemostatic,
antirheumatic, spasmolytic, antidiarrheal, antibacterial, anti-inflammatory, and anti-
hemorroidal; and for treating external wounds, ulcers, and acne (Houghton 1984;
Backhouse et al. 2008; Barboza et al. 2009; Gastaldi et al. 2016, 2018; Gallegos
Zurita and Gallegos Zurita 2015).

Chemical data  It has coumaric acid, p-methyl ester, echinacoside, buddledins A

and B, dehydrobuddledin A, zerumbone, buddledones A and B, acubin, catalpol,
and their related compounds, α- amyrin, linarin, verbascoside, lupeol, and mayte-
none (Fig. 9.6) (Houghton 2003; Barboza et al. 2009; Vogel et al. 2010).

Fig. 9.5  Buddleja globosa Hope (Buddlejaceae). MA Alvarez personal collection

238 9 Patagonia

Fig. 9.6  Some chemical compound found in Buddleja globosa Hope (Scrophulariaceae)

Biological activities  The antioxidant activity (DPHH and BCB assays) of a 5%

infusion was reported to be comparable to that of green tea (Mensah et al. 2001;
Sünter et al. 2012; Bustamante et al. 2015; Gastaldi et al. 2016, 2018).
Anti-inflammatory, cicatrizing, and analgesic activities as well as the regenera-
tion of the mucous membrane in case of ulcers were reported (Liao et  al. 1999;
Mensah et al. 1998, 2001; Vargas 2001; Rodríguez et al. 2001; Yañez 2001).
A recent in vitro study tested the aqueous extract of leaves on growth and protec-
tion from oxygen-free radical attack induced by hydrogen peroxide of cultured
fibroblasts. A weak and not statistically significant increase in growth was observed
at concentrations below 1 g ml−1; concentrations above 10 g ml−1 inhibited growth
and appeared to be cytotoxic. Antioxidant activity was significant at 50 g ml−1. The
activity was attributed to flavonoids and phenylethanoids, being verbascoside, echi-
nacoside, and linarin, twice as active as flavonoid aglycones (Mensah et al. 2001).
As for wound healing, the in vivo effect of this extract was tested in adult male
Sprague-Dawley rats showing no alteration of hemogram and clinical chemistry
parameters after 12 days of oral treatment. In addition, the topical treatment with the
same extract accelerated wound healing presumably by shortening the inflamma-
tory stage, as assessed by COX-2 levels, and by accelerating cellular proliferation
rate, as shown by the date of the proliferation marker Ki-67. This wound healing
effect was also attributed to the extract antioxidant activity (Letelier et al. 2012). B.
globose extract was a potent inhibitor of collagen-mediated platelet aggregation,
9.4  Buddleja globosa Hope (Scrophulariaceae) 239

