Plant Geography of Chile

Plant Geography of Chile
PLANT AND VEGETATION
Volume 5
Series Editor: M.J.A. Werger
For further volumes:

http://www.springer.com/series/7549
Plant Geography of Chile
by
Andrés Moreira-Muñoz
Pontificia Universidad Católica de Chile, Santiago, Chile
123
Dr. Andrés Moreira-Muñoz
Pontificia Universidad Católica de Chile
Instituto de Geografia
Av. Vicuña Mackenna 4860, Santiago
Chile
asmoreir@uc.cl
ISSN 1875-1318 e-ISSN 1875-1326

ISBN 978-90-481-8747-8 e-ISBN 978-90-481-8748-5
DOI 10.1007/978-90-481-8748-5
Springer Dordrecht Heidelberg London New York
© Springer Science+Business Media B.V. 2011

No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by
any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written
permission from the Publisher, with the exception of any material supplied specifically for the purpose
of being entered and executed on a computer system, for exclusive use by the purchaser of the work.
Cover illustration: High-Andean vegetation at Laguna Miscanti (23◦ 43′ S, 67◦ 47′ W, 4350 m asl)
Printed on acid-free paper
Springer is part of Springer Science+Business Media (www.springer.com)

Carlos Reiche (1860–1929)
In Memoriam
Foreword
It is not just the brilliant and dramatic scenery that makes Chile such an attractive
part of the world. No, that country has so very much more! And certainly it has a
rich and beautiful flora. Chile’s plant world is strongly diversified and shows inter-
esting geographical and evolutionary patterns. This is due to several factors: The
geographical position of the country on the edge of a continental plate and stretch-
ing along an extremely long latitudinal gradient from the tropics to the cold, barren
rocks of Cape Horn, opposite Antarctica; the strong differences in altitude from sea
level to the icy peaks of the Andes; the inclusion of distant islands in the country’s
territory; the long geological and evolutionary history of the biota; and the mixture
of tropical and temperate floras.
The flora and vegetation of Chile already drew the attention of the early adven-
turers and explorers and as from the eighteenth century attracted naturalists and
collectors from Europe. In the nineteenth century famous botanists explored and
studied the Chilean plant world, and gradually the flora and plant geographical pat-
terns became subjects of scientific analyses both by European and Chilean scholars.
Recently, the development of new scientific techniques have allowed to reveal the
remarkable evolutionary pathways in many Chilean plant groups, and have provided
clues to the origins of intriguing plant geographical patterns in the southern hemi-
sphere floras. This shall be of interest for botanists, plant geographers, ecologists
and evolutionary biologists worldwide.
I was very lucky to get into contact with Dr. Andrés Moreira-Muñoz. He is an
enthusiastic and outstanding Chilean plant scientist with historical roots in this sub-
ject area. Dr. Moreira-Muñoz here presents a modern and stimulating account of
the Plant Geography of Chile that analyses the floristic diversity and endemism
of the country. He interprets the origins of the fascinating plant geographical pat-
terns of Chile and explains the evolutionary background of the most important plant
groups. I am very pleased to present this book as a volume in the series “Plant and
Vegetation” to the international readership.
Utrecht, The Nederlands Marinus J.A. Werger
vii
Preface
One morning in 1897 at the Quinta Normal, Santiago: the Director of the Museo
Nacional de Historia Natural, Federico Philippi welcomes the new German botanist
responsible for taken the reins of the botanical section, Dr Carl Reiche. He has
been committed to maintain the National Herbarium, promoting exchanges, ana-
lyzing, increasing and organizing the collections of the Herbarium. He will be
also, and this is not a trivial thing, responsible for writing the new Flora de
Chile; and he has already published the first volume. Chilean botanical knowl-
edge showed at the end of the nineteenth century still many gaps, in spite of the
great achievements of Claudio Gay and R.A. Philippi, this latter the father and
mentor of the Museum’s Director. It took Reiche more than 15 years to system-
atize, revise and add the necessary information that finally encompassed the six
volumes of the Flora de Chile (Chap. 2). In the meantime, when Reiche was
already well familiarized with the Chilean flora, he got a request for writing a
synthetic book about the Chilean plant geography for the series Die Vegetation
der Erde, edited by the great German botanists Adolf Engler and Oscar Drude.
Reiche completed the assignment successfully, and 1907 published Grundzüge der
Pflanzenverbreitung in Chile, encompassing 222 pages with two maps and sev-
eral photographs (Vegetationsbilder). This was the first (and so far the only) Plant
Geography of Chile. This great effort, which put the Chilean plant world in a
renowned world series, only got a Spanish translation 30 years later, thanks to the
engagement of G. Looser, himself a botanist and notable scientific communicator
(Chap. 2).
Just as Reiche once did with the previous works of Gay and the Philippi, now it
seems to be time for a renewal of Reiche’s Plant Geography. No few things have
changed in a hundred years: plants have been renamed and reclassified; taxonomy
and systematics have suffered far-reaching changes; biology, geography, and bio-
geography have undergone paradigmatic vicissitudes. I underwent the challenge
of writing a “New Plant Geography of Chile” as a doctoral student in Erlangen,
Germany. In such an exponentially dynamic field, one and a half year after the
publication of the thesis many things had to be revised and updated for this book.
Regarding the subject, the reader may ask why to use the old concept of “plant
geography” rather than “phytogeography” or “geobotany”? As these terms are often
used indistinctly, I decided to use the oldest term “plant geography”, honouring
ix
x Preface
the seminal works from A. von Humboldt: Géographie des plantes, and A.P. de
Candolle’s Géographie Botanique (Chap. 4). The present book also takes inspiration
from Stanley Cain’s words in his book Foundations of Plant Geography: “This is
not a descriptive plant geography, but rather an inquiry into the foundations of the
science of plant geography” (Cain 1944, p xi) (Chap. 3).
What Is This Book Not About?

This book is not a traditional geobotanical textbook. It rather attempts to enter
into the discussion on the challenges that shape (post)modern biogeography in the
twenty-first century. A detailed vegetation description, which is sometimes mis-
understood as a main task of “plant geography”, is very far from the goal of the
book. The reader is redirected to recent advances in this specific field (Chap. 1).
Many new concepts and methods are currently emerging in biogeography. This
book doesn’t offer new conceptual or methodological advances; it rather wants to
be a “field guide” to the possibilities for the development of the discipline in Chile.
Consequently, several conflicting approaches that have been proposed for explaining
current biogeographic patterns are confronted throughout the text (e.g. vicariance
versus dispersal). The result is mostly not definitive, suggesting that a dichotomy is
just a too simple problem design of a much more complex problem.
What Is This Book Then About?

The present book intends to reflect the “state of the art” or a synthesis of the plant
geographical discipline in Chile. The challenge is seemingly overwhelming, since
in such a composite discipline like biogeography, today any intend to integrate the
different views that shape it, must confront the differences inherent to the diverse
approaches involved in the discipline. To what extent biogeography assumes and
reflects the conflicts, assumptions and challenges inherent to (post)modern science
must then be kept in mind while analysing the Chilean plant geography.
This approach leaves us the theoretical basis and practical lines of direction for
the endeavour of doing plant geography in the twenty-first century, in the constantly
“changing world” of biogeography (sensu Ebach and Tangney 2007) (Chap. 10).
Most efforts at the regional level concentrate rather on the descriptive or on the ana-
lytical. I would like to do both and also to present the few results in a more general
interpretative framework. I would like to accept the challenge posted by Morrone
(2009) (Chap. 10), touching methodological as well as more theoretical aspects that
will help the student build an own “road map” towards a future development of the
discipline in Chile, integrating methods, data, concepts, and interpretations from
different fields.
Applying one of the basic principles of geography, for a better comprehension
of the subject I have often put the eye beyond the Pacific and beyond the Andes,
touching aspects of the New Zealand biota, the Antarctic palaeobiomes, Argentinian
Patagonia. . . I apologize if I have mentioned these aspects in a superficial form.
Preface xi
Nevertheless, I suspect that several aspects of the book are applicable or of interest
for biogeographers in the other (once united) southern hemisphere territories; if so,
I will be deeply satisfied.
Structure of the Book

The book is divided in five parts that organize the different chapters.
The 1st part presents an overview of the geographical and botanical scenar-
ios that shape the Chilean vascular plant world, in the present as well as in the
geologic past. In chapter one, the main physical characteristics of the Chilean ter-
ritory are briefly exposed, especially the geological and tectonic origins of Chile
and their effects on the palaeogeography and the evolution of the Southern Cone
biomes. This contributes to a better understanding of the current climate and vege-
tation. The 2nd chapter makes a succinct revision of the historical development of
Chilean botany, and synthesizes the current knowledge regarding the composition of
the flora.
The 2nd part deals with Chilean plant geographical relationships, oriented to a
synthesis of the floristic elements of the extant flora. The classification of Chilean
genera into floristic elements in Chap. 3, will be the basis for the discussion of the
disjunct patterns that shape the Chilean flora. This analysis will be further comple-
mented with the task undertaken in the 4th chapter, regarding the biogeographical
regionalization of the Chilean territory.
The 3rd part provides an analysis of two close related subdisciplines: island bio-
geography and conservation biogeography. Chapter 5 presents a synthesis of the
plant world of the Chilean Pacific offshore islands, emphasizing their uniqueness
and threats, while the 6th chapter analyses the fragmentation in the mainland, related
to the impacts of human activities on the Chilean ecosystems. Concepts and tools
developed within the field of conservation biogeography are analyzed in relation to
current global changes.
The 4th part moves into the case studies, regarding specific groups that deserve
special attention in biogeography. Chapter 7 gets into the biogeography of one
of the most charismatic American families, the Cactaceae, of course regarding its
Chilean representatives. Chapter 8 turns to another not less interesting family, the
Asteraceae, the most genus/species-rich family in Chile. The last case study is pre-
sented in Chap. 9, devoted to a monogeneric family also called the “key genus in
plant geography”: Nothofagus.
The 5th and last part of the book announces several ways in which Chilean plant
geography can further develop; maybe more rapidly and effectively than during the
last 100 years? Chapter 10 is in this sense rather speculative, in an attempt to put
Chilean plant geography in a more general context of modern biogeography. Finally,
the 11th chapter only adds several digressions about the scientific endeavour and the
artificial distinction between nature and culture.
Santiago, Chile Andrés Moreira-Muñoz

Acknowledgments
The book was initially developed as a doctoral study at the Geographical Institute
of Erlangen-Nürnberg University, Germany. Support in form of a grant was fortu-
nately provided by the German Academic Exchange Service (DAAD). I am much
indebted to Prof. Dr. Michael Richter, who was from the first moment the main sup-
porter of the idea. He and his family, together with all the colleagues and workers
at the Geographical Institute in Erlangen made our family’s stay in Germany a great
life experience. From the Geography to the Botanical Garden in Erlangen there are
just several blocks, and the support and friendship we found there in the person of
Dr. Walter Welss and his family was also a foothold in our stay. Prof. Dr.Werner
Nezadal (Erlangen) and Prof. Dr. Tod Stuessy (Vienna) gently assumed the revision
of the thesis.
The thesis was improved by the attendance of several conferences thanks
to grants from the Zantner-Busch Stiftung (Erlangen). At the conference
“Palaeogeography and Palaeobiogeography: Biodiversity in Space and Time”,
NIEeS, Cambridge, UK, 10th–11th April, I attended the workshop for using the
program TimeTrek for plate tectonic reconstructions. I also could attend the XVII
International Botanical Congress in Vienna, 17th–23rd July 2005.
The idea of transforming the thesis into a book found absolute support in the
person of Prof. Dr. Marinus Werger. He acted not just as a language editor but as a
very patient reviewer guiding the editing process in all its stages. The early intention
was also promoted by Dr. Leslie R. Landrum and Dr. Juan J. Morrone.
Crucial for the positive development of the book has been Springer’s production
and editing team: first Inga Wilde and Ria Kanters, and lately Ineke Ravesloot and
Annet Shankary. Several colleagues and friends graciously read and commented on
draft chapters: Federico Luebert (Berlin), Hermann Manríquez (Santiago), Patrick
Griffith (Florida), Malte Ebach (Arizona), Michael Heads (Wellington), Michael
Dillon (Tal Tal), Carlos Lehnebach (Wellington), and Patricio Pliscoff (Lausanne).
Of course the errors and misconceptions that may still exist are exclusively my
responsibility.
In Chile, the project found early support in Dr. Belisario Andrade (Pontificia
Universidad Católica de Chile) and Dr. Roberto Rodríguez (Universidad de
Concepción). Once back in Chile, I can only express gratitude to the colleagues
xiii
xiv Acknowledgments
at the Pontificia Universidad Católica de Chile, which facilitated my incorporation

as an assistant professor by means of a grant for young doctors. I am especially
indebted to the Director of the Institute of Geography, Dr. Federico Arenas and the
dean of the Faculty, Dr. José Ignacio González.
Field work in Chile during 2008–2010, especially for research on Asteraceae
(Chap. 8), was supported by project Fondecyt Iniciación (2008) n◦ 11085016.
Speaking about field work, long ago I learned from Calvin and Linda Heusser the
“dirty side” of scientific field work. I will be always indebted to my old friends.
Vanezza Morales was a crucial helper in the final editing of most maps, and
with computer programs like NDM/VNDM. I gratefully mention also the important
advice provided by Tania Escalante (UNAM) and Claudia Szumik (U. de Tucumán).
Giancarlo Scalera (Roma), and Carlos Le Quesne (Valdivia) kindly provided articles
and figures. Sergio Elórtegui generously acceded to draw several original illustra-
tions for this work and also contributed many photographs. Carlos Jaña helped
finishing the most complicated figures. Sergio Moreira, Walter Welss, Hendrik
Wagenseil, Jeff Marso, María Castro, Francisco Casado, and Carlo Sabaini kindly
provided photos for illustrating this book.
Last but not least, I must acknowledge the life-long support of Mélica Muñoz-
Schick and Sergio Moreira, who could transfer to me their passion for nature
and beauty. Mélica, as ever, helped with the identification of species. Sergio also
helped providing scanned images of botanical specimens, thanks to a grant to the
National Herbarium provided by the Andrew W. Mellon Foundation trough the
Latin American Plants Initiative (LAPI).
When the doctoral thesis was still a draft project, my way crossed the one of
Paola, who soon turned to become my life companion. I would not have reached this
goal without her continuous support. I could also not imagine that the relationship
would be so fruitful: Sayén, Silene, Coyán, and Relmu remind me every evening
that there are other important things in life than just writing books. . . there is also
the possibility to read them!. . . especially when they deal not just with flowers but
also with rabbits, bears, elves and fairies.
1 May 2010 Limache

Contents
Part I Geobotanical Scenario

1 The Extravagant Physical Geography of Chile . . . . . . . . . . . . 3
1.1 Tectonics and Physiography . . . . . . . . . . . . . . . . . . . 6
1.1.1 Morphostructural Macrozones . . . . . . . . . . . . . 7
1.2 Past Climate and Vegetation . . . . . . . . . . . . . . . . . . . 11
1.2.1 The Palaeozoic (542–251 mya) . . . . . . . . . . . . . 12
1.2.2 The Mesozoic (251–65.5 mya) . . . . . . . . . . . . . 14
1.2.3 The Cenozoic (65.5 mya Onwards) . . . . . . . . . . . 20
1.3 Current Climate and Vegetation . . . . . . . . . . . . . . . . . 32
1.3.1 Bioclimatic Zones . . . . . . . . . . . . . . . . . . . . 33
1.3.2 Vegetation Formations . . . . . . . . . . . . . . . . . 35
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2 Getting Geobotanical Knowledge . . . . . . . . . . . . . . . . . . . 47
2.1 Romancing the South: The Discovery of a Virgin World . . . . 47
2.2 Classification and Phylogeny of the Chilean Vascular Flora . . . 56
2.2.1 Rich Families and Genera . . . . . . . . . . . . . . . . 59
2.2.2 Endemic Families . . . . . . . . . . . . . . . . . . . . 63
2.2.3 Phylogenetic Groups . . . . . . . . . . . . . . . . . . 65
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Part II Chorology of Chilean Plants

3 Geographical Relations of the Chilean Flora . . . . . . . . . . . . . 87
3.1 Floristic Elements . . . . . . . . . . . . . . . . . . . . . . . . 87
3.1.1 Pantropical Floristic Element . . . . . . . . . . . . . . 91
3.1.2 Australasiatic Floristic Element . . . . . . . . . . . . . 92
3.1.3 Neotropical (American) Floristic Element . . . . . . . 95
3.1.4 Antitropical Floristic Element . . . . . . . . . . . . . . 98
3.1.5 South-Temperate Floristic Element . . . . . . . . . . . 101
3.1.6 Endemic Floristic Element . . . . . . . . . . . . . . . 101
3.1.7 Cosmopolitan Floristic Element . . . . . . . . . . . . 106
xv
xvi Contents
3.2 To Be or Not To Be Disjunct? . . . . . . . . . . . . . . . . . . 109

3.2.1 Pacific-Atlantic Disjunctions . . . . . . . . . . . . . . 110
3.2.2 Antitropical (Pacific) Disjunctions . . . . . . . . . . . 113
3.3 In the Search for Centres of Origin: Dispersal v/s
Vicariance in the Chilean Flora . . . . . . . . . . . . . . . . . . 114
3.3.1 Revitalizing Long-Distance Dispersal . . . . . . . . . 117
3.3.2 Relativising Long-Distance Dispersal . . . . . . . . . 120
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
4 Biogeographic Regionalization . . . . . . . . . . . . . . . . . . . . 129
4.1 The Chilean Plants in the Global Concert . . . . . . . . . . . . 130
4.2 The Austral v/s the Neotropical Floristic Realm . . . . . . . . . 136
4.2.1 Floristic Elements in the Latitudinal Profile . . . . . . 137
4.2.2 Similarity Along the Latitudinal Gradient . . . . . . . 139
4.3 Regions and Provinces . . . . . . . . . . . . . . . . . . . . . . 142
4.3.1 Endemism as the Base for Regionalization . . . . . . . 144
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Part III Islands Biogeography

5 Pacific Offshore Chile . . . . . . . . . . . . . . . . . . . . . . . . . 153
5.1 Rapa Nui . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
5.2 Islas Desventuradas . . . . . . . . . . . . . . . . . . . . . . . . 157
5.3 Juan Fernández Archipelago . . . . . . . . . . . . . . . . . . . 159
5.3.1 The Unique Plant World of Juan Fernández . . . . . . 159
5.3.2 Floristic Similarity of Juan Fernández . . . . . . . . . 165
5.3.3 Origins of the Fernandezian Flora . . . . . . . . . . . 168
5.3.4 Conservation of Juan Fernández Plants . . . . . . . . . 171
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
6 Islands on the Continent: Conservation Biogeography
in Changing Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . 181
6.1 Fragmentation v/s Conservation on Chilean
Landscapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
6.2 Global Change Biogeography: A Science
of Uncertainties . . . and Possibilities . . . . . . . . . . . . . . . 185
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Part IV Case Studies on Selected Families

7 Cactaceae, a Weird Family and Postmodern Evolution . . . . . . . 197
7.1 Cacti Classification . . . . . . . . . . . . . . . . . . . . . . . . 198
7.2 Chilean Representatives and Their Distribution . . . . . . . . . 198
7.2.1 Cacti Distribution in Chile . . . . . . . . . . . . . . . 200
7.2.2 Areas of Endemism . . . . . . . . . . . . . . . . . . . 202
7.3 Notes on Cacti Conservation . . . . . . . . . . . . . . . . . . . 209
7.4 Biogeographic Insights: Spatial and Temporal Origins . . . . . 214
Contents xvii
7.5 Cactus Postmodern Evolution . . . . . . . . . . . . . . . . . . 215

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
8 Asteraceae, Chile’s Richest Family . . . . . . . . . . . . . . . . . . 221
8.1 Classification of Chilean Asteraceae . . . . . . . . . . . . . . . 221
8.2 Floristic Elements of Chilean Asteraceae . . . . . . . . . . . . 223
8.3 Biogeographic regionalization of the Chilean Asteraceae . . . . 229
8.4 Asteraceae Evolutionary Biogeography . . . . . . . . . . . . . 234
8.4.1 Origin and Dispersal Routes . . . . . . . . . . . . . . 238
8.4.2 Dispersal Capacities . . . . . . . . . . . . . . . . . . . 238
8.5 Conservation v/s Invasions . . . . . . . . . . . . . . . . . . . . 239
8.5.1 Invading Biogeography . . . . . . . . . . . . . . . . . 241
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
9 Nothofagus, Key Genus in Plant Geography . . . . . . . . . . . . . 249
9.1 Taxonomy and Phylogeny . . . . . . . . . . . . . . . . . . . . 249
9.2 Diversity and Distribution . . . . . . . . . . . . . . . . . . . . 254
9.3 Speciation v/s Extinction . . . . . . . . . . . . . . . . . . . . . 256
9.4 Vicariance v/s Dispersal and Centres of Origin . . . . . . . . . 260
9.5 Nothofagus and Associated Taxa . . . . . . . . . . . . . . . . . 262
9.6 Synthesis and Outlook . . . . . . . . . . . . . . . . . . . . . . 263
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263
Part V Where to from Here? Projections of Chilean Plant Geography

10 All the Possible Worlds of Biogeography . . . . . . . . . . . . . . . 269
10.1 The Fragmented Map of Modern Biogeography . . . . . . . . . 269
10.2 Postmodern Biogeography: Deconstructing the Map . . . . . . 270
10.3 Sloppy Biogeography v/s Harsh Geology? . . . . . . . . . . . . 273
10.4 Just Some Possible Worlds . . . . . . . . . . . . . . . . . . . . 275
10.4.1 Connections Over Land Bridges . . . . . . . . . . . . 276
10.4.2 And What About a Closer Pacific Basin? . . . . . . . . 276
10.4.3 Three Models of Gondwana Fragmentation +
One Dispersal . . . . . . . . . . . . . . . . . . . . . . 279
10.5 The “New Biogeography” . . . . . . . . . . . . . . . . . . . . 282
10.6 Coda: The Geographical Nature of Biogeography . . . . . . . . 284
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
11 Epilogue: The Juan Fernández Islands
and the Long-Distance Dispersal of Utopia . . . . . . . . . . . . . . 293
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294
Appendix A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
General Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
Vascular Chilean Plant Genera Index . . . . . . . . . . . . . . . . . . . 335
Abbreviations
AAO Antarctic Oscillation

ACC Antarctic circumpolar current
cfr. Refer to
Chap. Chapter
CONAF Corporación Nacional Forestal
CONC Herbario de la Universidad de Concepción
ENSO El Niño Southern Oscillation
Fig. Figure
GIS Geographic information systems
ITCZ Intertropical Convergence Zone
IUCN International Union for Conservation of Nature
K/T boundary Cretaceous/Cenozoic boundary
LDD Long-distance dispersal
LGM Last Glacial Maximum
m asl Metres above sea level
mya Million years ago
PAE Parsimony Analysis of Endemicity
PDO Pacific Decadal Oscillation
SEBA Systematic and Evolutionary Biogeographical Association
Sect. Section
SGO National Herbarium Santiago, Chile
SNASPE National public protected areas system
yr BP Years before present
xix
About the Author
Andrés Moreira-Muñoz was born in Los Angeles (Chile), studied at the German
School in Santiago and graduated as Professional Geographer at the Pontificia
Universidad Católica de Chile. Botanical interest was inherited from his grand-
father and mother, both renowned botanists at the Museo Nacional de Historia
Natural in Santiago. He obtained his doctoral degree in Geography from the
University Erlangen-Nürnberg, Germany, under the direction of the plant geogra-
pher Prof. Michael Richter.
He currently occupies a position as assistant professor at the Instituto de
Geografía, Pontificia Universidad Católica de Chile, and develops research projects
about the chorology of Chilean plants, conservation biogeography and field-based
education.
He is a member of several national and international associations like the
Systematics and Evolutionary Biogeographical Association (SEBA), the Society for
Conservation GIS (SCGIS), the IUCN Species Survival Commission, the Sociedad
de Botánica de Chile, and Corporación de Investigación y Divulgación Científica
Taller La Era (www.tallerlaera.cl).
xxi
Part I
Geobotanical Scenario
Chapter 1
The Extravagant Physical Geography of Chile
Abstract Current Chilean vascular flora and its biogeographical patterns are
strongly related to the geographical features of the territory, past and present. Main
characteristics of the physical geography of Chile are described, with emphasis on
the geologic and climatic changes that affected the biome configuration since the
Devonian onwards. Approaching the present time, the effects of the Pleistocene
glaciations in the distribution of several communities are discussed.
Chile has been characterized as “a geographic extravaganza” (Subercaseaux 1940)

due to its impressive geographical contrasts: it contains the driest desert on the
planet, formidable inland ice fields, active volcanoes, fjords, geysers, a vast coastline
and the major highs of the Andes.
The country stretches for 4,337 km along the south-western margin of South
America from the Altiplano highs at 17◦ 35′ S to Tierra del Fuego, the Islas Diego
Ramírez and Cape Horn at 56◦ S (Figs. 1.1 and 1.2). Chile’s boundary to the
west is the wide Pacific Ocean. The national territory includes several groups of
Pacific oceanic islands, principally Rapa Nui (Easter Island), the Juan Fernández
archipelago, and the Desventuradas Islands (Fig. 1.1) (Chap. 5). Besides this the
nation has a geopolitical claim on a portion of 1,250,000 km2 in Antarctica. Though
geopolitical interests are beyond the scope of this book, and despite the modest
presence of extant vascular plants in Antarctica (only Deschampsia antartica and
Colobanthus quitensis), the Continent of Ice is of high interest regarding the origin
of the Chilean plant world (Sect. 1.2, Box 9.1).
A. Moreira-Muñoz, Plant Geography of Chile, Plant and Vegetation 5, 3

DOI 10.1007/978-90-481-8748-5_1, ⃝
C Springer Science+Business Media B.V. 2011
4 1 The Extravagant Physical Geography of Chile
90” W 72” W
17” S
Islas Desventuradas
Juan Fernández
archipelago
110” W
Isla Salas y Gómez
Rapa Nui (Isla de Pascua)
Islas Diego
Ramírez 53” W
Oceanic Chile
Continental / Antarctic Chile
0 1,000 2,000 km
90° S
Fig. 1.1 Chile including the American continental portion, the Pacific islands, and Antarctic
Peninsula. Polar stereographic projection with true scale at 71◦ S using ArcGIS 9. Base global
map provided by ESRI Labs
The eastern margin of mainland Chile is the Andes cordillera, which reaches to a
maximum of 6,962 m asl in the Monte Aconcagua at 32◦ 39′ S (Fig. 1.6). As its sum-
mit is located on the Argentinean side, the highest peak of the Chilean Andes is the
Ojos del Salado volcano at 27◦ 06′ S, reaching 6,893 m asl. Contrary to the long lati-
tudinal extent, in width Chile rarely extends more than 200 km, reaching a maximum
of 360 km at Mejillones (23◦ S) and a minimum of 90 km at Illapel (31◦ 37′ S). The
difference in altitude from the coast to the high Andes creates a series of bioclimatic
variations in the altitudinal profile (Fig. 1.6). These variations, coupled with the cli-
matic latitudinal gradient, create a variety of geographic conditions that dramatically
1 The Extravagant Physical Geography of Chile 5
a b
c d
e f
Fig. 1.2 Physical geography of Chile: a Valle de la Luna, Atacama desert, 23◦ S; b Cerro Las
Vizcachas, Cordillera de la Costa, 33◦ S; c rocky coast at Concón, Valparaíso (32◦ 50′ S); d Laguna
del Inca, Portillo, Andean pass to Argentina (32◦ 50′ S); e Glaciar Los Perros, Torres del Paine,
Campos de Hielo Sur (51◦ S); f southern fjords and Cordillera de Darwin (55◦ S) (photo credits: a,
b, d–f A. Moreira-Muñoz; c S. Elórtegui Francioli)
affect the Chilean vegetation from the arid North to the humid temperate rainforests
in the South (Sect. 1.3).
1.1 Tectonics and Physiography