probably for its high content of antiplatelet compounds such as luteolin and verbas-
coside. The cellular mechanism of that effect might be mediated by the inhibition of
PLC-g2 and PKC-b2 phosphorylation (Fuentes et al. 2017). The authors postulate a
potential use of B. globosa preparations to prevent thrombosis-related diseases
(Bustamante et al. 2015; Fuentes et al. 2017).
The leaf aqueous extract (1 mg ml−1) protected hepatocytes challenged with car-
bon tetrachloride, galactosamine, and complement-mediated cytotoxic medium
(CMC). From the isolated compounds, linarin was the most protective against car-
bon tetrachloride (40% reduction) and galactosamine (75% reduction). Echinacoside
and aucubin were also hepatoprotective (Houghton and Hikino 1989; Chang 1998;
Houghton 2003).
Polyphenol content was related to the inhibition of lipid peroxidation. It was
found that leaves from cultivated plants had the same or even higher polyphenol
content than that from wild plants. Extracts from young leaves showed the best
protection on thiol groups. Less watered plants showed higher polyphenol contents
and protection on thiol groups, whereas the inhibition of lipid peroxidation was not
affected by irrigation (Vogel et al. 2010).
The antiproliferative activity of the infusion (5 mg ml−1) was tested by the MTT
assay on T84 (tumoral) and HTR-8/SVneo (non-tumoral) cell lines. B. globosa had
a higher antiproliferative activity on HTR-8/SV neo cells than on T84 cells (Gentile
et al. 2016; Kogiannou et al. 2013; Gastaldi et al. 2018).
The sesquiterpenes buddledins A–C 15–17 (50 mg ml−1) inhibited both cyclo-
oxygenase (COX) and 5-lipoxygenase (5-LOX) enzymes being the activity stronger
against 5-LOX. Buddledin A15 showed the greatest inhibitory activity (COX 89%,
5-LOX 98% inhibition), while zerumbone 19 and dihydrobuddledin A 18 showed
no activity against COX and 5-LOX at the concentrations tested (Liao et al. 1999).
Lipophilic root extracts were tested against pathogenic, dermatophytes, and soil
fungi; the sesquiterpene components were selectively antifungal against dermato-
phytes (Epidermophyton floccosum, Trichophyton interdigitale, and E. rubrum) and
soil fungi with little effect on Penicillium notatum, Aspergillus niger, and Candida
albicans (Mensah et al. 2000). Aqueous and methanol extracts of B. globosa aerial
parts showed antiplasmodial activity, but the compounds responsible for this activ-
ity have not yet been isolated (Debenedetti et al. 2002). Also, a nematicidal effect
was reported (Bohm et al. 2009).
The antinociceptive activity of sequential ethanol extracts, verbascoside, and the
global methanol extract was tested using complementary chemical and thermal
mice models of acute pain. The analgesic effect was probably produced through a
mechanism partially linked to either lipoxygenase or cyclooxygenase enzymes via
the arachidonic acid cascade and/or opioid receptors. When verbascoside and luteo-
lin glycoside were tested, they showed similar topical analgesic effects, but verbas-
coside exhibited a much stronger oral analgesic effect than the flavonoid (Merfort
et al. 1994; Backhouse et al. 2008; Küpeli et al. 2007).
Leaf extracts were also used to green synthesis of silver nanoparticles with
potential biotechnological applications in medicine, agriculture, and bioengineering
among others (Carmona et al. 2017).
240 9 Patagonia

Finally, the hydroalcoholic extract was effective to prevent the oxidation of lipids
and thiol groups in Oncorhynchus mykiss (rainbow trout) stakes mediated by poly-
phenols and terpenes (Gormaz Araya 2005).

9.4.1  Toxicity

Mutagenic and antimutagenic activity of the aqueous extract (3.83–15.32 mg ml−1)

was tested by the wing-spot test in Drosophila melanogaster. At 15.32  mg  ml−1,
toxicity was high, but at lower concentrations (1.91, 3.83, and 7.66 mg ml−1), it did
not induce a significant increase in the frequency of mutant spot formation. When a
combined treatment, leaf extract/ethyl methanesulfonate (EMS), was tested, it was
reported a relevant reduction of single, large, and total mutant spots in somatic wing
cells in comparison to those induced by the EMS alone (Carmona et al. 2016).

9.5  Verbena litoralis Kunth var. litoralis (Verbenaceae)

Synonymy  Verbena approximata Briq., Verbena caracasana Kunth, Verbena cor-

dobensis Briq., Verbena litoralis Kunth var. caracasana, Verbena litoralis Kunth var.
leptostyachya, Verbena litoralis Kunth var. pycnostachya, Verbena affinis
M.  Martens & Galeotti, Verbena litoralis Kunth var. glabrior, Verbena nudiflora
Nutt. ex Turcz., Verbena litoralis Kunth f. angustifolia, Verbena litoralis Kunth var.
albiflora, Verbena integrifolia Sessé & Moç. f. albiflora, Verbena bonariensis L. var.
litoralis, Verbena integrifolia Sessé & Moç., and Verbena litoralis Kunth f.
albiflora.

Vernacular name  Verbena del litoral, yerba del acero, verbena encañutada, ver-
bena, gervãozinho-do-campo, and erva-de-pai-caetano (Portuguese).

General information  V. litoralis is widespread in the country, growing in the prov-

inces of BAI, CAT, CHA, COR, COS, ERI, FOR, JUJ, LPA, LRI, MEN, MIS, NEU,
RNE, SAL, SDE, SFE, SJU, SLU, and TUC. Flowering is from mid-spring until
mid-autumn.

Botanical characteristics  V. litoralis is a perennial herb with a quadrangular gla-

brous stem, opposite, narrow, ovate, ovate-lanceolate, or lanceolate  leaves with
irregular serrate margins. The blue, purple, or lavender small flowers are arranged
in groups of three in terminal cylindrical thin spikes, being the central larger than
the two laterals. Bracts are lanceolate; the tubular calix has short, triangular, uneven
teeth. Corolla has tubular lobules. Mericarps are cylindrical, brownish, with rounded
apex and base (Correa 1999; Tasso de Souza et al. 2005, Alonso and Desmarchelier
2015).
9.5  Verbena litoralis Kunth var. litoralis (Verbenaceae) 241

Ethnobotanical information  The decoction is used in cases of skin allergy and the
aerial parts as cicatrizing and anti-inflammatory and for neonatal and pediatrics
care, menorrhagia, and menopause. The decoction of dried branches and leaves is
used to treat diarrhea, and the decoction from leaves as hepatic, digestive, emmena-
gogue, anti-cephalea, febrifuge, and contraceptive to treat angina and pharyngitis
and to expel intestinal tapeworms. The root is considered as antitussive (Di Lullo
1929; Marzocca 1997; Toursarkissian 1980; Tasso de Souza et al. 2005; Barboza
et al. 2009).

Chemical data  The chemical compounds found in V. litoralis are verbenalin, ver-
benaloside, brasosid, aurantiamide acetate, jionoside, and verbenachalcone: dimeric
dihydrochalcon-60 (Fig. 9.7) (Barboza et al. 2009; Rodríguez Morcelle 2015).

Biological activity  Antioxidant activity was tested in V. litoralis hexanol extracts

by DPPH test, and (IC50 = 169.3 μg ml−1 ) (Braga et al. 2012). The hydroalcoholic

Fig. 9.7  Chemical compounds from Verbena litoralis Kunth var. litoralis
242 9 Patagonia

(1: 9 v/v) leaf extract showed activity against Staphylococcus aureus (ATCC 25923)
with an inhibition hallo of 11 mm diameter (Toribio et al. 2009).
Antinociceptive activity was tested on male Swiss mice using V. litoralis hexane
(100, 300 and 500 mg kg−1) extracts by writhing and hot plate tests. A good antino-
ciceptive potential was observed, but it was not possible to establish a dose-­
dependent relation. However, the potential supraspinal analgesic activity was only
proved to the highest dose (Braga et al. 2012). The anticonvulsant activity of the
ethanolic extract was tested for anticonvulsant activity in pentylenetetrazol (PTZ)
and maximal electroshock (MES) induced seizures in mice. The extract signifi-
cantly increased the period taken before the onset and decreased the duration of the
seizures induced by PTZ without signs of toxicity on animals (Rashidin et al. 2017).
The treatment of corn crops with different concentrations of V. litoralis (0.4, 0.8,
and 1.6 g per every 10 g of corn) showed an inhibitory activity on Sitophilus grana-
rius (maize beetle) with a 50% mortality rate at 1.6 g (Silva et al. 2001; Arango
Gutiérrez and Vásquez Villegas 2008).
Toxicity    Mice  treated with an  infusion (>25  g  kg−1) for 6  months did not have
micro or macroscopic histopathological lesions. A dose of 500 mg kg−1 of β-myrcene
was reported to cause skeletal fetus alterations in pregnant rats (Paumgartten et al.
1988). Subacute administration of the crude extract of V. litoralis at 400 mg kg−1
resulted in an increase in AST in males, whereas ALT enzyme showed a small
increase in males that received 200 mg kg−1 and 400 mg kg−1 of the extract (de Lima
et al. in press). Small farm animals that consumed V. litoralis suffered diarrhea or
died (Alonso and Desmarchelier 2015).