The main character of Chilean landscapes is driven by tectonic forcing: the geo-
logical evolution of Chile is related to the east-directed subduction of the Nazca
Plate beneath the South American Plate (Pankhurst and Hervé 2007) (Fig. 1.3).
The Chile Rise is an active spreading centre that marks the boundary between the
Nazca Plate and the Antarctic Plate at the so called Chile Triple Junction (Fig. 1.3).
The Nazca Plate is being subducted at a rate of ~65 mm/year (to the North of the
Triple Junction), while the Antarctic Plate is being subducted at a slower rate of
~18 mm/year (Barrientos 2007). According to Ranero et al. (2006), the amount of
sediments to the trench is variable in space and time: north of 28◦ S, due to aridity,
there is a relatively small amount of erosion and sediment supplied to the trench; in
the mid-latitude, the well developed river drainage system supplies much material
to the trench; south of ~ 40◦ S glacial-interglacial periods might have controlled the
amount of sediment supplied to the trench (Ranero et al. 2006).
Cocos
Plate
a
e
Galápagos
Ris
Pacific Plate
Pe
cific
r
u-
Ch
t Pa
ile
Nazca Plate
Tre
Eas
nch
b
Desventuradas
Rapa Nui
Pitcairn Is.
Juan Fernández
Ch c
ile South American
Ri
se Plate
Chile Triple
d
Junction
Scotia Plate
Antarctic Plate
Fig. 1.3 Tectonic main features and volcanic zones of South America: a northern volcanic zone;
b central volcanic zone; c southern volcanic zone; d austral volcanic zone (adapted from Orme
(2007), by permission of Oxford University Press; see also Stern et al. (2007))
1.1 Tectonics and Physiography 7
A prominent feature of the Nazca Plate is the Juan Ferrnández hot spot chain,
a series of disconnected seamounts that disappear into the trench at 33◦ S (Ranero
et al. 2006) (Fig. 1.3). Subduction is accompanied by intense magmatic and seismic
activity (Orme 2007). Great earthquakes occur somewhere along the western South
American margin every few years, and “no recorded human generation in Chile has
escaped the damaging consequences of large earthquakes” (Barrientos 2007, p 263).
Indeed, while writing these lines, on the 27th of February 2010, an earthquake with
a magnitude of 8.8 followed by a tsunami affected Central-south Chile, resulting in
hundreds of deaths and thousands homeless.
Together with earthquakes, the active volcanism along the length of the country
is also a good reminder of the active tectonic processes acting below the surface
(Box 1.1).
Box 1.1 Living Under the Volcano

Chilean active and inactive volcanoes comprise ca.10% of the circum-Pacific
“ring of fire” (Pankhurst and Hervé 2007). These are mostly andesitic stra-
tovolcanoes that occupy almost the entire length of the country, especially
at the “South Volcanic Zone”, that encompass most of the South American
active volcanoes (Stern et al. 2007) (Fig. 1.3). More than 150 potentially active
volcanoes have been detected, and 62 of them erupted in historical times
(González-Ferrán 1994). One of the most recent is the eruption of Volcán
Chaitén (43◦ S) on May 2008, which was responsible for the obligate aban-
donment of the homonymous town. The ash column reached a height of 15 km
and spread wide upon the Atlantic (Figs. 1.4 and 1.5). Apart from its conse-
quences and risks for human occupation, volcanism has been a constant source
of disturbance in the Chilean ecosystems, especially in the southern temperate
forests (Milleron et al. 2008).
1.1.1 Morphostructural Macrozones
Taking account of its tectonic and morphostructural features, Chile can be classified
in a broad sense in five macrozones (Fig. 1.6) (Charrier et al. 2007; Stern et al.
2007):
(a) The Coastal Cordillera occupies the western part of the profile from 18◦ S
to Chiloé Island (~ 42◦ S). It comprises the coastal batholith that consists
predominately of Late Palaeozoic and Mesozoic igneous rocks, with paired
belts of Palaeozoic metamorphic rocks cropping out south of Pichilemu
(34◦ 23′ S) (Pankhurst and Hervé 2007). Very impressive is the high riffs
(“acantilado”) that stretches from 0 to 800 m asl at Iquique (20◦ S).
(b) The Central Depression is a tectonic downwarp with a Mesozoic to Quaternary
sedimentary fill of volcanic, glacial and fluvial origin. This main agricultural
b c
Fig. 1.4 Examples of volcanic activity in historical times: a ash expulsion by Volcán Antuco on
the 1st March 1839, as represented in Claudio Gay’s Atlas (Chap. 2); b eruption of Volcán Carrán
in 1955 (from Illies 1959); c Volcán Chaitén eruption photographed on May 26, 2008 (photo by
J.N. Marso, courtesy of the USGS)
and urbanized region ranges from 18◦ S to Copiapó (27◦ S), and again from
Santiago (33◦ S) to Chiloé (42◦ S). It is absent between 27◦ and 33◦ S, in the
so called zone of transverse river valleys or “Norte Chico” (Weischet 1970;
Charrier et al. 2007). This zone corresponds also to the “flat slab” zone, a
zone free of recent volcanic activity, associated to the subduction of the Juan
Fernández Ridge (Fig. 1.3).
(c) The main Andean Cordillera is a chain of mountains that dates back to the
Miocene, whose emergence continues today (see Box 1.5). It can be subdivided
in three segments: Forearc Precordillera and Western Cordillera, between 18◦
and 27◦ S; High Andean Range, between 27◦ and 33◦ S (flat-slab subduction
1.1 Tectonics and Physiography 9
a c
Fig. 1.5 Chilean volcanoes: a Parinacota volcano, 18◦ 10′ S; b steam expulsion of Volcán Lascar
(23◦ 20′ S), on December 1996; c Volcán Chaitén (42◦ 50′ S), false colour Aster satellite image:
plume of ash and steam advancing ca. 70 km to the north-east on January 2009; d lava fields
around Nevados de Chillán (36◦ 50′ S) (photo credits: a H. Wagenseil; b, d A. Moreira-Muñoz; c
NASA Earth Observatory (www.earthobservatory.nasa.gov))
segment); and Principal Cordillera, between 33◦ and ca. 42◦ S (Charrier
et al. 2007).
(d) Patagonian Cordillera: the Andes’ continuation right down into Tierra del Fuego
at the southern tip of Chile, with a continuous reduction in height (Pankhurst
and Hervé 2007). The origin of this low portion of the Andes has been related
to an allochtonous Palaeozoic terrane (see Box 1.2). The west-southern margin
of the land (42◦ to the South) is modeled by recent glaciations that carved the
coastal areas into fjords and archipelagos comprising thousands of little islands
(Pankhurst and Hervé 2007). It has been calculated that the coastal extension
of Chile including these islands and southern archipelagos reaches 83,850 km!
(IGM 2005).
(e) The Andean foreland of the southern Patagonian Cordillera or Magallanes basin
consists of Upper Jurassic to Early Cenozoic sedimentary deposits (Charrier
et al. 2007; Fosdick 2007).
Profile 20° S
4,000
3,000
2,000
1,000
0
0 20 40 60 80 100 120 140 160 km
Profile 30°S
4,000
3,000
2,000
1,000
0
0 20 40 60 80 100 120 140 km
Profile 33°S
5,000
4,000
3,000
Cordillera de La Costa (La Campana)
2,000
1,000
0
0 20 40 60 80 100 120 140 km
Profile 37.5° S
2,500
2,000
Cordillera de Nahuelbuta
1,500
1,000
500
0
0 20 40 60 80 100 120 140 160 180 200 220 km
Profile 40°S
1,600
1,200
800
Cordillera Pelada
400
0
0 20 40 60 80 100 120 140 160 km
Profile 50°S
2,000 Campo de Hielo Sur

1,500 Fjords and channels zone
1,000
500
0
0 20 40 60 80 100 120 140 160 km
Fig. 1.6 Physiography of continental Chile, on the base of SRTM (Shuttle Radar Topography
Mission) data (http://www2.jpl.nasa.gov/srtm/), five morphostructural zones (see text for expla-
nation; for national political borders see Fig. 1.1). Altitudinal profiles have been produced with
ArcGIS 9 based on Aster GDM data (http://asterweb.jpl.nasa.gov/gdem.asp). Note variations in
the vertical scale, not homogeneous
Box 1.2 Patagonian Vicissitudes

The remarkable landscape and flora of Patagonia motivated early naturalists
like the Perito Francisco P. Moreno to propose an independent origin of this
microcontinent from the rest of South America (Moreno 1882, as quoted by
Ramos 2008). The characteristic landscape and rocks led Moreno to remark
strong affinities to other southern landmasses like Antarctica, Australia, and
New Zealand, suggesting that Patagonia was the rest of a sunken continent.
1.2 Past Climate and Vegetation 11
This view was retained even during the time of continental drift discussion
(e.g. Windhausen 1931). Current geologic and palaeomagnetic data suggests
that indeed, Patagonia has seen successive periods of breaking and drifting
during the whole Palaeozoic (Rapalini 2005; Ramos 2008). The TimeTrek
model (see also Pankhurst et al. 2006) shows an amalgamation of Patagonia
to Antarctic Peninsula during Late Carboniferous (300 mya), and a grad-
ual separation from Antarctica into the Cretaceous (120 mya) (Fig. 1.7).
Biotic exchange between South America and Antarctic Peninsula may have
been favoured (and then prevented) more than just one time, following rather
exchange cycles (Fig. 1.7).
a b c
Fig. 1.7 Positions of Patagonia: a in the Late Carboniferous (300 mya) aggregated to the
Antarctic Peninsula; b in the Early Cretaceous (120 mya), separated from Antarctica; c in
the Eocene (50 mya), again close to the Antarctic Peninsula. Modeled with TimeTrek v
4.2.5, Cambridge Paleomap Services
1.2 Past Climate and Vegetation

Tectonic and geomorphologic processes, coupled with the oceanic-atmospheric
system, have had enormous effects on the botanical evolution and its physiognom-
ical expression (i.e. the vegetation). The main aspects of the palaeogeographical
evolution of the territory will be resumed hereafter.
Palaeobotanical studies of Chile date back to Engelhardt (1891), Ochsenius
(1891), Dusén (1907), Berry (1922a, b), Fuenzalida (1938, 1966) among others.
More recent advances are centered in the Cenozoic (e.g. Cecioni 1968; Nishida
1984; Troncoso and Romero 1998; Hinojosa 2005). Constant improvement of the
methods applied to the study of “climatically sensitive” sediments (e.g. coals, salt
deposits, evaporites), together with studies in diversity patterns in global vegetation
through time, are benefiting our understanding of the evolution of plant biomes in
space and time (Willis and McElwain 2002).
The floristic and vegetational history of southern South America is strong related
to the tectonic and climatic history of the Gondwana continent (McLoughlin 2001)
COOL WARM COOL WARM COOL WARM COOL WARM COOL

30°C
20°C
10°C
Palaeogene
Ordovician
Cretaceous
Cambrian
Devonian
Neogene
Carbonif.
Quaternary
Permian
Jurassic
Silurian
Triassic
145
23
199
359
488
65
251
443
299
542
416
2.5
Fig. 1.8 Global climate change since the Cambrian onwards. Adapted from Frakes et al. (1992)
and Scotese et al. (1999). Dates have been updated with the 2004 Geologic Time Scale (Gradstein
et al. 2004)
(Box 1.3, Table 1.1). “During the 500 million years that Gondwana and its fragments
existed, the Earth’s global climate system has shifted from ‘Ice House’ conditions
to ‘Hot House’ conditions four times” (Scotese et al. 1999) (Fig. 1.8). These global
climatic fluctuations have constantly affected the biotic evolution and biogeography:
floristic regions can be tracked back even to the mid-late Silurian, the time when
according to most palaeobotanical evidence, the vascular plants have conquered the
land surface (Willis and McElwain 2002; Raymond et al. 2006) (Box 2.3).
1.2.1 The Palaeozoic (542–251 mya)

Several orogenic events affected the western margin of Gondwana from the Late
Proterozoic to the Palaeozoic (Ramos and Aleman 2000; Pankhurst et al. 2006).
The Famatinian orogeny in the Ordovician (~490–450 mya) is characterized by the
amalgamation of several allochtonous terranes, like Cuyania and Chilenia, imply-
ing that North America had collided with West Gondwana by that time (Astini et al.
1995). Mejillonia and Patagonia terranes amalgamated in the Early Permian, as the
last convergence episodes (Ramos 2009) (Box 1.2). The development of preAndean
foreland basins during the Palaeozoic, set the stage for the initiation of the Andes
long before the event that culminated in massive Cenozoic uplift (Orme 2007).
During the Late Palaeozoic, Gondwana became amalgamated to the supercontinent
of Laurussia to form the vast single landmass called Pangaea.
From the Early Devonian to the Late Carboniferous (400–300 mya), global veg-
etation evolved from one dominated by small, weedy plants, only several decimetres
in height, to fully forested ecosystems with trees reaching sizes of 35 m (Willis and
McElwain 2002). During the Middle to Late Devonian (390–360 mya) warm, humid
climates with high levels of atmospheric CO2 prevailed worldwide, favouring the
appearance of earliest arborescent forms of plants (see Box 2.3).
Subtropical desert
Cool temperate
Ice sheet
Fig. 1.9 Late Carboniferous biomes (adapted from Willis and McElwain (2002) on a TimeTrek
4.2.5 model, Cambridge Paleomap Services)
By the Late Carboniferous (330–299 mya) the southern flora consisted

mainly of likely pteridosperms, lycopsids, Cordaites and Ginkgophytes (Vega and
Archangelsky 1997). Diversity was rather low, and the southern flora was uni-
formly developed across Gondwana between 30◦ S and 60◦ S (Anderson et al.
1999; DiMichele et al. 2001). However, Cúneo (1989) suggests that floristic
differentiation was also apparent on the west coast of South America. The pres-
ence of Lepidodendron and Sigillaria (lycopod trees) has been reported from
the Carboniferous deposits of Chile (Charrier 1988). Late Carbonifeous ended
in a widespread glaciation, one of the most severe in Earth’s history. The
Permo-Carboniferous glaciation (310–290 mya) lasted for around 30 million years
(Beerling 2002); Gondwanan continents were locked in deep glaciation (Fig. 1.9).
The Permian (299–251 mya) was characterized by major global climate
changes, from glaciated (icehouse) to completely ice-free (hothouse) stages
(Fig. 1.8). “With the onset of glaciation in the Permian, the flora changed dra-
matically with the appearance of Glossopteris and the disappearance of most of
the Late Carboniferous elements” (DiMichele et al. 2001, p 467). By the Middle
Permian, one of the most striking vegetation changes was the relatively increased
proportion of seed plants together with a reduction of the swamp-dwelling lycopsids
and sphenopsids (Wnuk 1996, McAllister Rees et al. 2002). Glossopteris, a gym-
nosperm genus with many species, turned to be the characteristic plant of Gondwana
(DiMichele et al. 2001). Indeed, Glossopteris dominant presence across Gondwana
Subtropical desert
Mid latitude desert
Cold temperate
Cool temperate
Fig. 1.10 Middle Permian biomes (adapted from Willis and McElwain (2002) on a TimeTrek
4.2.5 model, Cambridge Paleomap Services)
is one of the keys that supported the continental drift theory of Alfred Wegener.
Botrychiopsis, another typical species from west Gondwana, went extinct when the
environmental conditions typical of a greenhouse stage were created by the end of
the Permian (Jasper et al. 2003).
The Permian flora of Gondwana was significantly more diversified than the one
of the Late Carboniferous (Cúneo 1989), and the floristic provinciality changed dur-
ing the course of the Permian. The belt located between 60◦ and 45◦ S in western
Gondwana was called the “Southern temperate semiarid belt of middle latitudes”,
characterized by Glossopteris and moderately thermophilic vegetation with abun-
dant tree-ferns and lycopods (McLoughlin 2001; Chumakov and Zharkov 2003)
(Fig. 1.10).
1.2.2 The Mesozoic (251–65.5 mya)

The transition from the Palaeozoic to the Mesozoic is characterized by a dramatic
event: the Permian-Triassic extinction event, which apparently saw the destruction
of 90% of marine life on Earth due to extensive volcanism, under other causes
(Benton and Twitchett 2003). The impacts on the terrestrial ecosystem were not
so drastic, or paradoxically even favorable for some plants (Looy et al. 2001).
The Triassic (251–199.6 mya) climate was relatively warm compared to today,
and continentality and aridity were more extended due to the permanence of the
single continent Pangaea. The Triassic flora remained broadly similar to that of the
Permian, dominated by gymnosperms (seed ferns, cycads, and ginkgos). During
the Triassic, Glossopteris-dominated communities were replaced by Dicroidium
(a seed fern) dominated floras across the Southern Hemisphere (McLoughlin 2001).
Also, the major radiation of conifers, e.g. the Araucariaceae began in the Triassic
(see Sect. 2.2). Other important components of the southern flora were ginkgo-
phytes, putative gnetales, bennettitales, and cycadales, plus many lycophytes and
osmundacean, gleicheniacean, dicksoniacean, dipteridacean and marattiacean ferns
(McLoughlin 2001, p 286; Artabe et al. 2003) (see Sect. 2.2).
The Jurassic (199.6–145.5 mya), better known for the diversification of charis-
matic faunal groups like the dinosaurs, is also considered one of the most important
periods in plant evolution. By the Early Jurassic, both composition and distribu-
tion of southern hemisphere vegetation had changed dramatically. Glossopteris and
Dicroidium no longer dominated the southern flora. Instead they were replaced by
cycads, bennettites, ginkgos, and conifers, and for the first time global floras con-
tained a significant portion of forms that are recognizable in our present floras. The
floral assemblage for Cerro La Brea, Mendoza, Argentina (Early Jurassic) shows the
presence of 14 taxa belonging to the Equisetaceae, Asterothecaceae, Marattiaceae,
Osmundaceae, Dipteridaceae, and several conifers (Artabe et al. 2005).
While Gondwana drafted towards the equator, five distinct biomes settled dur-
ing the Early Jurassic (McAllister Rees et al. 2000) (Fig. 1.11). Southern South
America must have been occupied by a “winterwet biome” with a climate similar
Subtropical desert
Winterwet
Warm temperate
Fig. 1.11 Early Jurassic biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5
model, Cambridge Paleomap Services)
to that of today’s Mediterranean-type one. The relatively increased proportion of

plants with small leaves and other xerophytic features clearly indicates seasonal
water deficits (Willis and McElwain 2002). In the Middle Jurassic, main compo-
nents of this biome, like Cycadales, Bennettitales, conifers, ferns, and Sphenopsids,
reached northernmost Chile, i.e. current arid Atacama (Fuenzalida 1966; Herbst and
Troncoso 1996).
Quattrocchio et al. (2007) listed more than a hundred species from the Jurassic of
the Neuquén basin, Argentina. Clearly dominant groups were the Cheirolepidiaceae,
Araucariaceae and Podocarpaceae, together with Cyatheaceae, Osmundaceae,
Marattiaceae, Dipteridaceae, Lycopodiaceae, Schizaeaceae, Anthocerotaceae,
Ricciaceae, Cycadales/Bennettitales, Caytoniaceae and Gnetales. The authors
further propose an environmental model in which the Araucariaceae and
Podocarpaceae occupied mostly high-altitude places, while ferns, cycads and
Cheirolepidiaceae may have been restricted to more low-lying and humid places
(Fig. 1.12). Let us keep in mind that there was still not such thing like an elevated
Andes (Box 1.5)
Cycadales Podocarpaceae
Cheirolepidiaceae Araucariaceae
v
v
Lycopodiaceae and ferns v v
v
v v v
v v v v
v v v
v Igneous rocks
v v v v
v v v
v v v
v v v v
Marine deposits v v v Continental v
v deposits v v v
v v v v v v
v v v v v v v v
v v v v
Fig. 1.12 Palaeoenvironmental reconstruction of middle Jurassic flora from southern South
America (adapted from Quattrocchio et al. 2007, with permission of the authors)
Box 1.3 Gondwana Breaks-Up

Most authors recognize three major separation events of Gondwana that
affected the evolution of the South American flora: the separation between
W and E Gondwana during the Jurassic (180–150 mya); the separation
America/Africa between 119 and 105 mya, and the split between Antarctica
and southern South America (32–28 mya) (Table 1.1). These ages serve as
reference; but there is no real consensus on the time of fragmentation of the
different components. The crucial separation of Australia from Antarctica
and South America from Antarctica and the development of the Drake
Passage is still a controversial issue: “South America may have separated
from Antarctica as early as the Late Jurassic, or as late as the Palaeocene
or Eocene” (Orme 2007, p 10) (see Box 9.1). The TimeTrek model shows
indeed an early separation of South America and Antarctica at around 120
mya (Early Cretaceous) (Fig. 1.7).
Table 1.1 Three stages in the break-up of Gondwana (as resumed by McLoughlin 2001)
Major separation Palaeoreconstructions on a TimeTrek

events Period and causes v. 4.5.2 model
(W Gondwana / During Middle to Late Jurassic

E Gondwana) (180–150 mya): breakup
associated with development
of a series of deep seated
mantle plumes beneath the
extensive Gondwanan
continental crust in S Africa
(c 182 mya) and the
Transantarctic mountains
(c 176 mya) (Storey 1995)
Africa–S Early Cretaceous (119–105

America mya): opening of the South
separation Atlantic Ocean, due to the
emplacement of
Plume-related
Parana-Etendecka
continental flood basalts in
Brazil and Namibia
(137–127 mya). Final
break-up of Africa and S
America was completed only
at 80 mya
West Early Oligocene (ca 30 mya):

Antarctica-S beginning at ~35–30.5 mya
America as a subsidence in the Powell
Basin followed by seafloor
spreading. Opening of the
Drake Passage between the
southern tip of South
America and the northern
end of Antarctic Peninsula
allowed deep water
circulation and the
installation of the Antarctic
Circumpolar Current (ACC)
between 41 and 24 mya (see
Box 9.1)
Southern Floras during Early Cretaceous did not differ much from the
Late Jurassic ones (Fig. 1.13). Most famous is the middle Cretaceous,
known as the period of expansion and radiation of the angiosperms (see
also Box 2.4). Angiosperms evolving during this time include a number of
18
Fig. 1.13 Illustration of the biotic assemblage from the limit Jurassic/Cretaceous (145.5 mya) of the Southern Cone. Theropod dinosaur on a swamp surrounded
1 The Extravagant Physical Geography of Chile
by ginkgos, araucarias, and arborescent ferns (original illustration by Sergio Elórtegui Francioli)
families that constitute a significant part of the present-day global flora (e.g.
Betulaceae, Gunneraceae, Fagaceae/Nothofagaceae). For the early Late Cretaceous
(Cenomanian to Coniacian), Troncoso and Romero (1998) reported a Neotropical
flora showing a notable change compared to the previous ones. They reported
the definitive replacement of the dominance of gymnosperms by angiosperms,
including representatives of extant families, such as the Lauraceae, Sterculiaceae,
Bignoniaceae, and Monimiaceae; and from extant genera like Laurelia, Peumus,
and Schinopsis (this last genus is currently not present in Chile).
By the Late Cretaceous, (Campanian-Maastrichtian) Troncoso and Romero
(1998) reported a Neotropical flora with marginal presence of Nothofagus
(Campanian first appearance of Nothofagus in Antarctica; Maastrichtian first
appearance of Nothofagus in the fossil record from Central Chile and Tierra del
Fuego) (see also Chap. 9). In spite of its marginal presence, it is the peak of northern
expansion of Nothofagus in South America, reaching 30◦ S (Torres and Rallo 1981)
(Fig. 1.14). This expansion of Nothofagus is challenging since the Late Cretaceous
is considered a rather greenhouse world (Box 1.4). It is but possible that transient
small icecaps existed during this mostly warm period. It has been proposed that rela-
tively large and short-term global sea level variations may have been connected with
small and ephemeral ice sheets in Antarctica, probably related to short intervals of
peak Milankovitch forcing (Gallagher et al. 2008).
Southern South America, already isolated from the rest of western Gondwana,
was occupied mainly by a “subtropical desert” and a “warm temperate” biome
Tropical summerwet
Subtropical desert
Mid latitude desert
Warm temperate
Cool temperate
Fig. 1.14 Late Cretaceous biomes; arrow shows northernmost expansion of Nothofagus (see text)
(adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5 model, Cambridge Paleomap
Services)
(Fig. 1.14), the latter being characterized by Araucariaceae, Nothofagaceae,

Proteaceae, and Winteraceae (Willis and McElwain 2002). “The presence of tropi-
cal elements in the austral margin of South America gives support to the expansion
of a warm climate towards high latitudes during the mid Cretaceous” (Barreda
and Archangelsky 2006). Troncoso and Romero (1998) also reported the pres-
ence of Neotropical palaeofloras in the mid- and Late Cretaceous from Magallanes
and Tierra del Fuego. Microfossils assigned to the Arecaceae (Palmae) have been
reported since the Maastrichtian (Hesse and Zetter 2005).
Box 1.4 Floral Extinction at the K/T Boundary ?

A permanent question is whether massive extinction events that mostly
affected the terrestrial fauna affected as well the global flora (McElwain and
Punyasena 2007). It seems that at the K/T boundary, at least several groups
suffered similar luck than dinosaurs, plesiosaurs, and ammonoids. For exam-
ple, the seed-ferns, a group that dominated the vegetation formations in many
parts of the world from the Triassic to the Cretaceous, are considered to have
disappeared at the end of the Cretaceous. Neverthless, exceptions are the rule,
and there is a seed-fern fossil recent discovered in Tasmania that has been
dated from the Early Eocene (McLoughlin et al. 2008).
Recent findings on the Lefipán Formation in NW Chubut province dated
as Maastrichtian, supports the catastrophic character of the K/T boundary
(Cúneo et al. 2007). The discovery of a highly diversified assemblage of
dicot leaves with probably more than 70 species, as well as several mono-
cots, podocarp conifers, and ferns, suggests that the latest Cretaceous floras
were probably more diverse than those known from Patagonia during the
Palaeocene. This means that the K/T event indeed affected the terrestrial
ecosystems of southern latitudes. The recovery of floral diversity must have
taken most of the Palaeocene until the recovering of plant richness by the early
Eocene (Cúneo et al. 2007).
1.2.3 The Cenozoic (65.5 mya Onwards)

The deep-sea oxygen isotope record permits a detailed reconstruction of the
Cenozoic global climate, that has suffered a number of episodes of global warm-
ing and cooling, and ice-sheet growth and decay (Zachos et al. 2001) (Fig. 1.15).
The most pronounced warming occurred from the Mid-Palaeocene (59 mya) to the
Early Eocene (52 mya), showing a peak in the so called Early Eocene Climatic
Optimum (52–50 mya) (Fig. 1.15). This period was one of the warmest periods in
the Earth’s history: temperature estimates of between 9 and 12◦ C higher than present
have been proposed (Zachos et al. 2001). This optimum was followed by a trend
toward cooler conditions in the Late Eocene. According to Zachos et al. (2001), ice-
sheets appeared in the Early Oligocene, and persisted until a warming phase that
reduced the extent of Antarctic ice in the Late Oligocene Warming (Fig. 1.15). From
this point (26–27 mya) until the middle Miocene (15 mya), the global ice volume
remained low with the exception of several brief periods of glaciation. This warm
phase peaked in the Middle Miocene Climatic Optimum (17–15 mya), and was fol-
lowed by a gradual cooling and reestablishment of a major ice-sheet on Antarctica
towards the Plio/Pleistocene (Zachos et al. 2001) (Fig. 1.15).
In the Early Palaeocene (~65–55 mya) the global position of South America
had moved close to the present-day position (Fig. 1.14). Nevertheless, the cold cir-
cumpolar ocean current had not yet developed, and Pacific Ocean currents carried
heated tropical waters to high latitudes. As a consequence, a permanent ice cover at
the poles was absent, and the prevailing low relief of the continents, coupled with
high seas, resulted in rain-bearing winds penetrating far into the interior of all the
main landmasses (Willis and McElwain 2002).
South America was mainly occupied by “tropical everwet”, “subtropical desert”
and “warm temperate” biomes. The warm temperate biome was composed of ever-
green and deciduous dicots (e.g. Nothofagus), and podocarps. South of 70◦ S, and
widespread in Antarctica, a “warm cool temperate biome” was established, com-
posed mainly by Araucaria, Podocarpus, Dacrydium, evergreen Nothofagus, and to
a minor extent members of the Loranthaceae, Myrtaceae, Casuarinaceae, Ericaceae,
Liliaceae, and Cunoniaceae (Truswell 1990).
Troncoso and Romero (1998) emphasized the neotropical character of
the Palaeocene palaeofloras of Central and Southern Chile. Zonal vegeta-
tion was composed mainly of rainforests with palms, mangroves, and in the
higher parts, azonal vegetation composed of Gymnosperms (Cheirolepidaceae,
Araucariaceae, Podocarpaceae, Zamiaceae) and Nothofagus, accompanied by
Myrtaceae, Proteaceae and Lauraceae. Fossil Boraginaceae related to extant Cordia
Early Eocene
Climatic Optimum
Temperature (°C) above present day
12
Late Oligocene
Warming
8 Middle Miocene
Late Palaeocene
Thermal Maximum Climate Optimum
4 West Antarctic
Ice-sheet
0 Glaciation Glaciation East Antarctic

Ice-sheet
K Palaeoc. Eocene Oligocene Miocene Plio/Pleist.
70 60 50 40 30 20 10 0
Age (mya)
Fig. 1.15 Global climatic fluctuations during the Cenozoic, based on global deep-sea oxygen and
carbon isotope records (adapted from Zachos et al. 2001)
have been described by Brea and Zucol (2006) from the Late Palaeocene of Chubut,
Argentina. A rich assemblage of micro- and megafossils has been described by
Troncoso et al. (2002) from the Ligorio Márquez Formation in Aisén (47◦ S). Of the
twenty leaf species reported, fourteen are from the Lauraceae; the rest corresponding
to the Melastomataceae, Myrtaceae, Sapindaceae, and others. Furthermore, seven
Pteridophyta, two conifers, and four angiosperms are represented by palynological
species. In spite of this predominantly tropical character, the presence of temperate
taxa like Nothofagus and Podocarpaceae confirms the warm temperate tendency at
47◦ S (Okuda et al. 2006).
Recently Iglesias et al. (2007) reported a greater species richness than was
previously known from Palaeocene Patagonia, including more than 43 species of
angiosperm leaves. At the end of the Palaeogene, representatives of most of the
angiosperm modern classes and many orders were already present in southern South
America (Gandolfo and Zamaloa 2003; Prámparo et al. 2007).
Eocene (55.8–33.9 mya) floras of Southern South America show subtropical
to fully tropical forests, with zones of seasonal dryness in Chile (Romero 1986).
The three extant South American tribes of the Proteaceae were already present
in the early Eocene, forming the Australia-Antarctica-South America connection
(González et al. 2007). Late Eocene fossil leaves, flowers and fruits assigned to the
Escalloniaceae have also been reported as being involved in this austral connection
(Troncoso and San Martín 1999).
Remarkable is the presence of Eucalyptus macrofossils in the Patagonian Early
Eocene (Gandolfo et al. 2007), since the genus shows an extant distribution
in Australasia, mainly Australia and Tasmania (not New Zealand). The South
American macrofossils reported by Gandolfo et al. (2007) are to date the most
ancient register for the genus.
The Laguna del Hunco palaeoflora in NW Chubut, Argentina, shows the most
complete example of Early Eocene vegetation in South America. This palaeoflora
is one of the world’s most diverse Cenozoic assemblages of angiosperms (Wilf
et al. 2005, 2007). This assemblage comprises tropical elements restricted today
to temperate and tropical Australasia (e.g. Dacrycarpus, Papuacereus, Eucalyptus);
tropical elements (e.g. Roupala, Bixa, Escallonia), and the disjunct element South
America/Australasia (e.g. Eucryphia, Orites, Lomatia) (see Fig. 3.5). Fossil plants
at Laguna del Hunco are extremely abundant, diverse (>150 leaf species), and well-
preserved. During the early Eocene the area was a subtropical rainforest with land
connections both to Australasia via Antarctica and to the Neotropics (Fig. 1.16).
Wilf et al. (2007) suggest that the Laguna del Hunco plant lineages retreated
to geographically disparate rainforest refugia following post-Eocene cooling and
drying in Patagonia. Only few lineages adapted and persisted in temperate South
America.
The continuous decrease in temperature during the Eocene allowed a new dis-
placement of Nothofagus towards South-Central Chile. Therefore this time-span is
characterized by a mixed tropical-subantarctic palaeoflora (Troncoso and Romero
1998). In spite of the prevalence of mixed palaeofloras during the Eocene, results
obtained by Gayó et al. (2005) at Bahía Cocholgüe (36,5◦ S) suggest that tropical
floras persisted in central Chile during the Early Eocene and formed a belt between
Tropical summerwet
Subtropical desert
Warm temperate
Cool temperate
Fig. 1.16 Early Eocene biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5
model, Cambridge Paleomap Services)
25◦ S and 37◦ S. This persistence of tropical floras (composed mainly by Lauraceae
and Myrtaceae) might be related to the influence of the Early Eocene Climatic
Optimum (Fig. 1.15) and to a shrinking tropical belt (Gayó et al. 2005).
The transition from the Eocene to the Oligocene (33.9–23.03 mya) was a period
of significant global climatic cooling and increased aridity, major changes in oceanic
circulation, and the initiation of ice on Antarctica (Zachos et al. 2001; Convey
et al. 2008) (Fig. 1.15). Major reorganization and redistribution of global vege-
tation followed these climatic trends, with a reduction of tropical forests and the
expansion of temperate vegetation toward the equator (Willis and McElwain 2002).
A Subantarctic palaeoflora expanded its distribution range across southern South
America, occupying an area that became to extend from the island of Tierra del
Fuego to the south of Central Chile (Romero 1993) (Fig. 1.17).
This implicates the massive retreat of tropical and subtropical compo-
nents from the Sapindaceae and Lauraceae, the generic replacement of gen-
era in the Rhamnaceae, Myrtaceae, Bignoniaceae, Flacourtiaceae/Salicaceae;
and the regional extinction of several families like Moraceae, Annonaceae,
Dilleniaceae, Malpighiaceae, Vochysiaceae, Tiliaceae, Sterculiaceae, Sapotaceae,
and Styracaceae (Troncoso and Romero 1998). Permanent ice sheets persisted on
Antarctica until the Late Oligocene (26–27 mya), when a warming trend reduced
the extent of Antarctic ice (Zachos et al. 2001).
From the “Late Oligocene Warming” (26–27 mya) (Fig. 1.15) until the Middle
Miocene (~15 mya), the global ice volume remained low and water showed
slightly higher temperatures, intermingled with brief periods of glaciation (Zachos
Tropical everwet
Subtropical summerwet
Woody savannah
Warm/cool temperate
Cool/cold temperate
Cold temperate
Ice sheet
Fig. 1.17 Early Oligocene biomes; arrows show mixture of tropical and austral floras (adapted
from Willis and McElwain (2002) on a TimeTrek 4.2.5 model, Cambridge Paleomap Services)
et al. 2001). This was followed by a gradual cooling and reestablishment of a

major ice-sheet on Antarctica by 10 mya (Fig. 1.18). The continental interior
became increasingly arid/cold and large areas of shorelines were exposed due
to a falling sea level. Outside the core a depauperate “cold temperate” biome
survived, having lost its main forests components and with some herbs and C3
grasses remaining. During the Late Miocene most of Western South America was
occupied by a “cool temperate” biome. The “winterwet” and “subtropical summer-
wet” biomes were restricted to a reduced proportion of today’s Atacama Desert
(Fig. 1.18).
The Miocene (23.03–5.33 mya) is characterized by a development of mod-
ern angiosperm families like Asteraceae, Poaceae, Malvaceae, Fabaceae and
Cyperaceae, related to more open communities replacing tropical forests in southern
South America (Barreda et al. 2007; Palazzesi and Barreda 2007). Tropical forests
were still abundant during the Early Miocene of Patagonia; the vegetation increas-
ingly acquired a more complex aspect, due to the wider distribution of grasses and
shrubs. Barreda et al. (2007) list 60 angiosperm families present in the fossil record
during the Miocene in Argentina. More diversified families are the Malvaceae (33
taxa), Fabaceae (32), and Asteraceae (25).
During the Early and Middle Miocene, the subantarctic flora reached the south-
ern part of Central Chile (Troncoso and Romero 1998). In the middle Miocene of
Central Chile a change from the previous subantarctic palaeoflora into a mixed
palaeoflora with a predominance of neotropical taxa and the retreat of subantarc-
tic taxa occurred (Hinojosa 2005). The subsequent subtropical palaeoflora that
Tropical everwet
(woody savannah)
Warm temperate
Cool temperate
Cold temperate
Ice sheet
Fig. 1.18 Miocene biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5 model,
Cambridge Paleomap Services)
occupied central Chile during the lower to mid Miocene, 20–15 mya, developed
under a warmer and more humid palaeoclimate, with an incipient Andean rain
shadow effect, is the nearest ancestor of the sclerophyllous modern vegetation of
central Chile (Hinojosa et al. 2006).
In northern Chile, the hyperarid climate became established at the Oligocene/
Miocene boundary (ca. 25 mya) (Dunai et al. 2005; Nalpas et al. 2008), and was
followed by more humid (semiarid periods interrupted by short arid events up to the
earliest Late Pliocene (Hartley and Chong 2002, Box 3.1)).
Global deep-sea oxygen and carbon isotope records indicate additional cooling
and small-scale icesheet expansion on west-Antarctica during the Late Miocene
until the Early Pliocene (6 mya). The early Pliocene is marked by a subtle warm-
ing trend between 3.3 and 3 mya. Afterwards cooling again increased (Zachos et al.
2001).
Box 1.5 Slow or Rapid Andean Uplift?
The Late Miocene has been proposed as the initial phase of the Andes uplift.
Gregory-Wodzicki (2000), on the base of palaeobotanical data, proposed a
surface uplift in the order of 2,300–3,400 m asl since the late Miocene at
uplift rates of 0.2–0.3 mm/year. More recently Ghosh et al. (2006) obtained
results that indicate a surprisingly rapid uplift of the Bolivian Altiplano at
an average rate of 1.03 ±0.12 mm per year between ~10.3 and ~6.7 mya (i.e.
from 0 to 4,000 m asl since the Middle/Late Miocene). These results challenge
the known forces responsible for the uplift and are in conflict with geological
evidence (e.g. Hartley 2003, proposed a proto-Central Andean mountain range
placed between 15 and 9 mya). Geomorphological evidence, i.e. lahar deposits
in the Coastal Cordillera of central Chile (33◦ 40′ –34◦ 15′ S) still supports an
Oligocene–Miocene uplift of the Andes (Encinas et al. 2006). New findings by
Garzione et al. (2008) and Hoke and Garzione (2008) based on isotope data,
suggest that the Andes elevation remained relatively stable for long periods
(tens of millions of years), separated by rapid (1 to 4 million years) changes
of 1.5 km or more.
Most families already present in the mid to late Miocene continue to be present
during the Pliocene (5.33–1.81 mya) (e.g. Arecaceae, Lauraceae, Myrtaceae,
Anacardiaceae, Asteraceae, Chenopodiaceae/Amaranthaceae). Several families like
the Fabaceae increasingly diversified (Barreda et al. 2007).
Southern South America was dominated by grasslands, steppes, and shrublands,
with rainforests restricted to the moist temperate forests of south-western Patagonia
(Dowsett et al. 1999; Haywood et al. 2002) (Fig. 1.19). Barreda et al. (2007)
Tropical everwet
Woody savannah
Desert (hyperarid / arid)
Cool temperate
Cold temperate
Ice sheet
Andes
Fig. 1.19 Pliocene biomes (adapted from Dowsett et al. (1999) and Haywood et al. (2002), on a
TimeTrek 4.2.5 model, Cambridge Paleomap Services)
recognize a neotropical palaeo-floristic province from 32◦ S to the north and a proto-
espinal/steppe province to the south, together with a Nothofagacean province at the
southwest.
The Cerro Centinela palaeoflora in Central Chile contains representatives of
more than 20 modern families of different affinities: tropical genera not found in
Chile any longer (e.g. Nectandra Ocotea, Miconia), subtropical genera (Schinus,
Schinopsis, Acacia) and temperate or austral genera (Araucaria sección Eutacta,
Nothofagus) (Troncoso and Encinas 2006). Appealing is the presence of the
fern genus Dicksonia, found today mainly in Australasia (Malesia, New Guinea,
Australia, New Caledonia) and in Juan Fernández (Chap. 5).
The end of the Cenozoic, traditionally treated as the “Quaternary”, has been
divided into two epochs: the Pleistocene (1.8 mya to 11,500 year BP) and the
Holocene (11,500 year BP to the present). The Pleistocene is vastly known as the
most recent epoch of glaciations. The last Pleistocene glaciation cycle is known
from southern Chile as the Llanquihue glaciation, which is correlated with the
Wisconsin/Weichselian glaciations in the northern hemisphere, according to global
cooling data (Andersen et al. 1995; Lowell et al. 1995; Moreno et al. 2001).
In Chile glaciations affected to some extent all the ecosystems ranging from
the arid north to the humid south. Several proxy-data used for Quaternaty palae-
oreconstructions are specific for each environment: rodent middens in the north
(Betancourt and Saavedra 2002; Maldonado et al. 2005), tree rings in central Chile
(Barichivich et al. 2009), and sediment cores containing fossil pollen in the for-
merly glaciated south (Heusser 2003; Moreno 2004) (Box 1.6). The pollen analysis
is complemented with the study of macrofossils (leaves), beetles (e.g. Ashworth
et al. 1991), and more recently, chironomid stratigraphies (Massaferro et al. 2009).
Box 1.6 Six Steps for Palaeoenvironmental Reconstruction

Hereafter the main steps for getting pollen sample cores are briefly exposed
(arbitrary extracted from Heusser 2003, Chap. 10):
1. Select a suitable site for sampling, on the base of aerial photographs and
topographic maps, referring to the glacial borders and little sedimentary
bogs or mires. Take account of accessibility for heavy coring equipment.
2. Get a piston sampler equipped with core tubes 5 cm in diameter and 1 m
in length, with 1.5 m long extension rods.
3. With the piston sampler managed by three to four people, get the samples.
You may need to build a wooden platform on the bog and a chain host to
lift the sampler to the surface.
4. Extrude increments onto clear plastic, examine and describe them (color,
texture, layers) and wrap them in aluminum foil. Take multiple cores at
each coring location to ensure overlap at core breaks.
5. Once in the laboratory, identify the pollen grains under a microscope, at

every < 5 cm interval in every sample core.
6. Voilá! You are ready to begin your own palaeoenvironmental
reconstruction.
Note: Calvin and Linda Heusser, together with an international research team,
worked for more than 40 years in southern Chile. They could get thousands
of samples from 50 coring sites to reconstruct the glacial history and discern
the palaeoecological factors responsible for vegetation changes over 50,000
years.
Calvin Heusser and coring team at Taiquemó site (Chiloé) in the late nineties. From left
to right: Tom Lowell, Patricio Moreno, Linda Heusser, Calvin Heusser, David Marchant
(Photo A. Moreira-Muñoz)
Glaciation effects were especially drastic from 42◦ (Chiloé) southward, were
glaciers and ice lobes virtually devastated the temperate forests at the Last Glacial
Maximum (LGM) between 29,400 and 14,450 year BP (Fig. 1.20). Vivid remnants
of this widespread glaciation are the Campo de Hielo Patagónico Norte and Campo
de Hielo Patagónico Sur, together with Cordillera de Darwin in southernmost
Patagonia (Fig. 1.20).
At the LGM, periglacial effects like solifluction and glaciofluvial activity also
should have affected the Andes, the longitudinal depression, and the coastal
Cordillera between 39 and 43◦ , affecting principally the Valdivian and evergreen
northpatagonian forests (Heusser 2003).
Glacial conditions forced forest formations to migrate equatorward and tree-
lines to lower in altitude (Villagrán et al. 1998; Heusser 2003). Vegetation close
76° 68°
–30°
LGM ice extension
Campos de Hielo
Distribution
Past New
–40°
Drapetes
Huperzia
2 –50°
0 250 500 km
Fig. 1.20 Maximal extension of the last cycle of the Llanquihue glaciation (after Denton et al.
1999; Heusser 2003). Remnants of the Pleistocene glaciations: (1) Campo de Hielo Patagónico
Norte, (2) Campo de Hielo Patagónico Sur, (3) Cordillera de Darwin. Also shown is the past and
current distribution of Huperzia fuegiana and Drapetes muscosus (adapted from Heusser (2003)
and Moore (1983), and collections of the National Herbarium SGO)
to the glaciated areas was structurally open, forming a steppe-tundra and turning
to parkland and open woodland towards north-central Chile. In the northern part
of the Central Depression (Tagua Tagua, 34.5◦ S), at ~14,500 year BP, Lateglacial
warmth and dryness induced the retreat of Nothofagus-Prumnopitys woodland first
by a spread of grassland and ultimately by herb-shrub communities composed by
xeric Amaranthaceae and Asteraceae (Heusser 1997). The presence of Nothofagus

dombeyi type pollen until ~10,000 year BP in the Central Depression exemplifies the
downward altitudinal migration of taxa: this species is today restricted to the Andes
at this latitude, which is also its northern distribution limit (see Sect. 9.1, Fig. 9.5).
Similar situation was suffered by conifers in the south: the current disjunct range of
several species in both cordilleras is a relict of a formerly wider distribution (before
the colder period at 30,000–14,000 year BP), as shown by the (fossil) presence of
Fitzroya and Pilgerodendron in the Central Depression (Villagrán et al. 2004).
Termination of the last glaciation was (differentiated locally) more or less at
15,000 year BP. Subantarctic species at low altitude in Los Lagos-Chiloé region,
Like Lepidothamnus fonkii (Podocarpaceae), Astelia pulima (Asteliaceae) and
Donatia fascicularis (Stylidiaceae), migrated to higher altitudes. Other species like
Huperzia fuegiana (Lycopodiaceae) and Drapetes muscosus (Thymelaeaceae) were
pushed to the south and are today restricted to southernmost Patagonia or Fuegia
(Fig. 1.20).
The impacts of the Quaternary glaciations were not restricted to the southern
Andes at all, and affected also the vegetation in the Central Chilean Andes (Villagrán
et al. 1998). The changes on vegetation associated to the last glaciations cycle, as
emphasized by these authors are:
(1) Existence of disjuct populations of conifers (Fitzroya, Araucaria, Prumnopitys,

Austrocedrus) on the Costal cordillera between 37 and 42◦ S, dissociated
from the main distribution on the Andes (Fig. 2.18). These are remnants of a
widespread distribution of these species in the central depression when the cli-
mate was cooler. Same for the Magellanic moorlands located on the summits of
the Cordillera de la Costa.
(2) Current concentration of endemic and monotypic species between 36 and 40◦ S
has been interpreted as the function of refugia for laurifolious forests during the
Pleistocene.
(3) High level of endemism at species level showed by the extant high Andean flora
of central Chile is the consequence of repeated cycles of isolation associated to
the adavance of Andean glaciers during the Pleistocene.
(4) The current presence of Andean scrub communities in the Coastal cordillera
between 32 and 34◦ , could be the expression of a range expansion of these
communities experimented during the last glaciaton.
The Andes of Central Chile were locally glaciated on the tops of the mountains,
and it is presumed that the Andean flora descended altitudinally and also migrated
northwards during the Pleistocene (Simpson 1971; Villagrán et al. 1998).
Especially the altitudinal migrations are hypothesised to have affected commu-
nities as a whole, but there is also possible that there was a recombination and
redistribution of the taxa forming today these communities (Patricio Moreno pers.
comm.). Whether the Quaternary glaciations affected only the distribution ranges
or had deeper evolutionary implications is still a matter of discussion (Box 1.7).

As example, as the glacial tongues advanced down from the Andes into the cen-
tral depression, valleys like the Río Maule (36◦ S) and Río Biobío (37◦ to 38◦ S)
could have acted as barriers, interrupting gene flow between plant populations
and communities, as resulting from the study of Hypochaeris acaulis populations
(Tremetsberger et al. 2003).
Box 1.7 How to Survive a Glaciation? The Refugial Debate

The Campos de Hielo Norte and Sur are considered the biggest inland ice-
caps after Greenland, current reminders of the maximal extension of the
Pleistocene glaciations. Figure 1.20 suggests almost complete depletion of the
southern biota at the LGM. But the relative rapid reoccupation of deglaciated
areas under warmer conditions by the rainforest taxa suggests the continued
permanence of exemplars somewhere not so far from the glacial lobes.
Debate continues on possible impacts from the ice ages and possible loca-
tions of refugial sites in the Southern Andes (Knapp and Mallet 2003). As
2/3 of the actual area of the southern forests was depleted, the traditional
view proposes that taxa mostly survived the glaciations in the foreland of the
glaciers and on several nunataks. This view has been recently challenged by
Fickert et al. (2007), who suggest, based on research on six active glaciers
(e.g. Monte Tronador in southern Chile), that the size of possible refugia
would be considerably enlarged if debris-covered glaciers are considered.
The nunatak theory just offers a too small area for a survival of viable plant
populations (Fickert et al. 2007). Debris-covered glaciers should be added to
the recent systematization of three main types of glacial refugia proposed by
Holderegger and Thiel-Egenter (2009): i.e. nunatak, peripheral and lowland
refugia. Concrete results provided by Premoli et al. (2000) suggest that the
populations of Fitzroya cupressoides survived the Last Glacial Maximum in
multiple refugia rather than in only one refugium, such as an ice-free area of
coastal Chile (Single Refugium hypothesis). Multiple refugia in the eastern
side of the Cordillera are also hypothesized for the survival of Austrocedrus
chilensis (Pastorino and Gallo 2002).
Pleistocene and Holocene changes have disrupted species ranges, extir-
pated local populations, and changed selective pressures (Premoli et al. 2000),
but it is doubtful that they affected speciation processes. Some authors have
emphasized the role of the last glaciations in speciation, but others call this a
“failed paradigm” (Klicka and Zink 1997). It is possible that molecular stud-
ies of populations of Quaternary species help define the relict characteristic of
these species (Willis and Niklas 2004) (see Sect. 9.6).
1.3 Current Climate and Vegetation

A few crucial features of the present climate and vegetation will be outlined here, as
a detailed description of these aspects is beyond the scope of this book. The reader
is redirected to the most updated references on these topics, especially Luebert and
Pliscoff (2006), Veblen et al. (2007), Garreaud et al. (2009). A fine synthesis of the
southern Andean vegetation is also provided by Heusser (2003).
South America is situated within the influence of the Intertropical Convergence
Zone (ITCZ) and related circulation systems (Orme 2007). This band is a major
feature of the global circulation and the Chilean climate certainly depends upon
this regional situation. The yearly N-S displacement of the Subtropical Anticyclone
(South Pacific High) is one of the principal factors affecting the climatic latitudinal
gradient in Chile (Garreaud and Muñoz 2004; Emck et al. 2006) (Fig. 1.21).
The displacement of the South Pacific High towards the south during the austral
summer promotes Mediterranean-type climatic conditions in Central Chile (Luebert
and Pliscoff 2006). During this displacement, tropical rainfall can reach the north-
ern Altiplano, but this influence decreases at around 23◦ S (Vuille and Baumgartner
1998) due to the influence of the Andes as a “climatic wall” (Garreaud and Aceituno
2007). The displacement of the Anticyclone towards the North in the austral winter
allows the entrance of the westerlies from the SW, promoting an intense and reg-
ular rainfall period with a maximal influence at around 47◦ S (Luebert and Pliscoff
2006). South of this latitude, the influence of the high pressures located around
Antarctica generates a decrease in precipitation (Endlicher and Santana 1988). From
Antarctica comes another important feature of the Chilean climate, i.e. the cool
Humboldt Current derived from the Antarctic Circumpolar Current (Orme 2007).
January (southern summer) July (southern winter)
South Pacific and South Atlantic Highs Relative position of the ITCZ
Polar front Surface dominant winds
Fig. 1.21 Atmospheric circulation over South America. Note the yearly N–S displacement of the
Intertropical Convergence Zone (ITCZ) (adapted from Emck et al. 2006)
1.3 Current Climate and Vegetation 33
The Humboldt Current is characterised by a predominant northward flow of sur-

face waters of subantarctic origin and by strong upwelling of cool nutrient-rich
subsurface waters of equatorial origin (Thiel et al. 2007). The current produces a
decrease in the coastal zone temperatures in North- and Central Chile, and aided
by tropospheric subsidence creates a temperature inversion that increases the con-
ditions of aridity in western South America from near the equator to beyond the
Tropic of Capricorn (Trewartha 1961; Orme 2007). Regarding precipitation, these
factors generate a gradient of decreasing rainfall from the SW towards the NE, from
ca. 5,000 mm in the SW to 0 mm in the Atacama (Fig. 1.22a). Only at the north-
ern Altiplano this tendency is reverted by the tropical rains reaching northern Chile
from the Amazonas basin.
1.3.1 Bioclimatic Zones

On the base of the early climatic classification done by W. Köppen (1930), Chilean
bioclimatic classifications were progressively developed by Di Castri (1968),
Quintanilla (1974), Di Castri and Hajek (1976), and Amigo and Ramírez (1998).
Coupling the pioneer plant geographical works of Reiche (1907) and Pisano
(1954), with the bioclimatic classifications and phytosociological information (e.g.
Oberdorfer 1960), several attempts have been made for a vegetation classification of
the country, like the ones from Schmithüsen (1956), Quintanilla (1983), and Gajardo
(1994).
A systematic revision of previous classification schemes integrated to the analy-
sis of global climatic surfaces on a GIS-based platform allowed Luebert and Pliscoff
(2006) to generate the most accurate bioclimatic and vegetation synthesis to date.
The classification considers bioclimatic and floristic data on the base of the concep-
tual framework promoted by Rivas-Martínez and Rivas-Sáenz (1996–2009). The
application of the method for Chile resulted in the identification of 17 vegetation
formations and 127 vegetational belts (Luebert and Pliscoff 2006). The highest rank
of the bioclimatic classification is the “macrobioclimate”, and the five units that
exist worldwide are found in Chile (Fig. 1.22b):
(a) Tropical macrobioclimate: it extends from the border with Perú at 17◦ 35′ S
towards the south till a diagonal limit at 23◦ S at the coast and at 31◦ S in the high
Andes. The southern limit reflects on the one side the maximal influence of the
polar fronts coming from the SW (Fig. 1.22b) and on the other side the maximal
influence of the tropical moisture that reaches the Altiplano and the north-
ern high Andes during the austral summer. Within this zone lies the Atacama
Desert.
(b) Mediterranean-type macrobioclimate: this macrozone is one of the most char-
acteristic features of Chile, as one of only five regions at the global scale that
share this type of climate, characterized mainly tdue to the marked seasonality in
the thermal and precipitation regimes (Di Castri 1981). The limits of this mac-
robiolimate in Chile have been controversial, but the proposal of Luebert and
Pliscoff (2006) is concordant with the findings of Amigo and Ramírez (1998). It
extends from the diagonal limit with the tropical bioclimate towards the South,
20°
a 2 1
30° 4
5
b
6
1
<10
7
10–50
40°
50–200
200–500 8a
500–1000
1000–3000 8b
3000–5000 c
> 5000 12
50°
10
11
d 9
a b e c
Fig. 1.22 a Annual precipitation, based on Schmithüsen (1956) updated with current available
data; b five Chilean macro-bioclimates (adapted from Luebert and Pliscoff 2006); c distribu-
tion of vegetation formations according Schmithüsen (1956): 1= Andean vegetation; 2= desert
core; 3= semi-desertic scrub; 4= xeric scrub and Fray Jorge fog-forest; 5= woody savanna;
6= sclerophyllous matorral; 7= deciduous (maulino) forest with conifers; 8a= Valdivian rain-
forest; 8b= northpatagonic rainforest; 8c=subantarctic rainforest; 9= subantarctic moorlands;
10=subantarctic deciduous forest, 11=east-patagonic steppe; 12=Campos de Hielo
till 37◦ S at coast and Andes, and till 39◦ S in the Central Depression. The
Mediterranean-type macrobioclimate appears also in disjunct patches further
South around 46–47◦ S, related to the western limit of the Patagonian steppe.
(c) Temperate macrobioclimate: it occupies the major area in continental Chile,
from the limit with the Mediterranean-type one at 37–39◦ S up to western
Patagonia and the Magallanes region in the southern territories. It is thermally
most homogeneous and precipitation can reach more than 5,000 mm on the
southwestern fjordland and island groups more exposed to the humid westerlies.
(d) Antiboreal macrobioclimate: it occupies a restricted portion of the southernmost
continental extreme, affecting the Magellanic archipelagos and the southern
part of Tierra del Fuego. Precipitation decreases notably toeads the east and
temperature decreases towards the south.
(e) Polar bioclimate: with increasing thermal cold conditions, it occupies the
territory outside the American continent towards Antarctica.
1.3.2 Vegetation Formations

Schmithüsen (1956) provided one of the most sinthetical and comprehensive
accounts of the Chilean vegetation, valid till today (Fig. 1.22c). He also illustrated
magistrally the latitudinal versus altitudinal distribution of the vegetation formations
(Fig. 1.23).
The principal vegetation formations and their main characteristics will be briefly
described here, based on Schmithüsen (1956) and Luebert and Pliscoff (2006)
(Figs. 1.22c, 1.23 and Fig. 1.24).
(a) The hyperarid desert formation or desert core extends from 18◦ S along the
coast and interior zones towards the south till around 24◦ . Approaching
the border with Peru, vegetation is restricted to the deep valleys of Azapa
North Snow limit Upper vegetation limit South

6000
5000
4000
3000
2000
1000
18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54 56
Andean vegetation Desert scrub Valdivian forest North Patagonian rainforest
Pre-Andean scrub Xerophytic scrub Austrocedrus chilensis Subantarctic rainforest and

moorlands
Browningia candelaris stands Sclerophyllous matorral Araucana araucana
Patagonian steppe
Deciduous forest Fitzroya cupressoides
Desertic core
Fig. 1.23 Distribution of vegetation formations along the altitudinal profile according
Schmithüsen (1956)
a b
c d
Fig. 1.24 Chilean vegetation: a high-Andean vegetation, Azorella compacta at Parinacota