In vitro culture  Micropropagation was attained transferring nodal segments of V.

litoralis to MS medium supplemented with 7.5 μM BAP and 0.0005 μM NAA to
induce multiple shoots and transferring to MS medium with 0.2 μM IAA for rooting
(Braga et al. 2012).

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Index

A synonymy, 230
Acacia caven (Molina) Molina var. caven toxicity, 231
biological activities, 141 vernacular name, 230
botanical characteristics, 140, 141 Adiantum raddianum C. Presl (Pteridaceae)
chemical data, 141 biological activities, 196
ethnobotanical information, 140 botanical characteristics, 196
general information, 140 chemical compounds, 197
in vitro culture, 142 chemical data, 196
synonymy, 140 ethnobotanical information, 196
vernacular name, 140 general information, 196
Acantholippia seriphioides (A. Gray) vernacular name, 196
Moldenke Allophylus edulis (A. St.-Hil., A. Juss. &
biological activities, 46 Cambess.) Hieron. ex Niederl.
botanical characteristics, 45 biological activities, 114
chemical data, 46 botanical characteristics, 112
ethnobotanical information, 46 chemical data, 113
general information, 45 ethnomedical information, 112
synonymy, 45 general information, 112
vernacular name, 45 synonymy, 111
Achyrocline satureioides (Lam.) D.C. toxicity, 114
biological activities, 49–52 vernacular name, 112
botanical characteristics, 47 Aloysia citriodora Palau (Verbenaceae)
chemical data, 48 biological activities, 201
ethnobotanical information, 47 botanical characteristics, 199
general information, 47 chemical compounds, 200
in vitro cultures, 52 chemical data, 199
legal status, 53 cytotoxicity, 203
synonymy, 47 ethnobotanical information, 199
vernacular name, 47 general information, 199
Adesmia boronioides Hook. f. in vitro culture, 203
biological activities, 230, 231 legal status, 203
botanical characteristics, 230 vernacular name, 199
chemical data, 230 Aloysia gratissima (Gillies & Hook. ex Hook.)
ethnobotanical information, 230 Tronc.
in vitro culture, 232 biological activities, 37
mapuche-tehuelche communities, 230 botanical characteristics, 36, 37

© Springer Nature Switzerland AG 2019 249

M. A. Alvarez, Pharmacological Properties of Native Plants from Argentina,
https://doi.org/10.1007/978-3-030-20198-2
250 Index