(18◦ 30′ S); b high-Andean vegetation, vegas de Zapaleri, border with Bolivia at 22◦ 50′ S; c Puya
coerulea, characteristic of the xeric matorral at Cerro La Campana (33◦ S); d sclerophyllous
forest at Río Clarillo (33◦ 47′ S); e temperate forest at Navarino island (55◦ S); f conifer forest
(Araucaria) with Nothofagus at El Cañi, Pucón (39◦ 20′ S) (photo credits: a Walter Welss; b–f
A. Moreira-Muñoz)
and Camarones, related to agriculture. At the heart of the Atacama, vegeta-

tion is almost completely lacking; nevertheless, there are stands of natural
and human induced forests of Prosopis tamarugo = Pampa del Tamarugal.
Towards the Andes, scrub vegetation consists mainly of a low scrub (mator-
ral) of Adesmia atacamensis, Cistanthe salsoloides, Atriplex imbricata, and
Acantholippia deserticola. In a very thin belt between 2,000 and 2,800 m asl,
big cacti of Browningia candelaris bear out the landscape (Fig. 7.1).
(b) The sparse coastal shrub vegetation, also characteristic for coastal southern
Peru, is called the “Lomas” formation, and consists of a rich assemblage of
Eulychnia, Nolana, Heliotropium, Tetragonia, and Euphorbia species. This for-
mation is highly dependent on fog and humidity to some extent related to the
El Niño phenomenon (Box 7.1).
The desert coastal scrub extends into the land’s interior between 24 and
32◦ S, generating a transition zone from the desert towards the winter-wet
Mediterranean climate of Central Chile. It encompasses an open scrub (mator-
ral) composed by Adesmia spp., Bulnesia, Balbisia, and Heliotropium species.
At 30◦ S, the vegetation gradually changes to a xerophytic scrub composed of
Haplopappus spp., Porlieria chilensis, Flourensia thurifera, Colliguaja odor-
ifera, Trichocereus cacti, and Puya species. This zone harbours one of the
most interesting botanical phenomena, the “desierto florido” (Box 7.1), and a
plant geographical icon = the fog forest of Fray Jorge, where the northernmost
remnants of southern floristic elements are to be found (see Box 3.1).
(c) The Andean vegetation is the formation occupying extreme, high environments,
ranging from 17◦ 30′ S to ca. 40◦ S along the western Andean slope. This wide
latitudinal extension encompasses a very different composition along the North-
South profile, and in the altitude. The intermediate altitudinal belts show the
structurally most developed vegetation: the lowest belts are affected by the
aridity, the highest by low temperature. These intermediate belts are composed
by tolares, dominated by Parastrephia species and pajonales (cushion grasses)
of Festuca orthophylla. Stands of Polylepis tarapacana trees are to be found.
Precordilleran belts are dominated by Fabiana ramulosa and Diplostephium
meyenii. Above 4,700 m asl bofedales (high Andean wetlands) support the long
tradition of Andean llama and alpaca pasture. Approaching the most arid part
of the Atacama Desert there are shrubs of Fabiana and Baccharis, together with
Atriplex and Acantholippia in the lower belts. Towards the south, sparse vege-
tation is composed of Jarava frigida cushion grasses, several Adesmia species,
with the addition of Mulinum, and Urbania species. The treeline changes con-
stantly along the latitude gradient: in the north it is composed by queñoales =
Polylepis tarapacana and P. rugulosa, in the central-north it is replaced by
Adesmia shrubs with Ephedra between 31 and 34◦ S the treeline reappears
by means of Kageneckia angustifolia accompanied by Guindilia trinervis.
At this latitudinal range the Andean scrub is composed of Adesmia species,
Tetraglochin alatum, Mulinum spinosum, and cushion Apiaceae like Azorella
spp. and Laretia acaulis. From 32◦ 60′ to the South, at the lower limit of the
Andean formation, appears the conifer Austrocedrus chilensis (Fig. 2.18). The
Andean formation disappears at around 37◦ S, where it is replaced by deciduous

forests.
(d) Entering into the Mediterranean climate zone the vegetation changes to a sclero-
phyllous high scrub or matorral esclerófilo. On favorable South oriented slopes
this scrub shows characteristics of woodland, with trees reaching 20–25 meter
in height. Typical species of the Central Chilean matorral are Peumus boldus,
Cryptocarya alba, Quillaja saponaria, Maytenus boaria. In the quebradas,
i.e. more humid stands, there appears a more hygrophyllous forest composed
of Crinodendron patagua, Beilschmiedia miersii, Drimys winteri, and Persea
lingue. In contrast, the most exposed and plain areas contain a woody savanna
(espinal) mainly composed of Acacia caven and Prosopis chilensis. North
exposed slopes show a rich array of annual species and characteristic bromeli-
ads, Puya chilensis, P. berteroniana and P. coerulea, together with the cactus
Trichocereus chiloensis.
(e) Around 33◦ the coastal cordillera reaches far inside the continent, and above
1,200 m asl, the sclerophyllous woodland leaves space for a deciduous forest
composed of deciduous Nothofagus species. The northernmost populations at
33◦ S seem to be remnants of an ancient distribution of the genus (Chap. 9).
Deciduous forests dominate along the Andes and the coast towards the South,
surrounding the Central Depression. The core of the deciduous forest between
35 and 36◦ S is known as the maulino forest, a mesic forest type, dominated by
the two broadleaved deciduous species Nothofagus alessandrii and N. glauca
(San Martín and Donoso 1996). At around 38◦ S, this forest shows signs of
the transition towards the temperate macrobiolimate, with the remarkable pres-
ence of the resinous or conifer forests of Araucaria araucana at the coast
(Nahuelbuta) and the Andes. Deciduous forests turn often into a krummholz
of Nothofagus antarctica and N. pumilio composes the treeline along the Andes
all the way to the Cape Horn.
(f) Located well into the temperate macrobioclimate, and related to high precip-
itation levels (>2,000 mm/year) is the broad-leaved (laurifolious) forest, also
known as the Valdivian forest. It shows, same as the maulino forest, a “U”
shape with a coastal and an Andean branch between 39◦ and 42◦ S. The his-
tory of this forest has been vastly debated, and some of its components, like
Aextoxicon punctatum, Laureliopsis philippiana, Dasyphyllum diacanthoides
(Fig. 8.2), Luma apiculata, Laurelia sempervirens, Eucryphia cordifolia, and
Weinmannia trichosperma seem to be old remnants of Palaeogene floras (Sect.
1.2).
(g) At around 41◦ S on the Andes and 41◦ 30′ on the coast, broad-leaved forests
are replaced by an evergreen northpatagonian rainforest mainly composed of
large trees pertaining to the Nothofagaceae: Nothofagus dombeyi, N. nitida,
and N. betuloides. These rainforests are intermingled with the conifer forests
of Podocarpus nubigenus, Fitzroya cupressoides and Pilgerodendron uviferum.
These evergreen forests dominate at the coast and interior, being replaced in
altitude by the deciduous forest.
References 39
(h) As the landscape gets more and more fragmented into fjords and little islands
south of 47◦ , and the precipitation exceeds the 4,000 mm/year the vegeta-
tion turns to a low physiognomy of moorlands, dominated by Astelia pumila,
Donatia fascicularis, Oreobolus obtusangulus. Towards the East the moorlands
get less humid and dominated by the moss Sphagnum magellanicum. Most of
the interior of Patagonia is covered by the two wide icefielkds Campo de Hielo
Norte and Campo de Hielo Sur. To the South of this last icefield, the decid-
uous forest of Nothofagus reappears, together with the subantarctic evergreen
rainforest. In accordance with the marked precipitation gradient ranging from
4,000 mm at the western side to 300 mm at the eastern side of the low Andes in
southern Patagonia and Tierra del Fuego, a gramineous steppe of Festuca spp.
dominates the landscape.
The scheme presented here, based on Schmithüsen (1956) and Luebert and Pliscoff
(2006) corresponds to the potential vegetation, but all the formations and most of the
vegetation belts that compose the Chilean vegetation are to a high degree affected by
the long history of human occupation, from localized mining impacts in the north to
extended forest substitutions in the south. The core of the deciduous forest at around
38◦ S to 41◦ S has been transformed into agriculture, and in the Central Derpression
only remnants of sclerophyllous forests remain in this mainly cultural landscape
(Chap. 6).
References
Amigo J, Ramírez C (1998) A bioclimatic classification of Chile: woodland communities in the
temperate zone. Plant Ecol 136:9–26
Andersen BG, Denton GH, Heusser CJ, Lowell TV, Moreno PI, Hauser A, Heusser LE, Schluchter
C, Marchant D (1995) Climate, vegetation, and glacier fluctuations in Chile between 40◦ 30′
and 42◦ 30′ s latitude: a short review of preliminary results. Quat Int 28:199–201
Anderson JM, Anderson HM, Archangelsky S, Bamford M, Chandra S, Dettmann M, Hill R,
McLoughlin S, Rösler O (1999) Patterns of Gondwana plant colonisation and diversification.
J Afr Earth Sci 28:145–167
Artabe AE, Morel EM, Spalletti LA (2003) Characterization of the phytogeographic Triassic
provinces from extratropical Gondwana. Ameghiniana 40:387–405
Artabe AE, Ganuza DG, Spalletti LA, Zúñiga A, Morel EM (2005) Revisión de la paleoflora
del cerro La Brea (Jurásico Temprano), provincia de Mendoza, Argentina. Ameghiniana 42:
429–442
Ashworth AC, Markraf V, Villagrán C (1991) Late Quaternary climatic history of the Chilean
Channels based on fossil pollen and beetle analysis, with an analysis of the modern vegetation
and pollen rain. J Quat Sci 6:279–291
Astini RA, Benedetto JL, Vaccari NE (1995) The early Paleozoic evolution of the Argentine
Precordillera as a Laurentian rifted, drifted, and collided terrane: a geodynamic model. Geol
Soc Am Bull 107:253–273
Barichivich J, Sauchyn DJ, Lara A (2009) Climate signals in high elevation tree-rings from
the semiarid Andes of north-central Chile: responses to regional and large-scale variability.
Palaeogeogr Palaeoclimatol Palaeoecol 281:320–333
Barreda V, Archangelsky S (2006) The southernmost record of tropical pollen grains in the mid-
Cretaceous of Patagonia, Argentina. Cretaceous Res 27:778–787
Barreda VD, Anzótegui LA, Prieto AR, Aceñolaza PG, Bianchi MM, Borromei AM, Brea M et al.
(2007) Diversificación y cambios de las angiospermas durante el Neógeno en Argentina. In:
Archangelsky S, Sánchez T, Tonni EP (eds) Ameghiniana 50◦ aniversario, Publicación Especial
11:173–191
Barrientos SE (2007) Earthquakes in Chile. In: Moreno T, Gibbons W, (eds) The geology of Chile.
Geological Society, London, pp 263–287
Beerling DJ (2002) Low atmospheric CO2 levels during the Permo-Carboniferous glaciation
inferred from fossil lycopsids. Proc Natl Acad Sci 99:12567–12571
Benton MJ, Twitchett RJ (2003) How to kill (almost) all life: the end-Permian extinction event.
Trends Ecol Evol 18:358–365
Berry EW (1922a) Contributions to the paleobotany of Peru, Bolivia and Chile. The John Hopkins
Press, Baltimore.
Berry EW (1922b) The flora of Concepción-Arauco coal measures of Chile. Stud Geol (John
Hopkins Univ) 4:73–142
Betancourt JL, Saavedra B (2002) Rodent middens, a new method for Quaternary research in arid
zones of South America. Rev Chil Hist Nat 75:527–546
Brea M, Zucol AF (2006) Leños fósiles de Boraginaceae de la Formación Peñas Coloradas
(Paleoceno superior), Puerto Visser, Chubut, Argentina. Ameghiniana 43:139–146
Cecioni G, (ed) (1968) El Terciario de Chile: zona central. Editorial Andrés Bello, Santiago.
Charrier R (1988) Condiciones Paleoclimáticas para el Carbonífero Superior y Pérmico Inferior en
la mitad austral de América del Sur. Bol Mus Nac Hist Nat (Chile) 41:105–116
Charrier R, Pinto L, Rodríguez M (2007) Tectono-stratigraphic evolution of the Andean orogen in
Chile. In: Moreno T, Gibbons W, (eds) The geology of Chile. The Geological Society, London,
pp 21–116
Chumakov NM, Zharkov MA (2003) Climate of the Late Permian and Early Triassic: general
inferences. Stratigr Geol Correl 11:361–375
Convey P, Gibson JAE, Hillenbrand C-D, Hodgson DA, Pugh PJA, Smellie JL, Stevens MI (2008)
Antarctic terrestrial life – challenging the history of the frozen continent? Biol Rev 83:103–117
Cúneo RC (1989) Phytogeography and paleoecology of Late Paleozoic Floras from Southern South
America and their relationship with other floral realms. Abstracts, 28th International Geological
Congress, p 351
Cúneo RN, Johnson K, Wilf P, Gandolfo MA, Iglesias A (2007) A preliminary report on the diver-
sity of Latest Cretaceous floras from Northern Patagonia, Argentina. Abstract GSA Denver
Annual Meeting
Denton GH, Heusser CJ, Lowell TV, Moreno PI, Andersen BG, Heusser LE, Schlüchter C,
Marchant DR (1999) Interhemispheric linkage of paleoclimate during the last glaciation. Geogr
Ann 81A:107–153
Di Castri F (1968) Esquisse écologique du Chili. In: Delamanre-Debouteville C, Rapoport E, (eds)
Biologie del’ Amérique Australe, 4. Editions du Centre National de la Recherce Scientfique,
Paris, pp 6–52
Di Castri F (1981) Mediterranean-type shrublands of the world. In: Di Castri F, Goodall DW,
Specht RL, (eds) Mediterranean-type Shrublands. Ecosystems of the World vol 11. Elsevier,
Amsterdam, pp 1–52
Di Castri F, Hajek E (1976) Bioclimatología de Chile. Vicerrectoría Académica, Universidad
Católica de Chile, Santiago.
DiMichele WA, Pfefferkorn HW, Gastaldo RA (2001) Response of late carboniferous and early
permian plant communities to climate change. Annu Rev Earth Planet Sci 29:461–487
Dowsett HJ, Barron JA, Poore RZ, Thompson RS, Cronin TM, Ishman SE, Willard DA (1999)
Middle pliocene paleoenvironmental reconstruction: PRISM2. US Geological Survey Open
File Report, 99–535. http://pubs.usgs. gov/of/1999/of99-535/
Dunai TJ, González López GA, Juez-Larré J (2005) Oligocene–Miocene age of aridity in
the Atacama Desert revealed by exposure dating of erosion-sensitive landforms. Geology
33:321–324
References 41
Dusén P (1907) Über die Tertiäre Flora der Magellansländer. In: Nordenskjold O (ed)
Wissenschaftlichen Ergebnisse der Schwedischen Expedition nach der Magellansländern,
1895–1897, vol I, pp 84–108
Emck P, Moreira-Muñoz A, Richter M (2006) El clima y sus efectos en la vegetación. In: Moraes
M, Øllgaard B, Kvist LP, Borchsenius F, Balslev H, (eds) Botánica Económica de los Andes
Centrales, Universidad Mayor de San Andrés, La Paz, pp 11–36
Encinas A, Maksaev V, Pinto L, Le Roux JP, Munizaga F, Zentilli M (2006) Pliocene lahar deposits
in the Coastal Cordillera of central Chile: implications for uplift, avalanche deposits, and
porphyry copper systems in the Main Andean Cordillera. J S Am Earth Sci 20:369–381
Endlicher W, Santana A (1988) El clima del sur de la Patagonia y sus aspectos ecológicos: un siglo
de mediciones climatológicas en Punta Arenas. An Inst De La Patagonia (Chile) 18:57–86
Engelhardt H (1891) Über Tertiärpflanzen von Chile. Abh Senckenb Naturforsch Ges 16:629–692
Fickert T, Friend D, Grüninger F, Molnia B, Richter M (2007) Did debris covered glaciers serve
as Pleistocene refugia for plants? A new hypothesis derived from observations of recent plant
growth on Glacier surfaces. Arct Antarct Alp Res 39:245–257
Fosdick JC (2007) Late Miocene exhumation of the Magallanes Basin and sub-Andean fold belt,
southern Chile: new constraints from apatite U-Th/He thermochronology. Abstracts 2007 GSA
Denver Annual Meeting
Frakes LA, Francis JE, Syktus JI (1992) Climate modes of the Phanerozoic. Cambridge University
Press, Cambridge.
Fuenzalida H (1938) Las capas de Los Molles. Bol Mus Nac Hist Nat 16:67–92
Fuenzalida H (1966) Historia vegetacional de Chile. Estudios geográficos . Facultad de Filosofía y
Educación, Universidad de Chile, Santiago, pp 21–39
Gajardo R (1994) La vegetación natural de Chile: clasificación y distribución geográfica. Ed.
Universitaria, Santiago.
Gallagher SJ, Wagstaff BE, Baird JG, Wallace MW, Li CL (2008) Southern high latitude climate
variability in the Late Cretaceous greenhouse world. Glob Planet Change 60:351–364
Gandolfo MA, Zamaloa MC (2003) Evolution of Upper Cretaceous and Tertiary Angiosperm
floras of Patagonia. 12◦ Simposio Argentino de Paleobotánica y Palinología (Buenos Aires),
Resúmenes, pp 40–41
Gandolfo MA, Gonzalez CC, Zamaloa MC, Cúneo NR, Wilf P, Johnson K (2007) Eucalyptus
(Myrtaceae) macrofossils from the early Eocene of Patagonia, Argentina. 5◦ Southern
Connection Conference, Adelaide, Australia, Abstracts p 32
Garreaud RD, Aceituno P (2007) Atmospheric circulation over South America: mean features and
variability. In: Veblen T, Young K, Orme A, (eds) The physical geography of South America.
Oxford University Press, Oxford, pp 45–66
Garreaud RD, Muñoz R (2004) The diurnal cycle of circulation and cloudiness over the subtropical
southeast Pacific: a modeling study. J Clim 17:1699–1710
Garreaud RD, Vuille M, Compagnucci R, Marengo J (2009) Present-day South American climate.
Paleogeogr Paleoclimatol Paleoecol doi: 10.1016/j.palaeo.200710.032
Garzione CN, Hoke GD, Libarkin JC, Withers S, MacFadden B, Eiler J, Ghosh P, Mulch A (2008)
Rise of the Andes. Science 320:1304–1307
Gayó E, Hinojosa LF, Villagrán C (2005) On the persistence of tropical paleofloras in central Chile
during the Early Eocene. Rev Palaeobot Palynol 137:41–50
Ghosh P, Garzione NC, Eiler JM (2006) Rapid uplift of the Altiplano revealed through 13C-18O
bonds in paleosol carbonates. Science 311:511–515
González CC, Gandolfo MA, Zamaloa MC, Cúneo NR, Wilf P, Johnson KR (2007) Revision of
the Proteaceae macrofossil record from Patagonia, Argentina. Bot Rev 73:235–266
González-Ferrán O (1994) Volcanes de Chile. Instituto Geográfico Militar (IGM), Santiago.
Gradstein FM, Ogg JG, Smith AG et al (2004) A geologic time scale 2004. Cambridge University
Press, Cambridge.
Gregory-Wodzicki KM (2000) Uplift history of the central and northern Andes: a review. Geol Soc
Am Bull 112:1091–1105
Hartley AJ (2003) Andean uplift and climate change. J Geol Soc 160:7–10
Hartley AJ, Chong G (2002) A late Pliocene age for the Atacama Desert: implications for the
desertification of western South America. Geology 30:43–46
Haywood AM, Francis JE, Sellwood BW (2002) Global middle Pliocene biome reconstruction: a
data/model synthesis. Geochem Geophys Geosy 3:1–18
Herbst R, Troncoso A (1996) La tafoflora de Juan de Morales del Jurásico Medio (Formación
Chacarilla), Región de Tarapacá, Chile. Rev Geol Chile 23:3–15
Hesse M, Zetter R (2005) Ultrastructure and diversity of recent and fossil zona-aperturate pollen
grains. Plant Syst Evol 255:145–176
Heusser CJ (1997) Deglacial setting of the southern andes following the last glacial maximum: a
short review. An Inst Patagonia Serie Cienc Nat 25:89–103
Heusser CJ (2003) Ice age southern andes: a chronicle of paleoecological events. Developments in
Quaternary Science 3. Elsevier, Amsterdam.
Hinojosa LF (2005) Cambios climáticos y vegetacionales inferidos a partir de paleofloras
cenozoicas del sur de Sudamérica. Rev Geol Chile 32:95–115
Hinojosa LF, Armesto JJ, Villagrán C (2006) Are Chilean coastal forests pre-Pleistocene relicts?
Evidence from foliar physiognomy, paleoclimate, and paleobiogeography. J Biogeogr 33:
331–341
Hoke GD, Garzione CN (2008) Paleosurfaces, paleoelevation, and the mechanisms for the late
Miocene topographic development of the Altiplano plateau. Earth Planet Sci Lett 271:192–201
Holderegger R, Thiel-Egenter C (2009) A discussion of different types of glacial refugia used in
mountain biogeography and phylogeography. J Biogeogr 36:476–480
Iglesias A, Wilf P, Johnson KR, Zamuner AB, Matheos SD, Cúneo RN (2007) Rediscovery of
Paleocene Macrofloras in Central Patagonia, Argentina. GSA Denver Annual Meeting (28–31
October 2007)
IGM (2005) Atlas de Chile. Instituto Geográfico Militar, Santiago.
Illies H (1959) Die Entstehungsgeschichte eines Maares in Süd-Chile: ein aktuo-geologischer
Beitrag zum Problem des Maar-Vulkanismus. Int J Earth Sci 48:232–247
Jasper A, Guerra-Sommer M, Cazzulo-Klepzig M, Iannuzzi R (2003) Biostratigraphic and pale-
oclimatic significance of Botrychiopsis fronds in the Gondwana realm. In: Wong TE (ed)
Proceedings of the XVth International Congress on Carboniferous and Permian Stratigraphy,
Utrecht, 10–16 August 2003
Klicka J, Zink RM (1997) The importance of recent ice ages in speciation: a failed paradigm.
Science 277:1666–1669
Knapp S, Mallet J (2003) Refuting refugia? Science 300:71–72
Köppen W (1930) Die Klimagebiete nach Köppens Klassifikation. In: Knoch K (ed) Klimakunde
von Südamerika vol. 2. Handbuch der Klimatologie (ed. by W Köppen, Geiger R), Berlin, part
G, pp 242–329
Looy CV, Twitchett RJ, Dilcher DL, Van Konijnenburg-Van Cittert JHA, Visscher H (2001) Life
in the end-Permian dead zone. Proc Natl Acad Sci 14:7879–7883
Lowell TV, Heusser CJ, Andersen BG, Moreno PI, Hauser A, Denton GH, Heusser LE, Schluchter
C, Marchant D (1995) Interhemispheric correlation of Late Pleistocene Glacial events. Science
269:1541–1549
Luebert F, Pliscoff P (2006) Sinopsis bioclimática y vegetacional de Chile. Editorial Universitaria,
Santiago.
Maldonado A, Betancourt JL, Latorre C, Villagrán C (2005) Pollen analyses from a 50 000-yr
rodent midden series in the southern Atacama Desert (25 degrees 30’s). J Quat Sci 20:493–507
Massaferro JI, Moreno PI, Denton GH, Vandergoes M, Dieffenbacher-Krall A (2009) Chironomid
and pollen evidence for climate fluctuations during the Last Glacial termination in NW
Patagonia. Quat Sci Rev 28:517–525
McAllister Rees P, Ziegler AM, Valdes PJ (2000) Jurassic phytogeography and climates: new data
and model comparisons. In: Huber BT, Macleod KG, Wing SL, (eds) Warm climates in earth
history. Cambridge University Press, Cambridge, pp 297–318
References 43
McAllister Rees P, Ziegler AM, Gibbs MT, Kutzbach JE, Behling PJ, Rowley DB (2002) Permian
phytogeographic patterns and climate data/model comparisons. J Geol 110:1–31
McElwain JC, Punyasena SW (2007) Mass extinction events and the plant fossil record. Trends
Ecol Evol 22:548–557
McLoughlin S (2001) The breakup history of gondwana and its impact on pre-Cenozoic floristic
provincialism. Aust J Bot 49:271–300
McLoughlin S, Carpenter RJ, Jordan GJ, Hill RS (2008) Seed ferns survived the end-Cretaceous
mass extinction in tasmania. Am J Bot 95:465–471
Milleron M, Gallo LA, Marchelli P (2008) The effect of volcanism and postglacial migration and
seed dispersal. A case study in Southern South America. Tree Genet Genomes 4:435–444
Moore DM (1983) Flora of Tierra del Fuego. Missouri Botanical Garden, St. Louis, MO.
Moreno FP (1882) Patagonia, resto de un antiguo continente hoy sumergido. An Soc Cient Argent
14:97–131
Moreno PI (2004) Millennial-scale climate variability in northwest Patagonia over the last
15 000 yr. J Quat Sci 19:35–47
Moreno PI, Jacobson GL, Lowell TV, Denton GH (2001) Interhemispheric climate links revealed
by a late-glacial cooling episode in southern Chile. Nature 409:804–808
Nalpas T, Dabard M-P, Ruffet G, Vernon A, Mpodozis C, Loi A, Hérail G (2008) Sedimentation
and preservation of the Miocene Atacama Gravels in the Pedernales–Chañaral area, Northern
Chile: climatic or tectonic control? Tectonophysics 459:161–173
Nishida M (1984) The anatomy and affinities of the petrified plants from the tertiary of Chile, III.
Petrified woods from Mocha Island, Central Chile. In: Nishida M, (ed). Contributions to the
Botany in the Andes, I. Academia Scientific Books, Tokyo, pp 98–110
Oberdorfer E (1960) Pflanzensoziologische Studien in Chile. Ein Vergleich mit Europa. Flora Et
Vegetatio Mundi 2:1–208
Ochsenius CC (1891) Über Tertiärpflanzen von Chile von H. Engelhardt (Nachtrag). Abh Senckenb
Naturforsch Ges 16:1–4
Okuda M, Nishida H, Uemura K, Yabe A (2006) Paleocene/Eocene pollen assemblages from the
Ligorio Márquez Formation, Central Patagonia, XI Region, Chile. In: Nishida H, (ed) Post-
Cretaceous floristic changes in southern patagonia, chile. Chuo University, Tokyo, pp 37–43
Orme AR (2007) The tectonic framework of South America; tectonism, climate, and landscape
change. In: Veblen TT, Young KR, Orme AR, (eds) The physical geography of South America.
Oxford University Press, New York, NY, pp 3–44
Palazzesi L, Barreda V (2007) Major vegetation trends in the Tertiary of Patagonia (Argentina): a
quatitative paleoclimatic approach based on palynologial evidence. Flora 202:328–337
Pankhurst RJ, Hervé F (2007) Introduction and overview. In: Moreno T, Gibbons W, (eds) The
geology of Chile. The Geological Society, London, pp 1–4
Pankhurst RJ, Rapela CW, Fanning CM, Márquez M (2006) Gondwanide continental collision and
the origin of Patagonia. Earth Sci Rev 76:235–257
Pastorino MJ, Gallo LA (2002) Quaternary evolutionary history of Austrocedrus chilensis, a
cypress native to the Andean–Patagonian forest. J Biogeogr 29:1167–1178
Pisano E (1954) Fitogeografía. La vegetación de las distintas zonas geográficas chilenas. Rev
Geogr Terra Australis 11:95–107
Prámparo MB, Quattrocchio M, Gandolfo MA, Zamaloa MC, Romero E (2007) Historia evolutiva
de las angiospermas (Cretácico-Paleógeno) en Argentina a través de los registros paleoflorísti-
cos. In: Archangelsky S, Sánchez T, Tonni EP (eds) Ameghiniana 50◦ aniversario, Publicación
Especial 11:157–172
Premoli AC, Kitzberger T, Veblen TT (2000) Isozyme variation and recent biogeographical history
of the long-lived conifer Fitzroya cupressoides. J Biogeogr 27:251–260
Quattrocchio ME, Martínez MA, Volkheimer W (2007) Las floras jurásicas de la Argentina In:
Archangelsky S, Sánchez T, Tonni EP (eds) Ameghiniana 50◦ aniversario, Publicación Especial
11: 87–100
Quintanilla V (1974) La carta bioclimática de Chile Central. Rev Geogr Valparaíso 5:33–58
Quintanilla V (1983) Biogeografía. Colección Geografía de Chile. Instituto Geográfico Militar

(IGM), Santiago.
Ramos VA (2008) Patagonia: a paleozoic continent adrift? J S Am Earth Sci 26:235–251
Ramos VA (2009) Anatomy and global context of the Andes: main geological features and the
Andean orogenic cycle. Gsa Memoirs 204:31–65
Ramos VA, Aleman A (2000) Tectonic evolution of the Andes. In: Cordani UJ et al (eds) Tectonic
evolution of South America, International Geological Congress, 31st, Rio de Janeiro (2000),
pp 635–685
Ranero CR, von Huene R, Weinrebe W, Reichert C (2006) Tectonic processes along the Chile
convergent margin. In: Oncken O, Chong G, et al (eds) The Andes: Active Subduction Orogeny.
Springer, Berlin, pp 91–121
Rapalini AE (2005) The accretionary history of southern South America from the latest Proterozoic
to the Late Palaeozoic: some palaeomagnetic constraints. In: Vaughan APM, Leat PT,
Pankhurst RJ, (eds) Terrane processes at the margins of Gondwana, 246. Geological Society,
London, Special Publications, pp 305–328
Raymond A, Gensel PG, Stein WE (2006) Phytogeography of Late Silurian macrofloras. Rev
Paleobot Palynol 142:165–192
Reiche C (1907) Grundzüge der Pflanzenverbreitung in Chile. In Series: Engler A, Drude O, (eds)
Die Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, vol. VIII. Verlag
von V. Engelmann, Leipzig.
Rivas-Martínez S, Rivas-Sáenz S (1996–2009) World Bioclimatic Classification System.
Phytosociological Research Center, Spain. http://www.globalbioclimatics.org
Romero EJ (1986) Paleogene phytogeography and climatology of South Maerica. Ann Mol Bot
73:449–461
Romero EJ (1993) South American paleofloras. In: Goldblatt P, (ed) Biological relationships
between Africa and South America. Yale University Press, New Haven, CT/London, pp 62–85
San Martín J, Donoso C (1996) Estructura florística e impacto antrópico en el bosque maulino de
Chile. In: Armesto JJ, Villagrán C, Arroyo MK, (eds) Ecología de los bosques nativos chilenos.
Editorial Universitaria, Santiago, pp 153–167
Schmithüsen J (1956) Die räumliche Ordnung der chilenischen Vegetation. Bonn Geogr Abh
17:1–86
Scotese CR, Boucot AJ, McKerrow WS (1999) Gondwanan palaeogeography and palaeoclimatol-
ogy. J Afr Earth Sci 28:99–114
Simpson BB (1971) Pleistocene changes in the fauna and flora of South America. Science
173:771–780
Stern C, Moreno H, López-Escobar L, Clavero J, Lara LE, Naranjo JA, Parada MA, Skewes MA
(2007) Chilean volcanoes. In: Moreno T, Gibson W, (eds) The geology of Chile. The Geological
Society, London, pp 147–178
Storey BC (1995) The role of mantle plumes in continental breakup: case histories from
Gondwanaland. Nature 377:301–308
Subercaseaux B (1940) Chile o una loca geografía. Ercilla S, (ed). English translation from 1943:
Chile, a geographic extravaganza. Macmillan Co., New York, NY
Thiel M, Macaya EC, Acuna E, Arntz WE, Bastias H et al. (2007) The humboldt current system
of northern and central Chile. Oceanogr Mar Biol 45:195–344
Torres T, Rallo M (1981) Anatomía de troncos fósiles del Cretácico Superior de Pichasca en
el norte de Chile. Anais II Congreso Latinoaméricano Paleontología. Porto Alegre, Brasil 2:
385–398
Tremetsberger K, Stuessy TF, Guo Y-P, Baeza CM, Weiss H, Samuel RM (2003) Amplified
Fragment Length Polymorphism (AFLP) variation within and among populations of
Hypochaeris acaulis (Asteraceae) of andean southern South America. Taxon 52:237–245
Trewartha G (1961) The Earth’s problem climates. University of Wisconsin Press, Madison, WI.
Troncoso A, Encinas A (2006) La tafoflora de cerro Centinela (Chile, VI Región): vegetación
y clima de Chile central a fines del Mioceno-comienzos del Plioceno. Ameghiniana 43:
171–180
References 45
Troncoso A, Romero EJ (1998) Evolución de las comunidades florísticas en el extremo sur de

Sudamérica durante el Cenofítico. In: Fortunato R, Bacigalupo N (eds) Proceedings of the
Congreso Latinoamericano de Botánica, Monographs in Systematic Botany from the Missouri
Botanical Garden 6: 149–172
Troncoso A, San Martín A (1999) Presencia del género Escallonia (Magnoliopsida,
Escalloniaceae) en el Terciario de Chile Central. Bol Mus Nac Hist Nat (Chile) 48:29–36
Troncoso A, Suarez M, De La Cruz R, Palma-Heldt S (2002) Paleoflora de la Formación
Ligorio Márquez (XI Región, Chile) en su localidad tipo: sistemática, edad e implicancias
paleoclimáticas. Rev Geol Chile 29:113–135
Truswell EM (1990) Cretaceous and tertiary vegetation of Antarctica: a palynological perspec-
tive. In: Taylor TN, Taylor EL, (eds) Antarctic paleobiology: its role in the reconstruction of
Gondwana. Springer-Verlag, New York, pp 71–88
Veblen TT, Young KR, Orme AR, (eds) (2007) The physical geography of South America. Oxford
University Press, Oxford.
Vega JC, Archangelsky S (1997) The first Gondwana Carboniferous compound cupules and
associated seeds. A preliminary note. Rev Palaeobot Palynol 99:55–59
Villagrán C, Le-Quesne C, Aravena JC, Jiménez H, Hinojosa F (1998) El rol de los cambios de
clima del Cuaternario en la distribución actual de la vegetación de Chile central-sur. Bamberg
Geogr Schr 15:227–242
Villagrán C, León A, Roig FA (2004) Paleodistribución del alerce y ciprés de las Guaitecas durante
períodos interestadiales de la Glaciación Llanquihue: provincias de Llanquihue y Chiloé,
Región de Los Lagos, Chile. Rev Geol Chile 31: 133–151
Vuille M, Baumgartner MF (1998) Monitoring the regional and temporal variability of winter
snowfall in the arid andes using digital NOAA/AVHRR data. Geocarto Int 13:59–67
Weischet W (1970) Chile: seine länderkundliche individualität und struktur. Wissenschaftliche
Buchgesellschaft, Darmstast.
Wilf P, Johnson KR, Cúneo NR, Smith ME, Singer BS, Gandolfo MA (2005) Eocene plant
diversity at Laguna del Hunco and Río Pichileufú, Patagonia, Argentina. Am Nat 165:634–650
Wilf P, Gandolfo MA, Johnson KR, Cúneo R (2007) Biogeographic significance of the Laguna
del Hunco flora, Early Eocene of Patagonia, Argentina. Abstract GSA Denver Annual Meeting
(28–31 October 2007)
Willis KJ, McElwain JC (2002) The evolution of plants. Oxford University Press, Oxford.
Willis KJ, Niklas KJ (2004) The role of Quaternary environmental change in plant macroevolution:
the exception or the rule? Philos Trans R Soc Lond B 359:159–172
Windhausen A (1931) Geología Argentina. Geología Histórica y Regional del Territorio Argentino,
Buenos Aires.
Wnuk C (1996) The development of floristic provinciality during the Middle and Late Paleozoic.
Rev Palaeobot Palynol 90:5–40
Zachos J, Pagani M, Sloan L, Thomas E, Billups K (2001) Trends, rhythms, and aberrations in
global climate 65 Ma to present. Science 292:686–693
Chapter 2
Getting Geobotanical Knowledge
Abstract The history of the discovery of the Chilean plant world can be traced
back to the Magallanes voyage around the globe. Since then dozens of naturalists
and botanists contributed to build the corpus of the geobotanical knowledge of the
country. According to most recent updates, the Chilean flora is composed of 56
orders, 171 families, 837 genera, and about 4,295 species. This includes 4 endemic
families, 84 endemic genera, and 1,936 species endemic to the Chilean continen-
tal and oceanic territory. Richest families in genera and species numbers are the
Asteraceae, the Poaceae, the Fabaceae, and the Solanaceae, while the species-richest
genera are Senecio, Adesmia, Viola, and Carex.
A detailed revision of early collectors, botanists, and naturalists that contributed

to the description of the Chilean flora and geobotany was done by Reiche (1907).
Newer references to geobotanical works appear in Marticorena’s taxonomic and
botanical blibliography (Marticorena 1992, 1996). For the current Flora de Chile,
he also provided an updated revision (Marticorena 1995). Some highlights and
principal actors in the discovery of the Chilean plant world are briefly exposed here.
2.1 Romancing the South: The Discovery of a Virgin World

The discovery of the Strait of Magellan by Hernando de Magallanes and his crew
(21 October 1520) set the starting point for the European exploration of South
America. Magallanes gave the name “Cabo de Las Vírgenes” to the eastern Cape
that allowed them to get into the Strait. During the first days of exploration, the
observations are about a native wood to make fire of which the smoke smelled well.
They refer undoubtedly to the wood of “canelo”, Drimys winteri. The sailors early
noted the properties of canelo’s bark and herbs like Cardamine and Apium against
scurvy (Pigafetta 1536). The first notes from sailors refer of course the medicinal,
nutritional and wooden properties of the new plants found.
During the long conquest of Chile, many chroniclers referred to the cultivated and
natural plants and their properties, such as Gerónimo de Bibar, who accompanied
Pedro de Valdivia’s enterprise writing his Crónica y Relación copiosa y verdadera

DOI 10.1007/978-90-481-8748-5_2, ⃝
48 2 Getting Geobotanical Knowledge
a b
c d
Fig. 2.1 a Juan Ignacio Molina; b illustration in Molina’s work: mapuche kids and native plants:
“araucaria” (Araucaria araucana), “palma” (Jubaea chilensis), and “culén” (Otholobium glandu-
losum); c cover of Flora Peruviana et Chilensis from Ruiz and Pavón; d Lapageria rosea in Ruiz
and Pavón’s work
del Reyno de Chile between 1539 and 1558 (Muñoz-Schick 1975). During the
seventeenth century, important advances in the description of the territory and their
plant world were made by Jesuits such as Alonso de Ovalle in his Histórica Relación
del Reyno de Chile, or Diego de Rosales in his Historia General del Reino de
Chile, Flandes Indiano. This tradition continued with the Jesuit Juan Ignacio Molina
(1737–1829) who is considered the first Chilean naturalist (Fig. 2.1a). After the
expulsion of the Jesuit Company from America (1768), Molina published in Italy his
Compendio della storia geografica, naturale, e civile del Regno del Chile (Molina
1776). This and later works (Molina 1782, 1810) were for a long time the main
sources of knowledge on natural sciences of Chile.
2.1 Romancing the South: The Discovery of a Virgin World 49
a b c
Fig. 2.2 a Eduard Poeppig; b Ombrophytum subterraneum; c Viola cotyledon from Poeppig and
Endlicher (1835–1845)
Towards the end of the eighteenth century, and with the purpose of extending the
collections of the Real Botanical Garden of Madrid, King Carlos III of Spain sent
several naturalist expeditions to different countries; Peru and Chile were assigned to
Hipólito Ruiz and José Pavón. They made collections at some localities between
Talcahuano and Santiago between 1782 and 1783. Their work Flora Peruviana
et Chilensis was published in 3 volumes and contains more than 300 illustrations,
including the copihue, Lapageria rosea, that was later declared as the national flower
(Fig. 2.1c, d).
In 1827 the German naturalist Eduard Poeppig arrived in Chile (Fig. 2.2). He
explored the country for two years and it was the first time that a foreign naturalist
stayed in Chile for a long period. He also visited other countries of South America,
and his work of 3 volumes and 300 illustrations included for the first time colour
paintings (Poeppig and Endlicher 1835–1845). During the night of 9th to 10th of
January 1827 Poeppig for the first time crossed the equator aboard the ship Gulnare:
“The day we left the northern hemisphere was one of the most important in my long
life of travel. . . I had the feeling of entering into a new world and also into a new
life. . .” (after Morawetz and Röser 1998).
The English naturalist Joseph Dalton Hooker (1817–1911) accompanied the
expedition to the South Pole of John Clark Ross (1839–1843). He was the first
who collected intensively in the southern lands: Kerguelen Islands, Tasmania, New
Zealand, Falkland (Malvinas) Islands, and Tierra del Fuego. He rapidly noticed the
close floristic relationships between these territories. His Botany of the Antarctic
Voyage was published in three volumes between 1844 and 1859 (Fig. 9.1)
Charles Darwin (1809–1882) travelled around the globe aboard the Beagle from
1832 to 1836. He stayed in Chile from 1834 to 1835 and collected almost 1,500
plant specimens from Argentina, Falkland (Malvinas) Islands and Chile that were
later studied by W.J. Hooker, G.A.W. Arnott and J.D. Hooker. It is widely believed
that his visit to the southern territories and his impressions about the southern biota
and geology were fundamental for his later writings on the theory of evolution
(Box 2.1).
Box 2.1 Darwin’s Impressions from Central Chile

“We spent the day on the summit, and I never enjoyed one more thoroughly.
Chile, bounded by the Andes and the Pacific, was seen as in a map. The
pleasure from the scenery, in itself beautiful, was heightened by the many
reflections which arose from the mere view of the grand range, with its lesser
parallel ones, and of the broad valley of Quillota directly intersecting the lat-
ter. Who can avoid admiring the wonderful force which has upheaved these
mountains, and even more so the countless ages which it must have required,
to have broken through, removed, and leveled whole masses of them?” Darwin
(1839), from his diary while climbing the Cerro La Campana in Central Chile
(33◦ S) (Fig. 2.3).
Fig. 2.3 Memorial tablet dedicated to Ch. Darwin, approaching the top of Cerro La
Campana
After its Independence from the Spanish dominion in 1810, the young Republic
of Chile had to better know its natural resources. The French professor of natural
sciences, Claudio Gay (1800–1873), already living in Chile, was contracted by the
government to do the first intensive scientific exploration of the territory (Fig. 2.4).
He intensively travelled between Copiapó and Chiloé (1830–1842) (Muñoz Pizarro
1944). Afterwards he published his masterpiece Historia Física y Política de Chile,
which consists of 28 volumes (8 on botany) and 2 illustration atlases (one cultural
a b
Fig. 2.4 a Claudio Gay; b illustration of Nassauvia lagascae in Gay’s Atlas; c species altitudinal
limits from Pissis (1875) (www.memoriachilena.cl)
and one physical) (Gay 1845–1854, reprint 2010). This was a work without prece-
dents in America to that date. The mission that the Government of Chile had trusted
on him also included the formation of a Cabinet of Natural Sciences. That became
the basis of the Museo Nacional de Historia Natural (Box 2.2). Before Gay, the
flora of Chile was almost unknown, consisting of only ca 300 species; Gay’s com-
pilation numbered 3,767 species. More than 600 were described by himself and his
collaborators.
The Historia Física. . . of Gay, including both illustrated Atlases, is one of the
most important milestones in the development of Chilean natural history. Naturalists
coming after him could count on this truly basal study, and it allowed them to rapidly
develop many research fields: botany and zoology by R.A. Philippi, geology by I.
Domeyko, geography by A. Pissis. It is worth mentioning that these advances were
not only basic for the scientific development in Chile, but also were fundamen-
tal for the consolidation of the Republic (Saldivia Maldonado 2003). Moreover,
according to Stuardo and Feliú (1973, p 318), Gay’s writings on Chilean plant
geography were the first American advances in the field after the milestone of
Alexander von Humboldt and Aimé Bonpland Essai sur la Géographie des Plantes
of 1805 (Chap. 4). The first representation of altitudinal and latitudinal limits of sev-
eral plants is the profile accompanying the Geographical Atlas from Pissis (1875)
(Fig. 2.4c).
a b e
c d
Fig. 2.5 a Rodulfo Amando Philippi; b Federico Philippi; c Federico Johow; d Carlos Reiche;
e cover of Gründzuge der Pflanzenverbreitung in Chile
Rudolf Amandus Philippi (1808–1904) (Fig. 2.5a) arrived in Chile in 1851 with
a recommendation from A. von Humboldt to the epoch’s government. He rapidly
began exploring the territory making discoveries and descriptions in botany, zool-
ogy, geology, paleontology, and ethnology. He gave a large impulse to the Museo
Nacional de Historia Natural, getting the palace that harbours till today the main
natural history collections of Chile (Box 2.2).
RA Philippi got the collaboration of his son Federico Philippi (1838–1910)
(Fig. 2.5b), exploring vast regions in Chile and the Norte Grande, territories just
annexed to Chile (Taylor and Muñoz-Schick 1994). RA Philippi’s (1860) descrip-
tion of the botanical exploration of the Atacama is a vivid document of the practical
issues to solve at those early stages of botanical development. RA Philippi described
more than 3,750 plant species! and is recognized as the main naturalist in the history
of Chilean science (Castro et al. 2006). The Museum conserves 90% of Philippi’s
type exemplars.
Federico Johow (1859–1933) (Fig. 2.5c) arrived in Chile in 1889 to teach natural
sciences at the Instituto Pedagógico, Instituto Nacional and at the Universidad de
Chile. He is the author of the milestone Estudios sobre la flora de las Islas de Juan
Fernández, still an obligate reference for the islands (Johow 1896) (Chap. 5).
Carlos Reiche (1860–1929) (Fig. 2.5d), a German professor of natural sciences,
was appointed at the botanical section of the Museo Nacional de Historia Natural
in 1897 by Federico Philippi, when the last took the direction of the Museo. Reiche
published the second Flora de Chile (after Gay) in 6 volumes (Reiche 1894–1911).
a d
Fig. 2.6 a Francisco Fuentes; b Ivan Murray Johnston; c Gualterio Looser; d Adiantum gertrudis
illustrated by E. Sierra for Looser’s publications
He also explored intensively the Chilean territory and published the first plant geog-
raphy of Chile: Gründzuge der Pflanzenverbreitung in Chile in 1907 (Fig. 2.5e).
Francisco Fuentes (1879–1934) (Fig. 2.6a) was in charge of the National
Herbarium in 1911, as Carlos Reiche left the country accepting an appointment
in Mexico. Fuentes made the first study on plants of Easter Island (Fuentes 1913)
(Chap. 5) and on Monocots that had not been treated in Reiche’s Flora.
Ivan Murray Johnston (1898–1960) (Fig. 2.6b), prominent US botanist, studied
the coastal desert flora and vegetation. His papers The coastal flora of the depart-
ments of Chañaral and Taltal, and The flora of the Nitrate Coast (1929a, b), are
considered classical studies of the northern Chilean coast, where vegetation is rather
sparse but the flora is one of the most interesting at all.
Gualterio Looser (1898–1982) (Fig. 2.6c), showed a wide interest in the natural
sciences, having been appointed as chief of Anthropology at the National Museum.
After a journey to Juan Fernández in 1925 he demonstrated more interest in botany,
publishing till 1971 about 140 works mostly dedicated to ferns. He became the
national authority in this field. He also made big efforts in the translation into
a c
Fig. 2.7 a Carl Skottsberg; b Carlos Muñoz Pizarro; c cover of Sinopsis de la Flora Chilena
Spanish of main texts from C. Reiche and C. Skottsberg, including Reiche’s Plant
Geography, translated into Spanish as Geografía Botánica de Chile (1934–1937,
reprint 2010).
Carl Skottsberg (1880–1963) (Fig. 2.7a), Swedish botanical eminence, president
of the 7th International Botanical Congress in 1950, came first with the Swedish
expedition to the South Pole, between 1901 and 1903 and afterwards several times
explored Patagonia and the Pacific islands. He published about 128 articles and sev-
eral books about Chilean botany and plant geography. His Vegetations-verhältnisse
längs der Cordillera de los Andes südlich von 41 Grad. . . (1916) and his Natural
History of Juan Fernandez and Easter Island (1920–1956) are till today the most
complete works that treat these territories (see Chaps. 4 and 5).
Carlos Muñoz Pizarro (1913–1976) (Fig. 2.7b) was a student of Francisco
Fuentes. After a Guggenheim scholarship in the USA, under the direction of IM
Johnston, he became in charge of the Botanical Section at the Museo Nacional de
Historia Natural in 1942. He initiated a program of organization of the collections:
around 30,700 specimens were revised, catalogued and organized. In 1961, with a
grant from the Rockefeller Foundation, Muñoz Pizarro explored, together with his
wife Ruth Schick, the principal European herbaria, carrying out the photographic
registry of the type collections of Chilean plants. The acquired knowledge allowed
him to publish several important books: Sinopsis de la Flora Chilena (1959, 2nd
ed. 1966) (Fig. 2.7c), the principal synthetical work on the Chilean flora still in use;
Flores Silvestres de Chile (1966); Chile: Plantas en extinción (1973) (see Sect. 6.1).
To this brief revision of the most prominent “Chilean” botanists we should
add Edmundo Pisano (1919–1997), one of the most prolific plant researchers of
Patagonia and Magallanes, and Otto Zöllner (1909–2007), indefatigable collector,
professor and connoisseur of the plants of Central Chile. Finally, we should not for-
get to mention the national and international collaborators to the current Flora de
Chile, a continuing work in progress directed by the Universidad de Concepción
(Marticorena and Rodríguez 1995 onwards). This University contains one of the
most important Herbarium in Chile (CONC) with 175.000 specimens, initiated by
A. Pfister and continued by M. Ricardi, C. Marticorena and R. Rodríguez. Chilean
botanists recently collaborated in the great effort that represented the compilation
of the Catálogo de las Plantas Vasculares del Cono Sur, an up to date checklist
encompassing Argentina, Chile, Paraguay, Uruguay, and southern Brazil (Zuloaga
et al. 2008) (Sect. 2.2).
Box 2.2 The National Herbarium (SGO)

The National Herbarium (SGO in the international code) is strongly related
to the discovery and description of the Chilean plants (Muñoz-Schick 1991).
Is it located in Santiago in the Museo Nacional de Historia Natural (Fig. 2.8).
The National Herbarium has increased its collections under the curatorship
of Mélica Muñoz-Schick, daughter of C. Muñoz Pizarro. Today it contains
almost 81,000 specimens which are a sample of the entire vascular plant
diversity of the country. Bryophytes, Fungi and Algae are also well repre-
sented in the Herbarium. The museum holds also a palaeobotanical collection.
Museums and Herbaria have funding problems everywhere, but their role in
systematic biology and nowadays in conservation is increasingly being recog-
nized (e.g. Graham et al. 2004; Wake et al. 2009). Together with a constant
increase of collections, the daily work in the Herbarium comprises orga-
nization, identification, digitizing and georeferencing of the specimens. In
accordance with the global trend, digitalization of the specimens is actually
on its way. This considers the scanning of more than 4,000 type specimens,
including the first collections of Carlo G. Bertero from 1828. These are the
first steps towards a Virtual Herbarium (Muñoz-Schick and Moreira-Muñoz
2008), a challenge taken up by many countries worldwide (Wang and Qin
2008).
a b
c d
Fig. 2.8 a View of the old palace containing the Museo Nacional de Historia Natural since
R.A. Philippi’s directorship; b early collector C.G. Bertero; c type Specimen from Laurelia
sempervirens, collected by C.G. Bertero in the year 1828
2.2 Classification and Phylogeny of the Chilean Vascular Flora

While preparing the new Flora de Chile, Reiche’s account on the Chilean Flora
comprised 141 families and 716 genera (Reiche 1907). The previous account by F
Philippi numbered a total of 863 genera (Philippi 1881) (Table 2.1). Muñoz Pizarro
(1959) in his Sinopsis de la Flora Chilena gave a number of 182 families and 960
native genera, and Marticorena’s (1990) checklist comprised 175 families and 827
genera (Table 2.1).
For his Flora de Chile, Reiche (1894–1911) followed Engler and Prantl’s clas-
sification system. For his Sinopsis, Muñoz Pizarro (1959) followed Christensen
2.2 Classification and Phylogeny of the Chilean Vascular Flora 57
Table 2.1 Historical numbers for the Chilean native vascular plants
Orders Families Genera
F. Philippi (1881) – – 863

Reiche (1907) – 141 716
Muñoz Pizarro (1966) 91 182 960a
Marticorena (1990) – 175 827
a Including many naturalized taxa.
(1934) for the Pteridophyta, Engler and Diels (1936) for the Gymnosperms, and
Hutchinson (1926, 1934) for the Angiosperms. Marticorena and Quezada (1985),
and Marticorena (1990) presented their checklists in alphabetic order following
the criteria of several authors, like Cronquist (1981). The permanent revision of
taxa under the current cladistic paradigm leads to drastic changes in the traditional
classification of ferns and fern allies (Smith et al. 2006; Pryer and Schuettpelz
2009) and angiosperms (APGII 2003, APGIII 2009; Stevens 2001 onwards).
Gymnosperms are more constant in their classification, although relationships
between main groups are not resolved (Renner 2009).
According to these modern classification schemes, a comprehensive and updated
checklist of the Chilean flora is presented here. The basis for this account has
been the Southern Cone Checklist (Zuloaga et al. 2008), which has been revised
and updated according to Hassler and Swale (2002), Pryer et al. (2004) and Smith
et al. (2006) for the ferns and fern allies; and according to APGIII (2009) for the
angiosperms. Families’ classification differs slightly from APGIII (2009), consider-
ing the opinion of several experts. The resulting checklist is presented as Appendix,
including accepted genera and their global distribution, latitudinal range in Chile,
and the number of native and endemic species (not including infraspecific taxa).
According to this revision, the extant Chilean vascular native flora consists of 56
orders, 171 families, 837 genera, and about 4,295 species. Among the latter, ca.
1,936 species (45%) are endemic to the Chilean territory (continental or oceanic).
Numbers for traditional groups are presented in Table 2.2.
Table 2.2 Summary of Chilean vascular plants (updated January 2010)
Orders Families Genera Native species Endemic species
Ferns and fern allies 11 22 52 146 38

Gymnosperms 2 4 9 16 3
Monocots 6 28 176 906 330
Dicots 37 117 600 3,227 1,565
Total 56 171 837 4,295 1,936
This account includes the offshore oceanic flora. Pacific genera not represented
on the continent, many of them endemic to the islands, plus two endemic families
(in bold), are listed in Table 2.3 (see Chap. 5).
Table 2.3 Genera not found in continental Chile
Genera (family)
Juan Fernández Islands ferns and fern allies Dicksonia (Dicksoniaceae)a

Thyrsopteris (Thyrsopteridaceae)a
Arthropteris (Tectarinaceae)a
Gymnosperms –
Monocots Juania (Arecaceae)
Machaerina (Cyperaceae)
Megalachne (Poaceae)
Podophorus (Poaceae)
Dicots Centaurodendron (Asteraceae)
Dendroseris (Asteraceae)
Robinsonia (Asteraceae)
Yunquea (Asteraceae)
Selkirkia (Boraginaceae)
Haloragis (Haloragaceae)
Cuminia (Lamiaceae)
Lactoris (Lactoridaceae)a
Zanthoxylum = Fagara (Rutaceae)
Coprosma (Rubiaceae)
x Margyracaena (Rosaceae)
Santalum (Santalaceae) (1 extinct species)
Boehmeria (Urticaceae)
Desventuradas Islands Ferns and fern allies –
Gymnosperms –
Monocots –
Dicots Lycapsus (Asteraceae)
Thamnoseris (Asteraceae)
Nesocaryum (Boraginaceae)
Sanctambrosia (Caryophyllaceae)
Isla de Pascua Ferns and fern allies Davallia (Davalliaceae)a
Microlepia (Dennstaedtiaceae)
Diplazium (Woodsiaceae)
Dryopteris (Dryopteridaceae)
Microsorum (Polypodiaceae)
Psilotum (Psilotaceae)a
Vittaria (Pteridaceae)
Doodia (Blechnaceae)
Gymnosperms –
Monocots Kyllinga (Cyperaceae)
Pycreus (Cyperaceae)
Axonopus (Poaceae)
Stipa (Poaceae)
Dicots Triumfetta (Malvaceae)
Ipomoea (Convolvulaceae)
a Families absent from continental Chile.
2.2.1 Rich Families and Genera

As in many floras (e.g. Flora of Ecuador, see Table 2.8), most families and genera
are represented by a few species: 41 families (24%) are represented by only one
species and 80 families (47%) contain 14 or fewer species. Fifty families (29%)
have 15 or more species (Fig. 2.9a). The reduced number of species is most marked
at the genus level: most genera (381) are represented by only one species (46%),
and almost the same percentage of genera contains 14 or fewer species (389 genera,
46%). Only 67 genera (8%) have 15 or more species (Fig. 2.9b).
The Asteraceae, usually the largest family in floras of arid or semi-arid regions
(Goldblatt and Manning 2000), is also the most species- and generic-rich family
in the Chilean flora (Table 2.4) (Chap. 8) (Fig. 8.1). The second largest family is
a b
Fig. 2.9 a Number of families (white bars) versus the number of species; b number of genera
(black bars) versus the number of species
Table 2.4 Ranking of the 20
largest families in the Chilean Family Native species Endemic species
flora by size
Asteraceae 838 374
Poaceae 380 73
Fabaceae 280 152
Solanaceae 169 87
Cyperaceae 150 32
Brassicaceae 145 66
Cactaceae 112 88
Boraginaceae 103 64
Apiaceae 99 42
Malvaceae 75 37
Violaceae 73 50
Caryophyllaceae 72 32
Calceolariaceae 63 45
Verbenaceae 60 18
Orchidaceae 56 32
Portulacaceae 55 22
Amaranthaceae 54 23
Oxalidaceae 53 26
Veronicaceae 53 13
Loasaceae 46 22
a b
c d
e f
Fig. 2.10 Genus-rich families, representative species of the Poaceae: a Cortaderia atacamensis,
Chusmiza; b Chusquea cumingii, Lircay; c Melica argentata, Cerro La Campana; d Deschampsia
antarctica; e Festuca ortophylla and Mélica Muñoz-Schick, Colchane; f Hordeum comosum,
Juncal (photo/illustration credits: a–c, e A. Moreira-Muñoz; d plate 133 in J.D. Hooker (1847))
d e
Fig. 2.11 Genus-rich families, representative species of the Fabaceae: a Balsamocarpon brevi-
folium, Cuesta Pajonales; b Anarthrophyllum cumingii, Laguna Negra; c Astragalus cruckshanksii,
Portillo; d Adesmia argentea, southern Atacama; e Geoffroea decorticans, Cuesta Pajonales (photo
credits: a, b, d, e A. Moreira-Muñoz; c Sergio Moreira)
the Poaceae, as can be expected from the global richness of the family (Fig. 2.10).
Fabaceae, Solanaceae, Cyperaceae, and Brassicaceae follow in size (Figs. 2.11,
2.12).
The species-richest genera are Senecio, Adesmia, Viola, Carex, Calceolaria,
Haplopappus (Table 2.5). Some of these genera are cosmopolitan species-rich
a b
c d
Fig. 2.12 Genus-rich families, representative species of the Solanaceae: a Schizanthus hookeri,
Portillo; b Lycium chilense, Concón; c Salpiglossis sinuata, Rio Clarillo; d Nolana coelestis;
Cuesta Buenos Aires (photo credits: a, c, d A. Moreira-Muñoz; b S. Elórtegui Francioli)
genera (e.g. Senecio, Carex, Viola), while others are strictly neotropical genera (e.g
Haploppapus, Calceolaria). Adesmia is a notable case, being the species-richest
genus restricted to the southern Andes of Chile/Argentina (with little representation
in Perú and southern Amazonas). Senecio and Adesmia are also ahead in endemic
species numbers.
Table 2.5 Ranking of the 20 largest genera in the Chilean flora
Genus Family Species Endemic species
Senecio Asteraceae 224 103