Aloysia gratissima (Gillies & Hook. ex Hook.) Paraná flooded savanna, 4

Tronc. (cont.) Patagonia forest, 4
chemical data, 37, 38 Patagonian steppe, 4
ethnobotanical information, 37 Puna grassland, 3
general information, 36 shrublands, 4
synonymy, 36 South Atlantic islands, 5
vernacular name, 36 Thistle of the Prepuna, 3
Alternanthera pungens Kunth. Yungas, 3
biological activities, 155, 157 geographical regions
botanical characteristics, 155 Antártida, 2
chemical data, 155, 156 Cuyo, 2
ethnomedical information, 155 Gran Chaco, 2
general information, 154 Mesopotamia, 2
synonymy, 154 Northwest region, 2
toxicity, 157 Patagonia, 2
vernacular name, 154 Región Pampeana, 2
Ampicillin (AMP), 235 South Atlantic Islands, 2
Anadenanthera colubrina (Vell.) Brenan var. medicinal plants, 10, 11
cebil (Griseb.) Altschul native medicinal flora, 10, 11
biological information, 101, 102 phytogeographical regions, 7
botanical characteristics, 100 Antarctic region, 8
chemical data, 101 biomes, 9
ethnobotanical information, 100, 101 Cabrera classification, 9
general information, 99 definition, 6
synonymy, 99 endemic and endemism, 9
vernacular name, 99 Neotropical region, 6–8
Anemia tomentosa (Savigny) Sw. var. Oceanic region, 9
anthriscifolia (Schrad.) Mickel physiognomic-floristic map, 9
biological activities, 149 Argentine Food Code, 236
botanical characteristics, 148 Argentine National Pharmacopoeia (FNA), 11
chemical data, 149, 150 Asparagine, 83
ethnobotanical information, 149
in vitro culture, 149
synonymy, 148 B
vernacular name, 148 Basal acrodrome venation, 179
Antipyretic, 233 Bauhinia forficata subsp. pruinosa (Vogel)
Apigenin-7-O-rhamnosyl-6-C-glucoside, 68 Fortunato & Wunderlin
Archeobotany, 20, 22 biological activities, 205
Argentina botanical characteristics, 204
description, 1 chemical compounds, 205
eco-regions chemical data, 204
Alto Paraná Atlantic forests, 5 ethnobotanical information, 204
Antarctic Peninsula, 5 general information, 204
Argentina Low Monte, 5 in vitro cultures, 208
Argentine Northwest Monte, 3 toxicity, 208
Arid Chaco, 4 vernacular name, 204
definition, 3 Bella sombra, 70
Espinal, 4 Benzylamine purine (BAP), 77
grasslands, 4 Berberine, 235, 236
high Andean Complex, 3 Berberis microphylla G. Forest
Humid Chaco, 4 biological activities, 235, 236
Iberá wetlands, 4 botanical characteristics, 232
Mar Argentino, 5 chemical compounds, 234
Pampa, 5 chemical data, 233
Index 251

ethnobotanical information, 233 interactions, 99

geographical distribution, 232 legal status, 99
in vitro culture, 236 synonymy, 95
legal status, 236 toxicity, 99
synonymy, 232 vernacular name, 95
vernacular name, 232 Ceibales, 66
Blechnum occidentale L. Central nervous system (CNS), 69
biological activities, 119 Chaco region
botanical characteristics, 118 characteristic vegetation, 193
chemical data, 119 forests, 194
ethnobotanical information, 119 Chagas-Mazza disease, 178
general information, 118 Chemoprotective effect, 202
synonymy, 118 Chenopodium ambrosioides L.
toxicity, 119 biological activities, 145–147
vernacular name, 118 botanical characteristics, 143
Buddleja globosa Hope chemical data, 144
antinociceptive activity, 239 ethnomedical information, 143
antioxidant activity, 238 general information, 142
antiproliferative activity, 239 legal status, 148
biological activities, 238 synonymy, 142
biotechnological applications, 239 toxicity, 147, 148
botanical characteristics, 237 vernacular name, 142
chemical compound, 238 Clinopodium gilliesii (Benth.) Kunze
chemical data, 237 biological activities, 172
ethnobotanical information, 237 botanical characteristics, 172
hydroalcoholic extract, 240 chemical compounds, 173
leaf aqueous extract, 239 chemical data, 172
lipophilic root extracts, 239 ethnobotanical information, 172
polyphenol content, 239 information, 172
sesquiterpenes buddledins A–C 15–17, 239 toxicity, 174
synonymy, 236 vernacular name, 171
toxicity, 240 Coronopus didymus (L.) Smith, 169
vernacular name, 236 Cuyo region
wound healing, 238 Acantholippia seriphioides (see
Butylated hydroxytoluene (BHT), 114 Acantholippia seriphioides
(A. Gray) Moldenke)
Achyrocline satureioides (see Achyrocline
C satureioides (Lam.) D.C.)
Caesalpinia gilliesii var. gilliesii (Wall. ex Aloysia gratissima (see Aloysia
Hook.) D. Dietr. gratissima (Gillies & Hook.
biological activities, 210 ex Hook.) Tronc.)
botanical characteristics, 209 arid climate, 33
chemical data, 210 dry weather and cold winters, 33
ethnobotanical information, 210 geographical characteristics, 33
general information, 209 Lippia integrifolia (see Lippia integrifolia
toxicity, 212 (Griseb.) Hieron.)
vernacular name, 208 livestock production, 34
Cationic antimicrobial peptides (CAMPS), 72 medicinal plant species, 34, 36
Cecropia pachystachya Trecul (Cecropiaceae) Minthostachys mollis (see Minthostachys
biological activities, 98 mollis (Griseb.) Epling.)
botanical characteristics, 96 San Juan, 35
chemical data, 96 San Luis, 35
ethnobotanical information, 96 vegetation, 34
general information, 95 Cyclooxygenases (COX), 231
252 Index