Adesmia Fabaceae 130 85
Viola Violaceae 72 50
Carex Cyperaceae 67 16
Calceolaria Calceolariaceae 60 42
Haplopappus Asteraceae 54 42
Oxalis Oxalidaceae 53 26
Astragalus Fabaceae 49 23
Nolana Solanaceae 44 40
Solanum Solanaceae 44 19
Valeriana Valerianaceae 44 21
Poa Poaceae 42 7
Leucheria Asteraceae 42 16
Baccharis Asteraceae 40 11
Cryptantha Boraginaceae 40 31
Chaetanthera Asteraceae 38 16
Dioscorea Dioscoreaceae 36 34
Alstroemeria Alstroemeriaceae 33 26
Chloraea Orchidaceae 32 18
Loasa Loasaceae 31 18
2.2.2 Endemic Families

The Chilean flora comprises four endemic families: Gomortegaceae, Francoaceae
(Fig. 2.13), Thyrsopteridaceae and Lactoridaceae, the latter two endemic to Juan
Fernández (Table 2.6) (Chap. 5). The flora also contains several subendemic
families, restricted to Chile and adjacent territories in Argentina and/or Peru:
Aextoxicaceae, Quillajaceae, Misodendraceae, Philesiaceae, and Malesherbiaceae
(Table 2.7); and one subendemic family restricted to Chile and Australia:
Berberidopsidaceae. The Chilean flora harbours 84 endemic genera, which are
discussed in Sect. 3.1.
The high level of endemism at the family and genus level is remarkably, even
resting the Francoaceae (Table 2.6). As a comparison, Argentina harbours one
endemic family (Halophytaceae) and 45 endemic genera (Zuloaga et al. 1999). The
flora of Perú, composed of more than 17,000 species, and around 2,400 native gen-
era, has 51 endemic genera but none endemic family (Brako and Zarucchi 1993, not
considering ferns). The flora of Ecuador (2,110 native genera, and around 15,300
species), has 23 endemic genera and also none endemic family (Jørgensen and
León-Yáñez 1999) (Table 2.8). Other distant territories worth of interest due to
their biogeographical relationships are New Zealand and the Cape Floristic Region,
which contain respectively 48 and 160 endemic genera (Table 2.8). Compared to all
Table 2.6 Endemic families of the Chilean floraa
Family Genera N◦ species
Gomortegaceae Gomortega 1
Francoaceaea Francoa, Tetilla 2
Lactoridaceae Lactoris 1 (Juan Fernández)
Thyrsopteridaceae Thyrsopteris 1 (Juan Fernández)
a Francoaceae has been alternatively included in Melianthaceae
(APGIII 2009).
Table 2.7 Subendemic families of the Chilean flora
Family Genera N◦ species N◦ species Chile Distribution
Aextoxicaceae Aextoxicon 1 1 Chile/Argentina

Philesiaceae Lapageria 1 1 Chile/Argentina
Philesia 1 1
Misodendraceae Misodendrum 8 8 Chile/Argentina
Quillajaceae Quillaja 2 1 Chile/Arg/Brazil/Uruguay
Malesherbiaceaea Malesherbia 24 18 Chile/Perú/Argentina
Berberidopsidaceae Berberidopsis 2 1 Chile/E Australia
a Alternatively considered in the Passifloraceae by APG III (2009).
Table 2.8 Generic and familiar endemism in several vascular floras

N◦ N◦
native Endemic native Endemic Native Endemic
Country families families genera genera species species Sources
Chile 171 4 837 84 4,293 1,933 Updated here from

Zuloaga et al.
(2008)
Argentina ~ 250 1 900 45 ~ 9,000 1,760 Zuloaga et al. (1999)
and (2008))
Perú ~ 200 0 2,400 51 ~ 15,000 5,354 Brako and Zarucchi
(1993) (not
considering ferns)
Ecuador 254 0 2,110 23 15,306 4,173 Jørgensen and
León-Yáñez
(1999)
New 90 0 446 49 1,936 1,591 Wilton and
Zealand Breitwieser (2000)
and De Lange
et al. (2006)
Cape 178 5 994 160 9,000 6,210 Goldblatt and
Floristic Manning (2000)
Region
Note: This comparison must be taken as illustrative but not definitive, since each of these floras
follows its own classification system.
Fig. 2.13 Francoa appendiculata, illustrated in Cavanilles (1801)
these territories, the Chilean flora has a remarkable floristic identity, not equalling
the overwhelming Cape flora, but surpassing most species-rich South American
countries as regards endemism at genus and family levels.
2.2.3 Phylogenetic Groups

The systematic arrangement of the plant world has seen a dramatic improvement
during the last decade, under the current paradigm of cladistics. The monophyly
of the land plants is nowadays well supported by morphological and molecular
characters (Magallón and Hilu 2009) (Fig. 2.15). Living land plants have been
placed in four major clades: antocerophytes (hornworts, ca. 100 species), bryophytes
(mosses, ca. 13,000 species), marchantiophytes (liverworts; 5,000–8,000 species),
and tracheophytes (vascular plants: ca. 285,000 species) (Magallón and Hilu 2009)
(Fig. 2.15).
Vascular plant evolution has been viewed traditionally as “a successive series
of incremental increases in complexity, from simple bryophytic ancestors through
vascularized spore producers, more complex seed plants, and ultimately to
angiosperms” (Pryer et al. 2004, p 1582) (Box 2.3).
Box 2.3 Origin of the Vascular Plants

The time of the origin of the early vascular plants can be the Mid-Ordovician
(ca. 475 mya) (Wellman et al. 2003) or the Late Silurian (ca. 425 mya) (Gensel
2008). A deep phylogenetic dichotomy occurred in the early-mid Devonian
(ca. 400 mya), separating the lycophytes from the euphyllophytes (Gensel and
Berry 2001; Pryer et al. 2004). Monilophytes have been dated back to the Late
Devonian (ca. 370 mya). The earliest known occurrence of fossil leptosporan-
giate ferns is in the Early Carboniferous (Galtier and Phillips 1996); by the end
of the Carboniferous six families were present. In subsequent major radiations
in the Permian, Triassic, and Jurassic, several families with extant represen-
tatives replaced these Carboniferous families (e.g. Osmundaceae, Schizaceae,
Matoniaceae, and Dipteridaceae) (Rothwell 1987). The more derived poly-
pod ferns dramatically diversified in the Cretaceous, accompanying the great
angiosperm diversification (Schneider et al. 2004) (see Sect. 1.2).
Molecular dating results are giving much older dates for the land plant
crown group, from the Ordovician, ca. 490 mya (Sanderson 2003), to the
Precambrian (Heckman et al. 2001). This result contrasts sharply with
palaeobotanical estimates (Kenrick and Crane 1997; Wellman et al. 2003).
The extant seed plants have been shown to be a monophyletic group, but exact
relationships among these lineages and the pattern and chronology of diver-
gence remain unclear, despite of the recent accumulation of molecular data
sets and techniques to analyse them (e.g. Burleigh and Mathews 2004) (see
Box 3.3).
Gnetales and modern conifer families appeared in the Triassic to
Jurassic, and angiosperms on the boundary of the Jurassic/Cretaceous peri-
ods. From the Permian through the late Jurassic many seed plant lin-
eages went extinct, including lyginopterids, medullosans, Callistophytaceae,
Glossopterids, Cordaitales, and Voltziales (Stewart and Rothwell 1993), and
their relationships with extant groups remain poorly resolved. During the
Cretaceous and Cenozoic, the diversity of all surviving seed plant lines, except
angiosperms, decreased (Knoll 1984; Willis and McElwain 2002) (Fig. 2.14).
During the past 20 years the phylogenetic relationships are beginning to be

clearer and the predominant paraphyletic former scheme is changing to a more com-
800
Angiosperms
600 Gymnosperms
Pteridophytes
Number of species
Early vascular plants
400
200
0
Silurian Devonian Carboniferous Permian Triassic Jurassic Cretaceous Cenozoic
400 300 200 100 0
Millions of years
Fig. 2.14 Species richness of different vascular plant groups from the Silurian into the Cenozoic
(from Willis and McElwain (2002), by permission of Oxford University Press)
prehensive system of evolutionary relationships (Pryer et al. 2004). Monilophytes

and lycophytes are all spore bearing and “seed-free”, and therefore, were tra-
ditionally lumped under terms like “pteridophytes” or “ferns and fern allies”,
that united paraphyletic assemblages of plants (Pryer et al. 2004). Now there
is the tendency to differentiate the earliest diverging lineage Lycopsida or lyco-
phytes from the real ferns or monilophytes (Fig. 2.15). The vascular plants
(Tracheophyta) are classified in four main groups: lycophytes, ferns, Gymnosperms
and Angiosperms. Lycophytes all possess lycophylls (leaves with an intercalary
meristem) and comprise three main clades: homosporous Lycopodiales (club-
mosses), heterosporous Isoetales (quillworts) and Selaginellales (spikemosses)
(Fig. 2.15). Extant lycophytes are mostly diminute plants, but fossil members that
dominated the Carboniferous landscape (like Lepidodendron) were huge arbores-
cent forms. These are today the major component of coal deposits (Stewart and
Rothwell 1993) (see Sect. 1.2).
2.2.3.1 Lycophytes
Lycophytes constitute the earliest diverging lineage under the vascular plants,
and are consequently the most “primitive” plants in the Chilean extant vascu-
lar flora. They comprise two orders (Lycopodiales and Isoetales), two families
(Lycopodiaceae and Isoetaceae), three genera (Isoetes, Huperzia, Lycopodium), and
Monophyletic Paraphyletic
s
rn
fe
se
e
Spermatophytes
..
yt
at
s
d.
lie
ph
gi
an
an
al
o
s
rid
rn
or
rn
e
fe
p
Fe
Pt
Eu
...
Euphyllophytes
Angiosperms
Gymnosperms
Monilophytes
Leptosporangiate ferns x x
Tracheophytes
Horsetails x x
(ferns)
Marattioid ferns x x x
Ophioglossoid ferns x x x
Wisk ferns x x
Embryophytes
Lycophytes
Quillworts (Isoetes) x x
Spikemosses (Selaginella) x x
Clubmosses (Huperzia) x x
Anthocerophytes (hornworts)
Bryophytes (mosses)
Marchantiophytes (liverworts)
Fig. 2.15 Cladogram showing relationships among major lineages of vascular plants as currently
and formerly understood (i.e. monophyletic v/s paraphyletic) (adapted from Pryer et al. 2004;
Magallón and Hilu 2009)
11 species. Huperzia appears in Chile only at subantarctic latitudes (Fig. 1.20),

while Isoetes and Lycopodium are to be found further north to 37ºS.
2.2.3.2 Monilophytes
Living euphyllophytes belong to two major clades (Fig. 2.15): seed plants (sper-
matophytes) and monilophytes (Kenrick and Crane 1997; Pryer et al. 2004).
Monilophytes, ferns s. str., include most former groups recognized as “fern and
fern allies”. Ferns are represented in the Chilean flora by 9 orders, 20 families, 49
genera, and 135 species. Among the last, the Chilean flora comprises 38 endemic
fern species. Basal classes among ferns s. str. are Psilotopsida, Equisetopsida, and
Marattiopsida (Fig. 2.16), represented in Chile by three families: Ophioglossaceae
(genera Botrychium and Ophioglossum), Psilotaceae (only Psilotum from Isla de
Pascua), and Equisetaceae (Equisetum). The Order Marattiales does not show mod-
ern representatives in Chile, but is represented in the fossil record by the genus
Asterotheca for the Upper Triassic of the La Ternera formation (Copiapó province,
Ophioglossaceae (Ophioglossoids) Ophioglossales

Psilotaceae (Wisk ferns) Psilotales
Ferns Equisetaceae (Horsetails) Equisetales
Marattiaceae (Marattioids) Marattiales
Osmundaceae (Osmundaceous ferns) Osmundales
Hymenophyllaceae (Filmy ferns) Hymenophyllales
Leptosporangiates Gleichenioids Gleicheniales
Schizaeoids Schizaeales
Marsileaceae
Salviniales
Salviniaceae
Tree ferns-1 (incl. Thyrsopteridaceae)
Cyatheales
Core leptosporangiates Tree ferns-2 (incl. Dicksoniaceae)
Lindsaeoids
Dennstaedtiaceae
Polypods Pteridaceae
Polypodiales
Eupolypods I
Eupolypods II
Fig. 2.16 Relationships between main fern groups (adapted from Smith et al. 2006; Pryer and
Schuettpelz 2009, by permission of Oxford University Press)
Herbst et al. 1998), and the genera Asterotheca and Rienitsia for the Upper Triassic
of the Biobío province (Leppe et al. 2006).
The most diverse of the monilophytes are the leptosporangiate ferns, a group
of more than 9,000 extant species (Pryer and Schuettpelz 2009), represented
in Chile by 125 species. The species-richest family is the Pteridaceae, with 10
genera and 26 species. The species-richest genus in Chile is Hymenophyllum
(filmy ferns), comprising 19 species. The closely related monospecific genera
Hymenoglossum and Serpyllopsis are considered by Ebihara et al. (2006) as nested
within Hymenophyllum. Important to mention is the high fern species-richness in
the oceanic islands, especially Juan Fernández and Rapa Nui. The Juan Fernández
archipelago harbours 55 native fern species, of which 50% are endemic to the islands
(Marticorena et al. 1998). A very remarkably species is Thyrsopteris elegans, the
only representative of the tree fern family Thyrsopteridaceae (Smith et al. 2006)
(Table 2.6, Fig. 5.8). Rapa Nui lacks such a huge species richness, but contains
several tropical genera that are not represented in the rest of the Chilean territory,
like Davallia, Diplazium, and Vittaria (Table 2.3). In the continent, ferns occupy
most habitats from the coast to the Andes, and from the arid North (e.g. Equisetum
giganteum) to the humid South (e.g. Lophosoria quiadripinnata). The generic fern
richness concentrates in Chile between 35 and 45◦ S.
2.2.3.3 Spermatophytes
The seed plants (spermatophytes) (Fig. 2.15) are the most diverse vascular plant
group in the world. Spermatophytes comprise Gymnosperms and angiosperms.
Extant seed plants likely number between 250,000 and 300,000 species (Scotland
and Wortley 2003). In Chile the seed plants are represented by 45 orders, 149
families, 785 genera and 4,144 species (Appendix A).
2.2.3.4 Gymnosperms
The seed-bearing plants or gymnosperms are characterized, contrary to the
angiosperms, by the presence of ovules on the edge of an open sporophyll.
Gymnosperms are classified into four main groups: conifers, cycads, ginkgos, and
Gnetales. Gymnosperms as a whole are recognized as being monophyletic, but the
relationships between the groups remain unresolved (Renner 2009) (Fig. 2.17).
The Chilean Flora is lacking extant cycads or Ginkgoales, but they were
characteristic of the Triassic and Jurassic landscapes (see Sect. 1.2).
Fossil cycads have been reported from the Triassic of the Santa Juana Formation
at the lower Biobío (Leppe and Moisan 2003; Nielsen 2005), from the Panguipulli
Formation (Zavattieri et al. 2003), and the La Ternera formation (also Triassic)
(Herbst and Troncoso 2000).
Ginkgoales are also found in the Chilean Triassic floras, like the ones from
Copiapó and Los Molles (Troncoso and Herbst 1999), from La Ligua (Torres and
Philippe 2002), and from the Lake District (39º30′ S) (Herbst et al. 2005). This last
palaeoflora is composed of ca. 25 vascular plant species, including representatives
from the Apocalamitaceae, Asterothecaceae, Gleicheniaceae, Corystospermaceae
(Dicroidium), Peltaspermaceae, Cycadophyta, and Coniferales (Podocarpaceae and
Voltziaceae).
Chilean Gnetales are represented by the family Ephedraceae, with the genus
Ephedra comprising 7 species, 2 of them endemic.
Conifers harbour a rich fossil record that goes back to the Late Carboniferous,
ca 300 mya (Hilton et al. 2003). Traditionally the conifers have been consid-
ered as the sister clade to Ginkgophyta and Cycadophyta, but this relationship
Podocarpaceae
Araucariaceae
Conifers (including Gnetales)
Cupressaceae
Cephalotaxaceae
Taxaceae
Welwitschiaceae
Gnetaceae
Ephedraceae
Pinaceae
Ginkgoaceae
Cycads
Cycadaceae
Zamiaceae
Fig. 2.17 Putative relations between main groups of gymnosperms (from Renner 2009, by
permission of Oxford University Press)
20
30
40
50
Araucaria Austrocedrus Fitzroya Lepidothamnus
20
30
40
P. salignus
50 P. nubigenus
Pilgerodendron Podocarpus Prumnopitys Saxegothaea
Fig. 2.18 Distribution maps of Chilean conifers (SGO collections)
has been constantly challenged (Henry 2005); new proposals place the Pinaceae
as sister to the Gnetales and both as sister to the other conifers (Renner 2009)
(Fig. 2.17). Conifers are a main issue in some of the most characteristic Chilean
landscapes, especially in the temperate rainforests. They are represented in the
a b
c d
Fig. 2.19 Chilean conifers: a Araucaria araucana, Nahuelbuta; b Podocarpus salignus, Antuco;
c Austrocedrus chilensis, Antuco; d Fitzroya cupressoides, continental Chiloé (photo credits: a–c
A. Moreira-Muñoz; d María Castro)
extant flora by 3 families and 8 genera: Araucaria (Araucariaceae); Lepidothamnus,

Saxegothaea, Podocarpus, Prumnopitys (Podocarpaceae); Austrocedrus, Fitzroya,
Pilgerodendron (Cupressaceae) (Fig. 2.18) (Fig. 2.19 ).
The Araucariaceae are a family that is abundant worldwide in deposits from
the Mesozoic (Stockey 1982) and became restricted to the southern hemisphere
(Gondwana) from the end of the Mesozoic (fine distribution map in Golte 1993).
The family expanded and diversified in both hemispheres in the Jurassic and Early
Cretaceous and remained a significant component of Gondwanan vegetation until
the end of the Cenozoic (see Sect. 1.2). The development of angiosperms in the
Middle Cretaceous probably assisted in the demise of some araucarian components
but there was also evolution of new genera. Recorded diversity in the early Cenozoic
of Australia is as high as it was in the Early Cretaceous (Kershaw and Wagstaff
2001). Araucaria araucana distribution in Chile shows a disjunction between the
coastal cordillera (Nahuelbuta) and the Andes, that is also expressed in the specie’s
genetic diversity (Ruiz et al. 2005).
2.2.3.5 Angiosperms
Flowering plants or Angiosperms, defined as plants with ovules enclosed in a
carpel, are the most intensively studied group due to their predominance in mod-
ern terrestrial ecosystems (Palmer et al. 2004; Magallón 2009). They comprise
around 223,000–260,000 living species classified in 61 orders and ca. 450 families
(Scotland and Wortley 2003; APGIII 2009). The fossil record of the angiosperms
extends back at least to the early Cretaceous, conservatively 130 mya (Crane et al.
2004). Molecular dating has but pushed this age to the Early – Middle Jurassic
(ca 170 mya) (Wikström et al. 2001; Magallón 2009) (Box 2.4). Relationships
between Angiosperm orders are presented in Fig. 2.20, including numbers of
Chilean representatives.
2.2.3.6 Magnoliids
The most basal groups with representatives in Chile correspond to the magno-
liid clade, including Piperales, Laurales, Canellales and Magnoliales (this lat-
ter with no presence in Chile). Chileans representatives of these basal orders
are the Lactoridaceae, Piperaceae, Aristolochiaceae, Winteraceae, Lauraceae,
Monimiaceae, Gomortegaceae, and Atherospermataceae.
Monimiaceae and Lauraceae were widespread by the Upper Cretaceous. The old-
est fossil of the Lauraceae has been dated at 109 mya (Crane et al. 1994) and of
the Monimiaceae at 83 mya (Poole and Gottwald 2001). Chilean Peumus suppos-
edly diverged from a Monimia/Palmeria line some 76 mya, perhaps by disruption
of a once continuous range that stretched from Chile across Antarctica and the
Kerguelen plateau to Madagascar (Renner 2004). Laurelia sempervirens seems to
have been present already since some 80 mya, while Gomortega keule is considered
a relict species that has been in existence without modification for 100 my (Renner
Amborellales
Nymphaeales
Austrobaileyales
angiosperms
Piperales 3/3
Canellales 1/1
magnoliids
Magnoliales
Laurales 4/7
Poales 7/109
commelinids
Arecales 1/2
Asparagales 8/41
monocots Liliales 4/7
Dioscoreales 1/2
Alismatales 7/15
Ceratophyllales 1/1
Ranunculales 4/12
Proteales 1/4
Gunnerales 1/1
Cucurbitales 2/2
eudicots Fagales 2/2
Rosales 3/21
Fabales 3/23
Celastrales 1/2 fabids
Oxalidales 3/6
Malpighiales 10/20
Zygophyllales 2/7
Malvales 2/15
rosids Brassicales 4/28
core eudicots Sapindales 3/9

Myrtales 3/17 malvids
Geraniales 3/8
Vitales 1/1
Saxifragales 4/8
Berberidopsidales 2/2
Santalales 4/13
Caryophyllales 12/75
Cornales 2/8
Ericales 4/18
Gentianales 3/17
asterids Lamiales 14/58 lamiids
Solanales 2/28
Boraginaceae 1/13
Aquifoliales 1/1
Escalloniales 1/3
Asterales 5/137
Dipsacales 1/2
campanulids
Apiales 3/26
Bruniales 1/1
Fig. 2.20 Cladogram showing relationships between angiosperm orders (adapted from Stevens
2001 onwards, APG III 2009). Numbers represent native families/genera present in Chile; only
basal orders not present in Chile are shown (without numbers)
2004) (see Sect. 3.3). The Lactoridaceae is a monospecific family endemic to Juan
Fernández, constituting a real biogeographic enigma (Chap. 5).
Monocots
Regarding the Monocots, considerable diversification took place during the
Early Cretaceous, with most families already present at the Cretaceous-Cenozoic
boundary (Janssen and Bremer 2004). Basal within the Monocots are the
Acorales and Alismatales, the former being absent from Chile and the latter
present by means of 6 families, 15 genera and 29 (mostly aquatic) species.
Dioscoreales are represented by the sole family Dioscoreaceae, with 2 gen-
era and 39 species. Liliales diverged from their sister-group in the Early
Cretaceous, 117 mya (Bremer 2000). Among them are the Chilean national
species Lapageria rosea, and species-rich Alstroemeriaceae (Table 2.5, Muñoz-
Schick and Moreira-Muñoz 2003). Monogeneric Luzuriagaceae has been recently
treated as nested within Alstroemeriaceae (APGIII 2009). Asparagales encom-
pass also a big amount of Chilean Monocots (8 families, 41 genera, 195 species,
and within them, 133 endemics). According to APGIII (2009), Asparagaceae
may include Laxmanniaceae (Trichopetalum), and Xanthorrhoeaceae shall include
Hemerocallidaceae (Pasithea). There is also the proposal to include Alliaceae into
the Amaryllidaceae, but this is not followed by Anderson and Janssen (2009).
Janssen and Bremer (2004) found an unexpected old age of 111 mya for the
Orchidaceae: “Traditionally, this family has been looked at as a very specialized
and hence, probably, a young group. . . Orchid diversity is not necessarily due
to a rapid and recent radiation” (Janssen and Bremer 2004).The Tecophilaeaceae
are represented in Chile by three endemic genera: Conanthera, Tecophilaea, and
Zephyra (see Sect. 4.3). The rest of the Monocots have been grouped in a clade
called the Commelinids that comprises four orders: under them is the Poales, which
comprises in Chile 7 families, 109 genera, 596 species, of which 128 endemic
species, mostly from the Poaceae and Cyperaceae. Poales has been dated at ca
115 mya (Mid-Cretaceous) (Bremer 2000). Bromeliaceae also appear to date back
to the Cretaceous (Linder and Rudall 2005), but the uncertainties in dating the
Bromeliaceae are considerable (Bremer 2002).
Eudicots
The probable sister of the eudicots is Ceratophyllum (Ceratophyllaceae), as appar-
ent from APGIII (2009), but alternative models propose a closer relationship with
Magnoliids (Goremykin et al. 2009). Ranunculales is the first diverging clade in
the eudicots, followed by Proteales. These basal eudicots comprise in Chile 5 fam-
ilies, 16 genera, 71 species, and among the Ranunculales, 18 endemic species.
Most genera in the Proteaceae show a Gondwanic relationship with Australasia
(Chap. 3).
The Gunnerales is the first diverging lineage in the core eudicots (APGIII
2009; Forest and Chase 2009). Pantropical Gunneraceae are represented in south-
ern Chile and Juan Fernández by 11 species, 6 of them endemic. Relationships
within the rest of the core eudicots are getting clearer: Saxifragales appears as
sister to the newly recognized Vitales + Rosids. Saxifragales are represented by
the Saxifragaceae and to a less extent by the Haloragaceae and Grossulariaceae.
Berberidopsidales appear as sister to Santalales + Caryophyllales + Asterids. The
recently recognized order Berberidopsidales (APG III 2009) is composed of two
monotypic families from Central Chile/Australia (Berberidopsidaceae) and Central
Chile/Argentina (Aextoxicaceae) (Table 2.7). Santalales are represented, together

with the Santalaceae, by several parasitic Loranthaceae mainly from North-central
Chile, and the Misodendraceae, distributed from 33ºS to the South as parasites
of extant Nothofagus species (Chap. 9). Caryophyllales is one of the orders with
the highest presence in Chile, encompassing 12 families that include the species-
rich Cactaceae in North-central Chile (Chap. 7), and the related Portulacaceae, as
well as several rich families like the Caryophyllaceae and Polygonaceae (72 and
32 species respectively). The Molluginacea and Nyctaginaceae are more sparsely
represented.
Rosids
Rosids have been recently organized in two main clades: fabids and malvids,
incorporating as sister clade the newly recognized Vitales (APG III 2009). These
last are represented in Chile only by Cissus striata (Vitaceae), a liana from cen-
tral Chilean sclerophyllous forests. Fabids are represented in Chile by 8 orders:
Zygophyllales (comprising the Zygophyllaceae and Krameriaceae), Celastrales
(including the Lepuropetalaceae under the Celastraceae), Oxalidales (including the
Cunoniaceae, Elaeocarpaceae, Oxalidaceae), Malpighiales (including 10 Chileans
families), Cucurbitales, Fabales, Fagales, and Rosales. Under the fabids, the best
representation in Chile rests on the Fabales, that comprises the largest Fabaceae,
the third largest family of angiosperms, comprising more than 19,000 species
(Lewis et al. 2005). Chilean species-richest genera from the Fabaceae are Adesmia
and Astragalus (Table 2.5). The Fabales also includes the Polygalaceae and
Quillajaceae, this latter comprising just two species from South America (Kubitzki
2007), being the Chilean endemic Quillaja saponaria one of the most characteristic
trees of Mediterranean Chile. The Fagales is represented in Chile by the Myricaceae
and Nothofagaceae, the latter represented by10 species that are highly characteris-
tic of Central/South Chilean landscapes (Chap. 9). The Rosales are represented in
Chile by 3 families, 21 genera and 71 species. The Malpighiales are represented
by 10 families, with most genera (7) in the Euphorbiaceae. The Passifloraceae
consist in Chile of only one species, while the monogeneric Malesherbiaceae
is represented by 16 species, 13 of them endemic. The Malpighiaceae are
represented by two endemic genera, Dinemandra and Dinemagonum. Within
Malpighiales the species-rich genus Viola (Violaceae) also is remarkable
(Table 2.5).
Malvids are represented in Chile by 5 orders: Geraniales, Myrtales, Malvales,
Brassicales, and Sapindales. The Geraniales includes the Geraniaceae, Vivianiaceae,
and Meliathaceae. This last one shall include the endemic Chilean Francoaceae,
according to APGIII (2009) (Table 2.6). Within the Malvales, the Malvaceae
comprise 75 species. The Sapindales include the Sapindaceae, Anacardiaceae,
Rutaceae. The Brassicaceae is the richest family in the Brassicales, with 145 Chilean
species.
Asterids
This clade constitues one if the major lineages within the Angiosperms. It comprises
about 100 families and more than 80,000 species (Bremer 2009). Asterids have been
recently organized in two main clades: lamiids and campanulids. Sister to them
are the Cornales and Ericales (APG III 2009), both with representatives in Chile
(47 and 31 species respectively). Former Empetraceae are considered as nested
within the Ericaceae (Kron and Luteyn 2005).
Lamiids include in Chile the Gentianales, Lamiales, Solanales, and the
Boraginaceae. In Chile one of the species-richest order is the Lamiales, that
comprises 14 families, 58 genera, and 270 species (124 endemics). Species-rich
families are the newly recognized Calceolariaceae (Olmstead et al. 2001), the
Verbenaceae, and the Veronicaceae (Table 2.4). Veronicaceae is an alternative name
to Plantaginaceae (Tank et al. 2006), and resulted from the disintegration of the
former Scrophulariaceae. Most Chilean former Scrophulariaceae have been trans-
ferred to the Calceolariaceae (Calceolaria, Jovellana), Linderniaceae (Lindernia),
Phrymaceae (Mimulus), Orobanchaceae (Agalinis, Bartsia, Castilleja, Euphrasia,
Orobanche), or Veronicaceae (14 genera, see Appendix). The Scrophulariaceae still
conserve the genera Alonsoa, Bluddleja (formerly Buddlejaceae), and Limosella.
The Solanales comprise only two families (Convolvulaceae and Solanaceae),
but 28 genera, and 193 species (93 endemics). The species-rich genus Nolana
(Table 2.5), formerly pertaining to the Nolanaceae, is considered currently as part
of the Solanaceae (Dillon et al. 2009).
The Boraginaceae have not yet gained order status, and comprise in Chile 13
genera and 103 species (66 endemic).
Campanulids comprise the Aquifoliales (including Citronella in the
Cardiopteridaceae, formerly classified in the Icacinaceae); the big Asterales,
Escalloniales, Bruniales, Dipsacales, and Apiales.
The Asterales include 11families, five of them present in Chile: Asteraceae,
Calyceraceae, Campanulaceae (including Lobeliaceae), Goodeniaceae, and
Stylidiaceae (including Donatiaceae). No wonder that the Asterales are the best
represented order in the flora of Chile, with 137 genera, and 883 species (395
of them endemic); the sole presence of representatives of the Asteraceae and the
closely related Calyceraceae and Goodeniaceae compose much of these numbers
(Table 2.4, Chap. 8). The Escalloniales consists of the family Escalloniaceae, rep-
resented in Chile by 3 genera (Escallonia, Tribeles, Valdivia), and 16 species
(7 endemic). Chilean Campanulaceae consist of 6 genera; remarkably are the
endemic genus Cyphocarpus from the Atacama Desert, and Wahlenbergia, with
several species endemic to Juan Fernández. The Bruniales include the monospe-
cific family Desfontainiaceae, alternatively nested within Columelliaceae (APGIII
2009). The Dipsacales include the Chilean Valerianaceae (genera Stangea and
Valeriana), alternatively classified under the Caprifoliaceae (APGIII 2009). The
Chilean Apiales include three families (Apiaceae, Araliaceae, and Griseliniaceae),
comprising 26 genera, and 106 species (46 endemic).
Box 2.4 Angiosperms Temporal and Spatial Origin(s)

The origin of the flowering plants is considered as the “Holy Grail” of botany
(Miller 2009), or “Darwin’s abominable mystery” (Crepet 1998, Davies et al.
2004). From fossil evidence, a major radiation of angiosperms is obvious in
the mid-Cretaceous, (ca. 130 mya) (Lidgard and Crane 1990).
The fossil record is but not unequivocal: Sanmiguelia, a Late Triassic fos-
sil described by Brown (1956), has been interpreted by Cornet (2002) as
a very primitive angiosperm that combines both monocot and dicot char-
acters, but this interpretation has been ignored by other palaeobotanists.
Even the genus Archaefructus proposed as a Jurassic primitive angiosperm
(Sun et al. 1998, 2002) has been very controversial. In this respect there
is still a high level of uncertainty: the Holy Gray of botany is still a great
mystery (Frohlich and Chase 2007). “If the gymnosperms are indeed mono-
phyletic, their sister-group the angiosperms must date from the same period,
the Carboniferous. This leaves a gap of over 150 million years with no fos-
sil record of angiosperms – a period longer than their entire known fossil
history. This could be either because the gymnosperms are not a natural
group or because the stem lineage of the angiosperms lacked distinguishing
angiosperm synapomorphies” (Hill 2005).
Molecular dating shows a middle Jurassic origin of flowering plants
(ca 175 mya) (Wikström et al. 2001; Magallón 2009), suggesting that the
angiosperms may be older than the fossil record indicates, and “predating the
oldest unequivocal fossil angiosperms by about 45 million years” (Bell et al.
2005). Other molecular dating analyses even suggest a pre-Mesozoic age for
the divergence of the angiosperm lineage from other seed plants (Miller 2009).
This is an old idea: “The angiosperms undoubtedly originated long before
the Cretaceous period. The specialised character and astonishingly modern
facies of many Cretaceous angiosperms confirm our belief in an antiquity
of angiosperms antedating by many millions of years, probably by several
geologic periods, the first appearance of recognisable pioneers of the present
ruling dynasty in the modern world” (Seward 1933 as quoted by Takhtajan
1961, p 122). Early authors postulated a Triassic or even Permo-Triassic origin
(Wieland 1933; Camp 1947; Zimmermann 1959): “Angiosperms originated
during the Permian as the aftermath of the so called Permo-Carboniferous
Glaciations” (Croizat 1960, p 1735).
Regards the geographic origin of the angiosperms, high contrasting views
developed (Axelrod 1952; Retallack and Dilcher 1981; Lu and Tang 2005).
Stebbins (1974) proposed that alpine biomes of northern latitudes might have
been the centre of early radiation of angiosperms. An eastern Asian centre
of origin was put forth by Sun et al. (2001, 2008). Others postulated that
flowering plants evolved from multiple, unrelated seed plant lineages and
areas, i.e. a polyphyletic-polychromic-polytopic hypothesis (Wu et al. 2002).
References 79
Stuessy (2004) proposed that angiosperms evolved slowly from seed ferns
in the Jurassic, beginning first with the carpel, followed later by double fer-
tilization, and lastly by the appearance of flowers. These three fundamental
transitions may have taken more than 100 million years to complete! This
topic is far from been solved, and others maintain that it is an illusion the
search for a strict centre of origin (see Sect. 3.3).
References
APG II [Angiosperm Phylogeny Group II] (2003) An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering plants: APGII. Bot J Linn Soc 141:
399–436
APG III (2009) An update of the Angiosperm Phylogeny Group classification for the orders and
families of flowering plants: APGIII. Bot J Linn Soc 161:105–121
Anderson CL, Janssen T (2009) Monocots. In: Hedges SB, Kumar S, (eds) The timetree of life.
Oxford University Press, New York, pp. 203–212
Axelrod AJ (1952) A theory of angiosperm evolution. Evolution 6:29–60
Bell ChD, Soltis DE, Soltis PS (2005) The age of the angiosperms: a molecular timescale without
a clock. Evolution 59:1245–1258
Brako L, Zarucchi JL (1993) Catalogue of the flowering plants and gymnosperms of Peru. Reprint
1996. Monographs in Systematic Botany from the Missouri Botanical Garden, vol. 45. Missouri
Botanical Garden, St. Paul, MO, 1–1286
Bremer K (2000) Early Cretaceous lineages of monocot flowering plants. PNAS 97:4707–4711
Bremer K (2002) Gondwanan evolution of the grass alliance of families (Poales). Evolution
56:1374–1387
Bremer B (2009) Asterids. In: Hedges SB, Kumar S, (eds) The timetree of life. Oxford University
Press, New York, pp. 177–187
Brown RW (1956) Palm-like plants from the Dolores Formation (Triassic), southwestern Colorado.
US Geol Surv Prof Pap 274:205–209
Burleigh JG, Mathews S (2004) Phylogenetic signal in nucleotide data from seed plants:
implications for resolving the seed plant tree of life. Am J Bot 91:1599–1613
Camp WH (1947) Distribution patterns in modern plants and the problem of ancient dispersals.
Ecol Monogr 17:159–183
Castro SA, Camousseight A, Muñoz-Schick M, Jaksic FM (2006) Rodulfo Amando Philippi, el
naturalista de mayor aporte al conocimiento taxonómico de la diversidad biológica de Chile.
Rev Chil Hist Nat 79:133–143
Cavanilles AJ (1801) Del género Nuevo Francoa. An Cienc Nat 4:236–239
Christensen C (1934) Index filicum. Copenhagen
Cornet B (2002) Last updated on 10/10/2008. When Did Angiosperms First Evolve? (http://www.
sunstar-solutions.com/sunstar/Why02/why.htm)
Crane PR, Friis EM, Pedersen KR (1994) Palaeobotanical evidence on the early radiation of
magnoliid angiosperms. Plant Syst Evol 8(Suppl):51–72
Crane PR, Herendeen P, Friis EM (2004) Fossils and plant phylogeny. Am J Bot 91:1683–1699
Crepet WL (1998) The abominable mystery. Science 282:1653–1654
Croizat L (1960) Principia Botanica. Published by the author. Caracas, Venezuela
Cronquist A (1981) An integrated system of classification of flowering plants. Columbia University

Press, Nueva York
Darwin C (1839) Narrative of the surveying voyages of His Majesty’s Ships Adventure and Beagle
between the years 1826 and 1836, describing their examination of the southern shores of South
America, and the Beagle’s circumnavigation of the globe. Henry Colburn, London
Davies TJ, Barraclough TG, Chase MW, Soltis PS, Soltis DE, Savolainen V (2004) Darwin’s
abominable mystery: Insights from a supertree of the angiosperms. Proc Natl Acad Sci USA
101:1904–1909
De Lange PJ, Sawyer JWD, Rolfe JR (2006) New Zealand indigenous vascular plant checklist.
New Zealand Plant Conservation Network, Wellington
Dillon MO, Tu T, Xie L, Quipuscoa Silvestre V, Wen J (2009) Biogeographic diversification in
Nolana (Solanaceae), a ubiquitous member of the Atacama and Peruvian Deserts along the
western coast of South America. J Syst Evol 47:457–476
Ebihara A, Dubuisson J-Y, Iwatsuki K, Hennequin S, Ito M (2006) A taxonomic revision of
hymenophyllaceae. Blumea 51:221–280
Engler A, Diels L (1936) Syllabus der Pflanzenfamilien, 11th ed. Gebrüder Borntraeger, Berlin
Forest F, Chase MW (2009) Eudicots. In: Hedges SB, Kumar S, (eds) The timetree of life. Oxford
University Press, New York, pp 169–176
Frohlich MW, Chase MW (2007) After a dozen years of progress the origin of angiosperms is still
a great mystery. Nature 450:1184–1189
Fuentes F (1913) Reseña botánica sobre la Isla de Pascua. Bol Mus Nac Hist Nat 5:320–337
Galtier J, Phillips TL (1996) Structure and evolutionary significance of palaeozoic ferns. In:
Camus JM, Johns RJ, Gibby M (eds) Pteridology in perspective: pteridophyte symposium
“95. Proceedings of the Holttum memorial pteridophyte symposium, Kew 1995. Royal Botanic
Gardens Kew, pp 417–433
Gay C (1845–1854) Historia Física y Política de Chile según documentos adquiridos en esta
República durante doce años de residencia en ella y publicada bajo los auspicios del Supremo
Gobierno (including two illustrated atlases). Paris
Gensel PG (2008) The earliest land plants. Annu Rev Ecol Evol Syst 39:459–477
Gensel PG, Berry CM (2001) Early lycophyte evolution. Am Fern J 91:74–98
Goldblatt P, Manning J (2000) Cape plants: a conspectus of the Cape Flora of South Africa.
Strelitzia (National Botanical Institute of South Africa) 9:1–743
Golte W (1993) Araucaria: Verbreitung und Standortanspruche einer Coniferengattung in vergle-
ichender Sicht. Erdwissenschaftliche Forschung vol 27. Steiner Verlag, Stuttgart
Goremykin VV, Viola R, Hellwig FH (2009) Removal of noisy characters from chloroplast
genome-scale data suggests revision of phylogenetic placements of amborella and ceratophyl-
lum. J Mol Evol 68:197–204
Graham CH, Ferrier S, Huettmand F, Moritz C, Townsend Peterson A (2004) New developments
in museum-based informatics and applications in biodiversity analysis. Trend Ecol Evol 19:
497–503
Hassler M, Swale B (2002 onwards) Checklist of World Ferns. (http://homepages.caverock.net.nz/
~bj/fern/list.htm)
Heckman-enlargethispage-12pt DS, Geiser DM, Eidell BR, Stauffer RL, Kardos NL, Hedges SB
(2001) Molecular evidence for the early colonization of land by fungi and plants. Science
293:1129–1133
Henry RJ (2005) Plant diversity and evolution. CABI Publishing, Cromwell Press, London
Herbst R, Melchor R, Troncoso A (1998) Las Pteridophyta y el paleoambiente de la parte media
de la Formación La Ternera (Triásico Superior), en Quebrada La Cachivarita, III Región, Chile.
Rev Geol Chile 25:85–107
Herbst R, Troncoso A (2000) Las Cycadophyta del Triásico de las Formaciones La Ternera y El
Puquén (Chile). Ameghiniana 37:283–292
Herbst R, Troncoso A, Muñoz J (2005) Las Tafofloras Triásicas de la Región de Los Lagos, Xma
Región, Chile. Ameghiniana 42:377–394
References 81
Hill K (2005) Diversity and evolution of gymnosperms. In: Henry RJ, (ed) Plant diversity and
evolution: genotypic and phenotypic variation in higher plants. CABI Publishing, Cromwell
Press, London, pp 25–44
Hilton J, Wang S-J, Tian B, Li C-H (2003) Evidence for conifer origins and early evolution from
the Late Palaeozoic Cathaysian flora of South East Asia. Acta Hort 615:59–65
Hooker JD (1844–1859) The botany of the Antarctic voyage of H.M. Discovery ships Erebus and
Terror in the years 1839–1843. Under the command of Captain Sir James Clark Ross/by Joseph
Dalton Hooker. Reeve, London
Hutchinson J (1926) The families of flowering plants. Vol I Dicotyledons. The Macmillan Co,
London
Hutchinson J (1934) The families of flowering plants. Vol II Monocotyledons. The Macmillan Co,
London
Janssen T, Bremer K (2004) The age of major monocot groups inferred from 800 rbcL sequences.
Bot J Linn Soc 146:385–398
Johnston IM (1929a) The coastal flora of the departments of Chañaral and Taltal. Contributions
from the Gray Herbarium of Harvard University 85:1–137
Johnston IM (1929b) The flora of the Nitrate Coast. Contributions from the Gray Herbarium of
Harvard University 85:138–163
Johow F (1896) Estudios sobre la flora de las Islas de Juan Fernández. Imprenta Cervantes,
Santiago
Jørgensen PM, León-Yáñez S, (eds) (1999) Catalogue of the vascular plants of Ecuador.
Monographs in Systematic Botany vol 75. Missouri Botanical Garden, St. Paul, MO, pp 1–1182
Kenrick P, Crane PR (1997) The origin and early diversification of land plants: a cladistic study.
Smithsonian Institution Press, Washington DC
Kershaw P, Wagstaff B (2001) The southern conifer family Araucariaceae: history, status, and value
for paleoenvironmental reconstruction. Annu Rev Ecol Evol Syst 32:397–414
Knoll AH (1984) Patterns of extinction in the fossil record of vascular plants. In: Nitecki MH, (ed)
Extinctions. University of Chicago Press, Chicago, pp 21–68
Kron KA, Luteyn JL (2005) Origins and biogeographic patterns in Ericaceae: new insights from
recent phylogenetic analyses. Biol Skr 55:479–500
Kubitzki K (2007) Quillajaceae. In: Kubitzki K, (ed) The families and genera of vascular plants,
vol IX. Springer, Berlin, pp 407–408
Leppe M, Moisan P (2003) New records of Upper Triassic Cycadales and Cycadeoidales of Biobio
river, Chile. Rev Chil Hist Nat 76:475–484
Leppe M, Moisan P, Abad E, Palma-Heldt S (2006) Paleobotánica del Triásico Superior del valle
del Río Biobío, Chile: Clase Filicopsida. Rev Geol Chile 33:81–107
Lewis GP, Schrire B, Mackinder B, Lock M (2005) Legumes of the World. Royal Botanic Gardens,
Kew
Lidgard SH, Crane PR (1990) Angiosperm diversification and Cretaceous floristic trends: a
comparison of palynofloras and leaf macrofloras. Paleobiology 16:77–93
Linder HP, Rudall PJ (2005) Evolutionary history of Poales. Annu Rev Ecol Syst 36:107–124
Looser G (1934–1937) Geografía Botánica de Chile. Imprenta Universitaria, Santiago (reprint
2010)Translation of Reiche (1907)
Lu A-M, Tang Y-C (2005) Consideration on some viewpoints in researches of the origin of
angiosperms. Acta Phytotaxonomica Sinica 43:420–430
Magallón S (2009) Flowering plants (Magnoliophyta). In: Hedges SB, Kumar S, (eds) The timetree
of life. Oxford University Press, New York, pp 161–165
Magallón S, Hilu KW (2009) Land plants (Embryophyta). In: Hedges SB, Kumar S, (eds) The
timetree of life. Oxford University Press, New York, pp 133–137
Marticorena C (1990) Contribución a la estadística de la flora vascular de Chile. Gayana Botánica
47:85–113
Marticorena C (1992) Bibliografía Botánica Taxonómica de Chile. Monographs in Systematic
Botany vol. 41. Missouri Botanical Garden, St. Louis, MO
Marticorena C (1995) Historia de la exploración botánica en Chile. In: Marticorena C, Rodríguez

R, (eds) Flora de Chile, vol 1. Universidad de Concepción, Concepción, pp 1–62
Marticorena C (1996) Bibliografía Botánica Taxonómica de Chile. Gayana Botánica
53(Suplemento 1):1–263
Marticorena C, Quezada M (1985) Catálogo de la flora vascular de Chile. Gayana Bot 42:1–157
Marticorena C, Rodríguez R, (eds) (1995 onwards) Flora de Chile, 2 vols. Universidad de
Concepción, Concepcion
Marticorena C, Stuessy TF, Baeza CM (1998) Catalogue of the vascular flora of the Robinson
Crusoe or Juan Fernández Islands, Chile. Gayana Bot 55:187–211
Miller JM (2009) The Origin of Angiosperms. Online, updated January 15, 2009, URL
http://www.gigantopteroid.org/html/angiosperm.htm
Molina JI (1776) Compendio della storia geografica, naturale, e civile del Regno del Chile.
Molina JI (1782) Saggio sulla storia natural del Chili. Bologna, 2d edition (1810), Bologna.
Spanish translation (1987) by R Jaramillo, Santiago
Morawetz W, Röser M, (eds) (1998) Eduard Friedrich Poeppig 1798–1868. Gelehrter und
Naturforscher in Südamerika anlässlich seines 200. Geburtstages. Universität Leipzig, Leipzig
Muñoz Pizarro C (1944) El itinerario de don Claudio Gay. Bol Mus Nac Hist Nat (Chile) 22:
27–44
Muñoz Pizarro C (1959) Sinopsis de la Flora Chilena, 2nd ed (1966). Ediciones Universidad de
Chile, Santiago
Muñoz Pizarro C (1966) Flores Silvestres de Chile. Ediciones de la Universidad de Chile, Munich
Muñoz-Schick M (1975) Geronimo de Bilbar, notable observador naturalista en la alborada de la
Conquista. Bol Mus Nac Hist Nat (Chile) 34:5–27
Muñoz-Schick M (1991) 100 años de la Sección Botánica del Museo Nacional de Historia Natural
(1889–1989. Bol Mus Nac Hist Nat (Chile) 42:181–202
Muñoz-Schick M, Moreira-Muñoz A (2003) Alstroemerias de Chile: diversidad, distribución y
conservación. Taller La Era, Santiago
Muñoz-Schick M, Moreira-Muñoz A (2008) La colección de Tipos del Herbario SGO: avances
hacia el Herbario Nacional Virtual. Resúmenes XX Reunión Nacional de Botánica, Olmué,
25–27 septiembre 2008
Nielsen SN (2005) The Triassic Santa Juana Formation at the lower Biobio River, south central
Chile. J S Am Earth Sci 19:547–562
Olmstead RG, DePamphilis CW, Wolfe AD, Young ND, Elisons WJ, Reeves PA (2001)
Disintegration of the Scrophulariaceae. Am J Bot 88:348–361
Palmer JD, Soltis DE, Chase MW (2004) The plant tree of life: an overview and some points of
view. Am J Bot 91:1437–1445
Philippi RA (1860) Viaje al desierto de Atacama hecho por orden del Gobierno de Chile en el
verano 1853–1854 por el Dr. Rodulfo Amando Philippi. E. Anton, Halle
Philippi F (1881) Catalogus plantarum vascularium chilensium adhuc descriptarum. Ann Univ
Chile 59:49–422
Pigafetta A (1536) Primer Viaje en torno del globo. Spanish edition from 1963. Espasa-Calpe,
Madrid
Pissis JA (1875) Atlas de Geografía Física de la República de Chile. Instituto Geográfico de Paris,
Ch. Delagrave, Paris
Poeppig E, Endlicher S (1835–1845) Nova genera ac species plantarum, quas in regno Chilensi
Peruviano et in terra Amazonica :annis MDCCCXXVII ad MDCCCXXXII /legit Eduardus
Poeppig et cum Stephano Endlicher descripsit iconibusque illustravit. Sumptibus F. Hofmeister,
Leipzig
Poole I, Gottwald H (2001) Monimiaceae sensu lato, an element of Gondwanan polar forests:
evidence from the Late Cretaceous – early Tertiary wood flora of Antarctica. Aust Syst Bot
14:207–230
Pryer KM, Schuettpelz E (2009) Ferns. In: Hedges SB, Kumar S, (eds) The timetree of life. Oxford
University Press, New York, pp. 153–156
References 83
Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R (2004) Phylogeny and
evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. Am
J Bot 91:1582–1598
Reiche C (1894–1911) Flora de Chile, 6 vols. Ediciones de la Universidad de Chile, Santiago
Reiche C (1907) Grundzüge der Pflanzenverbreitung in Chile. In: Engler A, Drude O, (eds) Die
Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, Vol VIII. Verlag von
V. Engelmann, Leipzig
Renner SS (2004) Variation in diversity among Laurales, Early Cretaceous to Present. Biol Skr
55:441–458
Renner SS (2009) Gymnosperms. In: Hedges SB, Kumar S, (eds) The timetree of life. Oxford
University Press, New York, pp. 157–160
Retallack G, Dilcher DL (1981) A coastal hypothesis for the dispersal and rise to dominance of
flowering plants. In: Niklas K, (ed) Paleobotany, paleoecology and evolution, vol 2. Praeger,
New York, pp 27–77
Rothwell GW (1987) Complex Paleozoic Filicales in the evolutionary radiation of ferns. Am J Bot
74:458–461
Ruiz EA, Fuentes G, Samuel R, González F et al. (2005) Genetic diversity and differentiation
within and among Chilean populations of Araucaria araucana (Araucariaceae) base don AFLP
markers and allozyme variability. Abstracts XVII International Botanical Congress, Vienna, nº
13.3.5
Saldivia Maldonado Z (2003) La visión de la naturaleza en tres científicos del siglo XIX en Chile:
Gay, Domeyko y Philippi. Instituto de Estudios Avanzados, Universidad de Santiago de Chile,
Santiago
Sanderson MJ (2003) Molecular data from 27 proteins do not support a Precambrian origin of land
plants. Am J Bot 90:954–956
Schneider H, Schuettpelz E, Pryer KM, Cranfill R, Magallón S, Lupia R (2004) Ferns diversified
in the shadow of angiosperms. Nature 428:553–557
Scotland RW, Wortley AH (2003) How many species of seed plants are there?. Taxon 52:101–104
Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG (2006) A classification for
extant ferns. Taxon 55:705–731
Stebbins GL (1974) Flowering plants: evolution above the species level. Belknap Press, Cambridge
Stevens PF (2001 onwards) Angiosperm Phylogeny Website. Version 9, June 2008. URL http://
www.mobot.org/MOBOT/research/APweb/
Stewart WN, Rothwell GW (1993) Paleobotany and the evolution of plants, 2d ed.. Cambridge
University Press, Cambridge
Stockey RA (1982) The Araucariaceae: an evolutionary perspective. Rev Palaeobot Palynol
37:133–154
Stuardo C, Feliú G (1973) Vida de Claudio Gay (1800–1873). Editorial Nacimiento, Santiago de
Chile, vol 2, pp 318–329
Stuessy TF (2004) A transitional-combinational theory for the origin of angiosperms. Taxon 53:
3–16
Sun G, Dilcher DL, Zheng S-L, Zhou Z (1998) In search of the first flower: a jurassic angiosperm,
Archaefructus, from Northeast China. Science 282:1692–1695
Sun G, Zheng S-L, Dilcher DL, Wang YD, Mei SW (2001) Early Angiosperms and their associ-
ated plants from Western Liaoning, China. Shanghai Scientific and Technological, Education
Publishing House, Shanghai
Sun G, Ji Q, Dilcher DL, Zheng S-L, Nixon KC, Wang X (2002) Archaefructaceae, a new basal
angiosperm family. Science 296:899–904
Sun G, Ji Q, Dilcher DL, Zheng S-L (2008) A review of recent advances in the study of early
angiosperms from northeastern China. Palaeoworld 17:166–171
Takhtajan A (1961) The origin of Angiospermous Plants, 2nd ed. (original Russian edition,
Moscow). Traslated as Flowering plants, origin and dispersal by C. Jeffrey, Kew (1969). Reprint
1981. Otto Koeltz Science Publishers, Koenigstein
Tank DC, Beardsley PM, Kelchner SA, Olmstead RG (2006) Review of the systematics of
Scrophulariaceae s.l. and their current disposition. Aust Syst Bot 19:289–307
Taylor CM, Muñoz-Schick M (1994) The botanical works of Philippi, father and son, in Chile.
Ann Mo Bot Gard 81:743–748
Torres T, Philippe M (2002) Nuevas especies de Coniferales y Ginkgoales del Lías de La Ligua
(Chile) con una evaluación del registro paleoxilológico en el Jurásico de Sudamérica. Rev Geol
Chile 29:151–165
Troncoso A, Herbst R (1999) Triassic Ginkgoales from northern Chile. Rev Geol Chile 26:255–273
von Humboldt A, Bonpland A (1805) Essai sur la Géographie des Plantes. Chez Levrault, Schoell
et compagnie, Paris
Wake DB, Hadly EA, Ackerlya DD (2009) Biogeography, changing climates, and niche evolution.
PNAS 106:19631–19636
Wang L, Qin H (2008) The Chinese Virtual Herbarium (CVH) and its future role for biogeographic
research. Biogeografía 3:15–20. URL http://www.uac.pt/~seba/?q=taxonomy/term/7
Wellman CH, Osterloff PL, Mohiuddin U (2003) Fragments of the earliest land plants. Nature
425:282–285
Wieland GR (1933) Origin of Angiosperms. Nature 131:360–361
Wikström N, Savolainen V, Chase MW (2001) Evolution of the angiosperms: calibrating the family
tree. Proc R Soc Lond B 268:2211–2220
Willis KJ, McElwain JC (2002) The evolution of plants. Oxford University Press, Oxford
Wilton AD, Breitwieser I (2000) Composition of the New Zealand seed plant flora. New Zealand
J Bot 38:537–549
Wu Z-Y, Lu A-M, Tang Y-C, Chen Z-D, Li W-H (2002) Synopsis of a new ‘polyphyletic-
polychromic-polytopic’ system of the angiosperms. Acta Phytotaxon Sin 40:289–322
Zavattieri AM, Herbst R, Muñoz BJ (2003) Microflora de la Formación Panguipulli (Triásico
Superior), en Lincán Ray, Lago Calafquén, 10◦ Región, Chile. Ameghiniana 40:585–600
Zimmermann W (1959) Die Phylogenie der Pflanzen, 2d edn. Gustav Fischer Verlag, Stuttgart
Zuloaga FO, Morrone O, Rodríguez D (1999) Análisis de la biodiversidad en plantas vasculares
de la Argentina. Kurtziana 27:17–167
Zuloaga FO, Morrone O, Belgrano MJ, (eds) (2008) Catálogo de las Plantas Vasculares del Cono
Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monographs in Systematic Botany
from the Missouri Botanical Garden, vol. 107:3 vols. Missouri Botanical Garden, St. Paul, MO
Part II
Chorology of Chilean Plants
Chapter 3
Geographical Relations of the Chilean Flora
Abstract The chorological approach attempts to classify taxa according to their

geographic relationships. The Chilean native genera have been classified in 7 floris-
tic elements and 10 generalized tracks. The origin of major disjunctions in the
flora is intensively discussed from the viewpoint of under contrasting biogeographic
paradigms like dispersal and vicariance. Attention is also put on the influence of
the Andean Dry Diagonal on the evolution of the Chilean flora and its distribution
patterns.
Chorology is an old approach in biogeography that deals specifically with com-

paring distribution maps. In Cain’s (1944) opinion, the most comprehensive con-
cept encompassing the analysis of distributions would be the old German term
Arealkunde, the “science of area” (sensu Meusel 1943; Meusel and Jäger 1992).
Used more or less like a synonym, the term “chorology” is “still in use, in quite a
variety of different ways, but many modern studies do appear to be within that remit”
(Williams 2007, p 29). Chorology has it strength in its comparative nature . . .” area
maps will actually show interesting distribution conditions, others will only obtain
its value in the relationship with other areas” (Hannig and Winkler 1926–1940)
(Fig. 3.1).
Chorology is in line with Parenti and Ebach’s comparative biogeography (see
Chap. 10), and is closer linked to systematics (Rothmaler 1955; Stuessy et al. 2003;
Parenti and Ebach 2009). It is still the most suited term for remitting to the science
of geographic distributions.
3.1 Floristic Elements