E nonrenewable petroleum-based
Ecological ethnobotany, 16 substances, 75
Emmenagogue, 83 synonymy, 75
Environmental pollution, 1 vernacular name, 75
Epoxylignanes, 83
Erythrina crista-galli L. var. crista-galli
biological activities, 69 H
botanical characteristics, 66 Human embryonic stem cells (hESC), 69
chemical compounds, 68 Human-induced pluripotent stem cells
chemical data, 67, 68 (hiPSC), 69
epiphytes, 65
essential oils, 68
ethnobotanical information, 66 I
hummingbirds, 65 Ilex paraguariensis A. St.-Hil var.
isoquinoline alkaloids, 67 paraguariensis
medicinal usage, 65 biological activities, 105–107
synonymy, 65 botanical characteristics, 103
toxicity, 70 chemical data, 104
vernacular name, 65 ethnobotanical information, 103
Espinal forests, 195 general information, 103
Ethnobotany in vitro culture, 108
ancient communities, 16 legal status, 108
archeological sites, 15 synonymy, 102
in Argentina, 17–19 toxicity, 107
catholic reductions, 15 vernacular name, 103
Code of Ethics, 17 Indolbutiric acid (IBA), 77
definition, 15 Intergovernmental Platform Scientific and
different regions of Argentina, Technological Advice on
21–26 Biodiversity and Ecosystem
indigenous and criollos communities, Services (IPBES), 16
19, 21
quantitative studies, 16
traditional communities, 16 J
Ethyl methanesulfonate (EMS), 240 Jarrillas, 77
Jesuit missions, 93–95
Jesuits, 17
F Jodina rhombifolia (Hook. & Arn.) Reissek
5-Lypoxygenases (LOX), 231 biological activities, 216
Flavonoids, 238 botanical characteristics, 215
Foehn wind, 33 chemical compounds, 217
chemical data, 216
ethnobotanical information, 216
G general information, 215
Good agricultural and collection practices vernacular name, 215
(GACP), 10
Grindelia pulchella Dunal
biological activities, 76 K
botanical characteristics, 75 Kaurenoic acid methyl ester, 182
chemical compounds, 76
chemical data, 75
ethnobotanical information, 75 L
grindelic acid, 77 Larrea cuneifolia Cav.
in vitro cultures, 77 biological activities, 78, 79
labdane, 77 chemical data, 78
Index 253

ethnobotanical information, 78 M
flavonoids, 79 Maximal electroshock (MES), 242
synonymy, 78 Maytenus ilicifolia Mart. ex Reissek
vernacular name, 78 biological activities, 122, 123
Larrea divaricata Cav. botanical characteristics, 121
anti-inflammatory activity, 81 chemical data, 122
antimicrobial and antimitogenic ethnobotanical information, 122
activity, 82 general information, 121
biological activities, 80 in vitro cultures, 125
botanical characteristics, 80 legal status, 125
chemical compounds, 81 synonymy, 119
chemical data, 80 toxicity, 125
common name, 80 vernacular name, 119
ethanolic and chloroformic Medical ethnobotany
extracts, 82 Argentina, 22–23
ethnobotanical information, 80 Criollos, 21
in vitro culture, 82, 83 ethnic groups, 20
leaf aqueous extract, 82 Medicinal plant regulatory framework, 11
synonymy, 80 Mendoza (MEN), 33, 35
Larrea nitida Cav. Mesopotamia-Paranaense forest
biological activities, 84 Allophylus edulis (A. St.-Hil., A. Juss. &
chemical compounds, 84 Cambess.) Hieron. ex Niederl. (see
chemical data, 83 Allophylus edulis (A. St.-Hil., A. Juss.
common name, 83 & Cambess.) Hieron. ex Niederl.)
ethnobotanical information, 83 Anadenanthera colubrina (Vell.) Brenan
synonymy, 83 var. cebil (Griseb.) Altschul (see
Larrea spp. Anadenanthera colubrina (Vell.)
botanical characteristics, 77 Brenan var. cebil (Griseb.) Altschul)
jarrillas, 77 in Argentina, 91
L. cuneifolia, 78, 79 Blechnum occidentale L. (see Blechnum
L. divaricata, 80–83 occidentale L.)
L. nitida, 83, 84 Cecropia pachystachya Trecul
Lepidium didymum L. (Cecropiaceae) (see Cecropia
biological activities, 170 pachystachya Trecul
botanical characteristics, 169 (Cecropiaceae))
chemical compounds, 171 ecoregion of the Misiones Forest, 92
chemical data, 170 Iberá Wetlands natural reserve, 91
ethnobotanical information, 170 Iguazú Falls, 92
information, 169 Ilex paraguariensis A. St.-Hil var.
toxicity, 171 paraguariensis (see Ilex
vernacular name, 169 paraguariensis A. St.-Hil var.
Leukotriene B4 (LTB4), 231 paraguariensis)
Lipopolysaccharide (LPS), 72–73 Jesuit missions, 93–95
Lippia integrifolia (Griseb.) mammals, 92
Hieron. Maytenus ilicifolia Mart. ex Reissek
biological activities, 40, 41 (see Maytenus ilicifolia Mart. ex
botanical characteristics, 39 Reissek)
chemical data, 39 Passiflora caerulea L.
ethnobotanical information, 39 (see Passiflora caerulea L.)
general information, 39 plant species, 92
in vitro culture, 41 Tabebuia impetiginosa (Mart. ex DC.)
legal status, 41 Standl. (see Tabebuia impetiginosa
synonymy, 39 (Mart. ex DC.) Standl.)
vernacular name, 39 Micropropagation, 242
254 Index

Minimum inhibitory concentration (MIC), 43 flora, 229

Minthostachys mollis (Griseb.) Epling. hydroelectric power, 229
biological activities, 42–44 medicinal species, 229
botanical characteristics, 42 natural forest resources, 228
chemical data, 42 steppe vegetation, 229
ethnobotanical information, 42 volcanoes, 227
general information, 41 Pentylenetetrazol (PTZ), 242
legal status, 45 Peripheral blood mononuclear cells (PBMCs), 44
synonymy, 41 Phaseolus vulgaris, 72
toxicity, 44, 45 Phenylethanoids, 238
vernacular name, 41 Phytohemagglutinin (PHA), 44
Phytolacca dioica L.
biological activities, 72, 73
N botanical characteristics, 70
National Administration of Drug, Food and chemical data, 71
Medical Technology (ANMAT), 11 ethnobotanical information, 71
Nicotiana tabacum, 72 herbaceous ancestors, 70
N-methyl-D-aspartate (NMDA) receptors, 38 ombuoside, 72
Nordihydroguaiaretic acid (NDGA), 82 synonymy, 70
toxicity, 73
vernacular name, 70
O water storage tissues, 70
Obstructive sleep apnea syndrome (OSAS), 50 Phytolaccagenins (PG), 72
Picloram (PIC), 236
Pilocarpus pennatifolius Lemaire (Rutaceae)
P biological activities, 214
The Pampa plain botanical characteristics, 212
agricultural economy, 63 chemical compounds, 213
agro-industry region, 63 chemical data, 213
description, 63 ethnobotanical information, 213
E. crista-galli, 65–70 general information, 212
fauna, 64 in vitro culture, 214
flora, 64 vernacular name, 212
G. pulchella, 75–77 Plantago australis Lam. subsp. Australis
horticultural production, 64 biological activities, 152, 153
landscape, 64 botanical characteristics, 151
Larrea (see Larrea spp.) chemical data, 151, 152
P. dioica, 70–73 ethnomedical information, 151
Pampero wind, 63 general information, 151
plant communities, 65 toxicity, 153, 154
S. humboldtiana, 73–74 vernacular name, 150
Sudestada wind, 63 Pokeweed mitogen (PWM), 44
Passiflora caerulea L. Prostaglandin E2 (PGE2), 231
biological activities, 116, 118 Puna region
botanical characteristics, 115 Acacia caven (see Acacia caven (Molina)
chemical data, 116, 117 Molina var. caven)
ethnobotanical information, 116 Alternanthera pungens Kunth. (see
general information, 115 Alternanthera pungens Kunth.)
in vitro culture, 118 Anemia tomentosa (see Anemia tomentosa
legal status, 118 (Savigny) Sw. var. anthriscifolia
synonymy, 114 (Schrad.) Mickel)
vernacular name, 115 Chenopodium ambrosioides L. (see
Patagonia Chenopodium ambrosioides L.)
description, 227 cordillera de los Andes, 139
fauna, 229 fauna, 137
Index 255

flora, 138 T
Plantago australis (see Plantago australis Tabebuia impetiginosa (Mart. ex DC.) Standl.
Lam. subsp. Australis) biological activities, 110, 111
province of JUJ, 137 botanical characteristics,
provinces of JUJ, 137, 138 108, 109
salinas grandes, 139 chemical data, 109, 110
vegetation, 139 ethnobotanical information, 109
general information, 108
in vitro cultures, 111
R synonymy, 108
Ribosome-inactivating proteins (RIPs), 71 toxicity, 111
vernacular name, 108
Thidiazuron (TDZ), 236
S Thiobarbituric acid-reactive species
Salix humboldtiana Willd. (TBA-RS) assay, 49
biological activities, 74 Thromboxane-B2 (TXB2), 231
botanical characteristics, 73 Tobacco mosaic virus (TMV), 72
chemical data, 74 Tobacco necrosis virus (TNV), 72
ethnobotanical information, 74 Traditional medicine (TM), 10
rural constructions, 73 2,3,5-Triphenylltetrazolium chloride
synonymy, 73 (TTC), 50
vernacular name, 73
San Juan (SJU), 33, 36, 39, 45
San Luis (SLU), 33 U
Schinus molle L. var. molle (Anacardiaceae) United States Department of Agriculture
biological activities, 176 (USDA), 236
botanical characteristics, 175 Urban ethnobotany, 22–23
chemical compounds, 176
chemical data, 175
common name, 174 V
ethnobotanical information, 175 Verbascoside, 202
general information, 175 Verbena litoralis Kunth var. litoralis
toxicity, 178 biological activity, 241
Smallanthus macroscyphus (Baker ex Martius) botanical characteristics, 240
A. Grau chemical compounds, 241
biological activities, 182 chemical data, 241
botanical characteristics, 182 ethnobotanical information, 241
chemical compounds, 183 in vitro culture, 242
chemical data, 182 synonymy, 240
ethnobotanical information, 182 toxicity, 242
general information, 181 vernacular name, 240
toxicity, 183
Smilax campestris Griseb.
biological activity, 180 W
botanical characteristics, 179 Water remaining after hydrodistillation
chemical compounds, 180 (WRHD), 43
chemical data, 180 Woody Plant Medium (WPM), 111
ethnobotanical information, 179 World Health Organization (WHO), 10
general information, 179
in vitro cultures, 181
toxicity, 181 Y
Somatic mutation and recombination test Yungas
(SMART), 111 amphibians, 167
Systemic acquired resistance (SAR), 72 temperature and humidity, 167