A product of the comparison of distribution maps is the classification of floras
into floristic elements (Christ 1867; Engler 1882; Wangerin 1932; Good 1947;
Takhtajan 1986). Wulff (1950) described five different approaches to define floristic
elements, but recognized that “most investigators have been inclined to believe that
the geographical factor is of primary importance and that the term element should

DOI 10.1007/978-90-481-8748-5_3, ⃝
88 3 Geographical Relations of the Chilean Flora
Fig. 3.1 Donat’s distribution maps of the genera Ourisia and Caltha, in Hannig and Winkler
(1926–1940)
be applied to it” (Wulff 1950, p 204). In any case Wulff (1950) was of the opin-
ion that geographical [floristic] elements are fundamental for an understanding of
a flora, and that an analysis of a flora should begin with these elements. Recently,
floristic elements have been analysed for the North American flora north of México
(Qian 1999), the East Asian flora (Qian et al. 2003), the flora of the Bolivian Andean
valleys (López 2003), and the Ecuadorian superpáramo (Sklenar and Balslev 2007).
In the case of the Chilean flora, the first attempt is the one of Reiche (1907). He
was able to identify seven elements or Kontingente (Table 3.1). Later, Villagrán
3.1 Floristic Elements 89
Table 3.1 Floristic elements in the Chilean Flora
Kontingente (Reiche 1907) Elements (Villagrán and Hinojosa 1997)
Tropical-American Neotropical (wide/disjunct)

Andes Chile/Argentina Endemic subantarctic forests/Chilean endemic
California/Mexican –
Antarctic Australasiatic (warm/cool)
Boreal (Europa, S Chile) Wide
Pantropic Pantropical
Introduced flora –
and Hinojosa (1997) applied a similar classification to the woody genera of the
Chilean temperate forests. They described eight floristic elements, including fossil
taxa (Table 3.1).
A complementary approach to floristic elements is the so called “track analy-
sis”, which has been integrated into biogeographical analysis by Croizat (1952,
1958).1 This approach has been successfully used for the analysis and interpre-
tation of disjunct distribution patterns (see Craw et al. 1999). Worth of mention
here are the assessment of the Andean biota by Katinas et al. (1999); the biogeo-
graphic analysis of the Australian flora (Crisp et al. 1999); the analysis of North
American Onagraceae (Katinas et al. 2004), the analysis of the Mexican cloud
forests (Luna Vega et al. 2000) and the evolution of the Andean genus Adesmia
series Microphyllae (Mihoc et al. 2006).
Hereafter, Reiche’s (1907) attempt to analyse the geographical relationships of
the Chilean vascular flora as a whole will be replicated under current floristic and
chorological knowledge. Floristic elements have been usually recognized at the
genus level (e.g. Good 1947), and this approach will be followed here. The use
of the genus as analytical unit is not free of critique, but has the advantage of being
systematic and replicable. The global distribution for each genus has been obtained
from Wielgorskaya (1995) and Mabberley (1997, 2008). These global accounts
have been checked with available monographs. As resumed in Sect. 2.2, by cur-
rent knowledge, the Chilean flora is composed of 837 genera. Analysing the global
distribution of each genus, 7 floristic elements were identified, plus 10 generalized
tracks that eminently represent wide disjunct distributions (Table 3.2 and Fig. 3.17).
The geographical classification of each genus is presented in Appendix A.
The dominant element is clearly the Neotropical one, with almost one third
of the genera. It is followed by the Antitropical and Cosmopolitan element.
The Pantropical, South-temperate and Endemic elements are equally important
1 Croizat apparently took the idea from van Steenis (1962) or maybe from C Skottsberg? (see Craw
1988 in Chap. 10).
90
Table 3.2 Floristic elements for the total vascular flora (updated January 2010)
Floristic elements N◦ Genera
1. Pantropical 89 Pantropical
Cosmopolitan
2. Australasiatic 58 11%
16%
3. Neotropical 221 Austral-asiatic
4. Antitropical 156 7%
5. South-temperate 91
6. Endemic 83
7. Cosmopolitan 139
Total 837
Endemic
10%
South- Neotropical
temperate 26%
11%
Anti-tropical
19%
3 Geographical Relations of the Chilean Flora
reasching around 10%, while the smallish element is the Australasiatic. All elements
will be revised hereafter.
3.1.1 Pantropical Floristic Element

Genera grouped in this element grow mainly in tropical regions (pantropics),
sometimes extending into subtropical and/or temperate areas (Fig. 3.2). This
element includes 89 genera. The families with the greatest number of gen-
era are Asteraceae (9 genera), Fabaceae (6), Poaceae and Cyperaceae (5 each).
The most species-rich genera are Dioscorea, Conyza, and Nicotiana. Few gen-
era occur over the whole country (e.g Conyza), other are restricted to northern
Chile (e.g. Notholaena, Spilanthes, Gomphrena). Most are found in central-south
Chile, disjunct from the rest of their distribution area (e.g. Cryptocarya, Cissus,
Glinus, Passiflora, Dodonaea, Dennstaedtia, Sigesbeckia, Pelletiera, Pouteria,
Beilschmiedia, Wolffiella, Mikania) (see examples in Fig. 3.3).
Note: Megalastrum and Lippia are included in this element despite being
restricted to the American and African tropics. Alonsoa was included here, too,
even though the majority of its species are found in the Neotropics, because two of
its species occur in South Africa.
a b
c d
e f
Fig. 3.2 Distribution of several genera exemplifying the pantropical floristic element: a Acacia
(Fabaceae); b Bacopa (Veronicaceae); c Cotula (Asteraceae); d Lobelia (Campanulaceae);
e Nicotiana (Solanaceae); f Passiflora (Passifloraceae) (diverse sources in Moreira-Muñoz (2007))
a b
c d
Fig. 3.3 Representative species of genera pertaining to the pantropical element: a Passiflora pin-
natistipula, Quebrada del Tigre; b Lobelia excelsa, Ocoa; c Aristolochia bridgesii; Río Hurtado;
d Croton chilensis, Paposo (photo credits: a–d A. Moreira-Muñoz)
3.1.2 Australasiatic Floristic Element

This element comprises genera from the South Pacific, i.e. Australasia as well
as South America, and the Pacific islands (Fig. 3.4). Some genera are restricted
to temperate subantarctic latitudes; others extend their distribution into tropical
regions in South-East Asia or in South America. The components of this element
a b
c d
e f
g h
Fig. 3.4 Distribution of several genera exemplifying the australasiatic floristic element:
a Berberidopsis (Berberidopsidaceae); b Hebe (Veronicaceae); c Jovellana (Calceolariaceae);
d Lagenophora (Asteraceae); e Leptinella (Asteraceae); f Oreomyrrhis (Apiaceae); g Coprosma
(Rubiaceae); h Rytidosperma (Poaceae) (diverse sources in Moreira-Muñoz (2007))
can be traced to the Gondwana era, as a once-continuous cool-temperate flora, now

scattered into a relict distribution by tectonic movements. Therefore some authors
recognize explicitly the austral floras as a “Gondwanan element” (Barlow 1981;
Nelson 1981; Hinojosa 2005), building an austral floristic realm (see Sect. 4.2).
The australasiatic floristic element comprises 58 genera (see examples in
Fig. 3.5). The families with the greatest number of genera are Asteraceae (4) and
Poaceae, Apiaceae, Cyperacea, Rubiaceae, and Proteaceae, all with 3 genera. Within
these genera, 3 generalized tracks based on superimposing distributions have been
identified.
Austral-antarctic track
This track comprises genera occurring in southernmost South America, New
Zealand, Eastern Australia and Tasmania. It was previously described as
South Pacific track by Crisp et al. (1999).
b c
d e
Fig. 3.5 Representative species of genera pertaining to the australasiatic element: a Berberidopsis
corallina, Concepción; b Embothrium coccineum, Vilches; c Eucryphia glutinosa, Antuco;
d Orites myrtoidea, Antuco; e Fuchsia magellanica, Los Angeles (photo credits: a M. Muñoz-
Schick; b–e A. Moreira-Muñoz)
It numbers 30 genera, e.g. Luzuriaga, Eucryphia, Prumnopitys, Jovellana,

Raukaua. Most of these genera are restricted to the southern temper-
ate forests, but some of them reach northernmost Chile (e.g. Azorella,
Cortaderia). Some genera reach Central Chile, disappear in the Atacama
and reappear in the northern Andes (e.g. Fuchsia, Lomatia, Prumnopitys).
Haloragis is an austral genus that reaches the Juan Fernández islands, but
not the continent.
Tropical Pacific track
This track comprises genera occurring in southernmost South America, New
Zealand, Eastern Australia and/or Tasmania, some also extending to New
Guinea, Malesia, and even to East Asia.
It numbers 24 genera, e.g. Abrotanella, Araucaria, Nothofagus,
Lagenophora, Coprosma, and Hebe. Many genera are restricted to the south-
ern temperate forests, but a few reach northernmost Chile (e.g. Colobanthus,
Muehlenbeckia). Some reach central Chile, disappear in the Atacama and
reappear in the northern Andes (e.g. Citronella, Uncinia, Oreobolus). Doodia
is widespread in the Pacific till Isla de Pascua, but does not occur in conti-
nental Chile. Dicksonia, Arthropteris, Coprosma, and Santalum are austral
genera that are present in the Juan Fernández Islands, and are also absent
from the continent. The genus Santalum is considered in this account for
analytical purposes, in spite of the fact that due to human pressure, Santalum
fernandezianum is extinct from Juan Fernández (see Box 5.1).
Circum-austral track
This track comprises a few genera that extend their distribution through
Australasia to southern Africa and/or Madagascar. These are only 4 genera:
Ficinia, Nertera, Rumohra, and Wahlenbergia.
3.1.3 Neotropical (American) Floristic Element

Strictly speaking, this is an American element, but Neotropical has been used
for a long time in the phytogeographical literature (Fig. 3.6). This element is the
most important one in the Chilean flora: it includes 221 genera, mostly from the
Asteraceae (53), followed by Cactaceae and Poaceae (each 12), Brassicaceae and
Solanaceae (each 11) (see examples in Fig. 3.7). Four generalized tracks have been
identified within this element:
Wide Neotropical track

This track comprises genera from South America, extending to Mexico, to
the south-western USA or even to southern Canada. The main massing, how-
ever, lies in the intertropics. Many species also occur in Brazil. It numbers
65 genera, e.g. Ageratina, Baccharis, Calceolaria, Dalea, Nasa, Nassella.
Some are found in all of Chile (e.g. Baccharis, Gamochaeta, Calceolaria,
Calandrinia), other just in northern Chile (e.g. Coreopsis, Bouteloua), and
many genera occur in central/south Chile, disjunct from the core neotropical
JF
a b c
d e f
JF
g h i
j k l
Fig. 3.6 Distribution of several genera exemplifying the neotropical floristic element:
a Alstroemeria (Alstroemeriaceae); b Azara (Salicaceae); c Calceolaria (Calceolariaceae);
d Cremolobus (Brassicaceae); e Desfontainia (Desfontainiaceae); f Eremodraba (Brassicaceae);
g Escallonia (Escalloniaceae); h Eudema (Brassicaceae); i Malesherbia (Malesherbiaceae);
j Mecardonia (Veronicaceae); k Nototriche (Malvaceae); l Vasconcellea (Caricaceae) (diverse
sources in Moreira-Muñoz (2007))
a b c
d e
f g
Fig. 3.7 Representative species of genera pertaining to the neotropical element: a Tweedia biros-
trata, Concón; b Chloraea alpina, Vilches; c Caiophora chuquitensis, Chusmiza; d Colletia
ulicina, Los Queñes; e Escallonia pulverulenta, Granizo; f Mentzelia scabra, Río Hurtado;
g Calceolaria cana, Vilches (photo credits: a S. Elórtegui Francioli; b–f A. Moreira-Muñoz;
g Francisco Casado)
distribution area (e.g. Piptochaetium, Stenandrium, Chusquea, Calydorea,

Chaptalia, Ugni).
Wide Andean track
This track comprises genera ranging from southern Chile to Colombia
(and Costa Rica), but does not reach North America. It numbers 29
genera, e.g. Chuquiraga, Myrteola, Polylepis, Geoffroea. Some are found
in all of Chile (e.g. Perezia, Escallonia); others only in northernmost

(Dunalia, Cremolobus) or north-Central Chile (Exodeconus, Nototriche);
many distributions are disjunct between southern Chile/Argentina and north-
ern South America (e.g. Llagunoa, Myrcianthes, Dysopsis, Blepharocalyx,
Desfontainia, Myrteola).
Central Andean track
This track comprises genera found in the Central Andes, in Perú, Bolivia,
northern Argentina and north-central Chile.
It numbers 86 genera, e.g. Loasa, Philibertia, and Oxychloë, com-
prising by far many Asteraceae (18 genera) and Cactaceae (11). Most
genera occur in a continuous range from northern Chile to Peru/
Bolivia (e.g. Acantholippia, Balbisia, Neowerdermannia, Oreocereus,
Tunilla, Pycnophyllum, Philibertia), others occur disjunctly between cen-
tral Chile/Perú (Eccremocarpus, Kageneckia, Tetraglochin, Nassauvia,
Weberbauera). 10 genera are restricted to central/northern Chile and adjacent
Argentina (e.g. Cruckshanksia, Monttea, Urmenetea, Werdemannia, Lenzia,
Kurzamra).
South-Amazonian track
This track comprises genera found in central Chile, northern Argentina,
Uruguay, Paraguay, and south-eastern Brazil.
It numbers 41 genera, e.g. Colliguaja, Quillaja, Azara, Tweedia,
Myrceugenia, Viviania, and Dasyphyllum. Most of these genera occur dis-
junctly between Central Chile and SE Brazil (see Sect. 3.2).
3.1.4 Antitropical Floristic Element

This element comprises genera found both in the northern and in the southern tem-
perate regions, but are absent from the intervening tropics. This pattern is commonly
referred to as the amphi-tropical pattern in the literature, but as W. Welß (pers.
comm.) correctly noted, the most appropriate term would be “antitropic”, since
“amphitropical” means “both tropics”. In Cox’s (1990) opinion, the most appro-
priate term to be used would be “amphitemperate”, but since it includes subtropical
distributions, the most suited term is antitropical (see also Wen et al. 2002; Glasby
2005; Parenti 2007) (Fig. 3.8).
This element includes 156 genera. The families with the greatest number of gen-
era are Asteraceae (26), Poaceae (23), Fabaceae and Polemoniaceae (each 8), and
Boraginaceae (7) (see examples in Fig. 3.9).
Note: Menodora (Oleaceae) is considered as an antitropical genus, since its dis-
tribution range includes both subtropical Americas, and subtropical South Africa
(map in Muñoz-Schick et al. 2006).
One of the naturalists referring to this element was G. Treviranus (1803), even
though he did not recognise it formally. While discussing the components of his
Antarktische Flor, he noticed that there was a floristic relationship between southern
South America and New Zealand, and that some of these genera have their main
ID
a b c
d e f
g h
i j
Fig. 3.8 Distribution of several genera exemplifying the antitropical floristic element:
a Agoseris (Asteraceae); b Frankenia (Frankeniaceae); c Hoffmannseggia (Fabaceae); d Larrea
(Zygophyllaceae); e Mancoa (Brassicaceae); f Tropidocarpum (Brassicaceae); g Androsace
(Primulaceae); h Antennaria (Asteraceae); i Artemisia (Asteraceae); j Euphrasia (Orobanchaceae)
(diverse sources in Moreira-Muñoz (2007))
distribution in the northern and southern temperate zones (e.g. Pinguicula, Salix,
Ribes) (Treviranus 1803, pp 131–132). Du Rietz (1940) and many others called
it the pattern of “bipolar plant distribution”. The antitropical element was treated
in depth in several papers that arose from a symposium at the beginning of the
1960s (Constance 1963) (see Sect. 3.2). The antitropical element includes many
variable distribution patterns, some restricted, and others very wide. Within this
element three generalized tracks were identified.
Wide antitropical track (bipolar-temperate element)

This track comprises genera found in Eurasia, North America, southern
South America, some also ranging into the montane American tropics.
a b
d e
Fig. 3.9 Representative species of genera pertaining to the antitropical element: a Ribes puncta-
tum, Rio Clarillo; b Amblyopappus pusillus, Los Vilos; c Clarkia tenella, Los Molles; d Larrea
nitida, Illapel; e Bahia ambrosioides, Huentelauquén (photo credits: a–e A. Moreira-Muñoz)
It numbers 85 genera, e.g. Astragalus, Fagonia, Valeriana, Vicia. Many of

these genera are found in all of Chile (e.g. Hypochaeris, Cystopteris, Bromus,
Valeriana). Few are restricted to northern Chile (e.g. Woodsia), while most
are restricted to southern Chile (e.g. Rhamnus, Potentilla, Adenocaulon,
Saxifraga), some of them even to the southernmost Magallanes region
(Androsace, Botrychium, Chrysosplenium).
Antitropical Pacific track
All the genera grouped in this track have a disjunct distribution between
south-western North America and South America, occurring mostly in
subtropical and tropical deserts.
This track numbers 58 genera, e.g. Agalinis, Hoffmannseggia, Camissonia,

Tiquilia, and several Polemoniaceae (Gilia, Microsteris, Ipomopsis,
Leptosiphon, Navarretia). Few genera are found along the whole lati-
tudinal gradient in Chile (e.g. Phacelia). Some occur in northern Chile
and the Central Andes (e.g. Mancoa, Cistanthe, Tarasa), but many
are found only in Central Chile and south-western North America
(e.g. Blennosperma, Errazurizia, Hoffmannseggia, Lastarriaea, Navarretia,
Plectritis, Tropidocarpum).
Circum-Pacific track
This track comprises genera with a disjunct distribution in North/Central
America and South America that have also representatives in Australasia. It
numbers 12 genera: Distichlis, Flaveria, Gochnatia, Gaultheria, Lilaeopsis,
Microseris, Plagiobothrys, Sicyos, Soliva, and Weinmannia. Only two
circum-Pacific genera reach South Africa: Acaena and Carpobrotus, while
Weinmannia reaches Madagascar.
3.1.5 South-Temperate Floristic Element

This element comprises genera found in the Andes cordillera in Chile and in
adjacent Argentina (Fig. 3.10).
It includes 91 genera. The families with the greatest number of genera are
Asteraceae (11), Apiaceae (6), and Alliaceae, Brassicaceae, Solanaceae (each with
4 genera) (see examples in Fig. 3.11). The half of the Chilean conifer genera belong
to this element (Austrocedrus, Fitzroya, Pilgerodendron, Saxegothaea). Most gen-
era within this element are restricted to the southern temperate forests (e.g. Boquila,
Drapetes, Embothrium, Laureliopsis) and Patagonia (Eriachaenium, Lecanophora,
Lepidophyllum, Saxifragella, Xerodraba). Some are represented in the temperate
zone but reaching more arid environments to the north, like Triptilion, Nastanthus,
Trichopetalum. The genus Mulinum reaches further north, at 21◦ S.
3.1.6 Endemic Floristic Element
This element comprises genera endemic to continental Chile, and the Chilean Pacific
islands.
It numbers 83 genera, of which 16 pertain to the Asteraceae, 6 to the Cactaceae,
and 4 to the Alliaceae (Table 3.3). Species-rich genera are Copiapoa (22 species),
Leucocoryne (14),2 Dendroseris (11), and Robinsonia (8). Remarkably, most
2 Leucocoryne is an interesting case of taxonomic inflation, because the Southern Cone Checklist
mentions 45 species, most of them lacking justification.
20°
30°
40°
50°
a b c
20°
30°
40°
50°
d e f
20°
30°
40°
50°
g h i
Fig. 3.10 Distribution of several genera exemplifying the South-temperate floristic element:
a Aextoxicon (Aextoxicaceae); b Chiliotrichum (Asteraceae); c Embothrium (Proteaceae);
d Laretia (Apiaceae); e Laureliopsis (Atherospermataceae); f Nardophyllum (Asteraceae);
g Notopappus (Asteraceae); h Philesia (Philesiaceae); i Tepualia (Myrtaceae) (source: monographs
and SGO collections)
c d
Fig. 3.11 Representative species of genera pertaining to the south-temperate element:

a Schizopetalon dentatum, Concón; b Cruckshanksia pumila, Llanos de Challe; c Aextoxicon punc-
tatum, Fray Jorge; d Philesia magellanica. Photo/illustration credits: a, b A. Moreira-Muñoz;
c Walter Welss; d illustration from Macloskie G. (1903–1906) In: Scott WB. Flora Patagonica.
Reports of the Princeton University Expeditions to Patagonia, 1896–1899, Princeton
endemic genera are monospecific, an aspect that was already noticed by Reiche
(1905) (Fig. 3.12) (Table 3.3).
Most genera (67) are endemic to continental Chile (e.g. Adenopeltis,
Balsamocarpon, Huidobria, Leontochir, Pitavia) (Figs. 3.13, 3.14), while 16 gen-
era are endemic to the Chilean Pacific islands, especially Juan Fernández (e.g.
Dendroseris, Lactoris, Cuminia, Juania) (Chap. 5, Tables 5.2 and 5.3). The endemic
element will be further discussed in Chap. 4, and the Juan Fernández flora in
Chap. 5.
Table 3.3 Genera endemic to continental Chile
Genus Family N◦ species Genus Family N◦ species
Legrandia Myrtaceae 1
Adenopeltis Euphorbiaceae 1 Leontochir Alstroemeriaceae 1
Anisomeria Phytolaccaceae 2 Leptocarpha Asteraceae 1
Araeoandra Vivianiaceae 1 Leucocoryne Alliaceae 14
Avellanita Euphorbiaceae 1 Leunisia Asteraceae 1
Bakerolimon Plumbaginaceae 1 Marticorenia Asteraceae 1
Balsamocarpon Fabaceae 1 Metharme Zygophyllaceae 1
Bridgesia Sapindaceae 1 Microphyes Caryophyllaceae 3
Calopappus Asteraceae 1 Miersia Alliaceae 1
Cissarobryon Vivianiaceae 1 Miqueliopuntia Cactaceae 1
Conanthera Tecophilaeaceae 5 Moscharia Asteraceae 2
Copiapoa Cactaceae 22 Neoporteria Cactaceae 6
Cyphocarpus Campanulaceae 3 Notanthera Loranthaceae 1
Desmaria Loranthaceae 1 Ochagavia Bromeliaceae 4
Dinemagonum Malpighiaceae 1 Oxyphyllum Asteraceae 1
Dinemandra Malpighiaceae 1 Peumus Monimiaceae 1
Epipetrum Dioscoreaceae 3 Phrodus Solanaceae 1
Ercilla Phytolaccaceae 2 Pintoa Zygophyllaceae 1
Eriosyce Cactaceae 5 Pitavia Rutaceae 1
Eulychnia Cactaceae 6 Placea Amaryllidaceae 5
Fascicularia Bromeliaceae 1 Pleocarphus Asteraceae 1
Francoa Francoaceae 1 Podanthus Asteraceae 2
Gethyum Alliaceae 2 Reicheella Caryophyllaceae 1
Gomortega Gomortegaceae 1 Sarmienta Gesneriaceae 1
Guynesomia Asteraceae 1 Scyphanthus Loasaceae 2
Gymnachne Poaceae 1 Speea Alliaceae 1
Gypothamnium Asteraceae 1 Tecophilaea Tecophilaeaceae 2
Hollermayera Brassicaceae 1 Tetilla Francoaceae 1
Homalocarpus Apiaceae 6 Thelocephala Cactaceae 6
Huidobria Loasaceae 2 Traubia Amaryllidaceae 1
Ivania Brassicaceae 1 Trevoa Rhamnaceae 1
Jubaea Arecaceae 1 Valdivia Escalloniaceae 1
Lapageria Philesiaceae 1 Vestia Solanaceae 1
Latua Solanaceae 1 Zephyra Tecophilaeaceae 1
60
50
Number of genera
40
30
20
10
Fig. 3.12 Number of 0

endemic genera versus the 1 2 3 4 5 6 8 11 14 22
number of species Number of species
20°
30°
40°
50°
a b c d e
20°
30°
40°
50°
f g h i j
Fig. 3.13 Distribution of several genera exemplifying the endemic floristic element: a Adenopeltis
(Euphorbiaceae); b Balsamocarpon (Fabaceae); c Cyphocarpus (Campanulaceae); d Fascicularia
(Bromeliaceae); e Francoa (Francoaceae); f Huidobria (Loasaceae); g Lapageria (Philesiaceae);
h Leontochir (Alstroemeriaceae); i Peumus (Monimiaceae); j Pitavia (Rutaceae) (source: collec-
tions SGO)
a b c
e f
Fig. 3.14 Representative species of genera pertaining to the endemic element: a Zephyra elegans,
Llanos de Challe; b Pleocarphus revolutus, Cuesta Pajonales; c Bridgesia incisifolia, La Serena; d
Leontochir ovallei, Llanos de Challe; e Speea humilis, Caleu; f Placea amoena, Cuesta el Espino
(photo credits: a–f A. Moreira-Muñoz);
3.1.7 Cosmopolitan Floristic Element

There are not many genera that can be considered as really cosmopolitan (Good
1947), e.g. some aquatic or semi-aquatic plants (Sagittaria, Landoltia, Wolffia),
g j
Fig. 3.14 (continued) g Jubaea chilensis, Limache; h Lapageria rosea, Los Angeles; i Gomortega
keule, specimen at the National Herbarium SGO; j Scyphanthus elegans, Concón (photo credits: g,
h, j A. Moreira-Muñoz; i specimen scanned by Sergio Moreira)
some ferns and fern allies (Adiantum, Isoetes, Lycopodium, Huperzia,) or some
really widespread terrestrial angiosperm genera like Cyperus, Ceratophyllum,
Gnaphalium, Geum, Juncus, Rubus, Ranunculus, Senecio or Scirpus. Most genera
included in this element have a wide distribution in more than two continents and
more than two principal climatic zones (e.g. tropical and temperate) (Fig. 3.15). In
fact, the element should be called subcosmopolitan, but cosmopolitan is more often
used.
a b
c d
e f
g h
Fig. 3.15 Distribution of several genera exemplifying the cosmopolitan floristic element:
a Apium (Apiaceae); b Bidens (Asteraceae); c Eryngium (Apiaceae); d Gnaphalium (Asteraceae);
e Hydrocotyle (Apiaceae); f Lycium (Solanaceae); g Mimulus (Phrymaceae); h Viola (Violaceae)
(sources: Meusel et al. (1978) and Meusel and Jäger (1992))
It numbers 139 genera, some of them occupying the whole country (e.g.
Asplenium, Gnaphalium, Ranunculus, Chenopodium, Silene), some being restricted
to the north (e.g. Salvia, Chloris, Portulaca), and most being found in central
and southern Chile (Ceratophyllum, Samolus, Aphanes, Geum, Glyceria, Panicum,
Hypericum) (see examples in Fig. 3.16). A few genera are restricted to the
Magallanes region (Huperzia, Landoltia).
At the moment, the variety of distributions in this element prevents any attempt
to classify these genera into different tracks, but this will certainly provide an
interesting task for the future.
3.2 To Be or Not To Be Disjunct? 109
c
b
d e
Fig. 3.16 Representative species of genera pertaining to the cosmopolitan element: a Oxalis peni-
cillata, Portillo; b Salvia paposana, Paposo; c Viola montagnei, Laguna Negra; d Euphorbia
lactiflua, Paposo; e Polygala gnidioides, Concón (photo credits: a–e A. Moreira-Muñoz)
3.2 To Be or Not To Be Disjunct?

Analysis of disjunct patterns has been one of the main tasks in biogeography
since the earlier attempts to unravel biogeographic histories (e.g. Hofsten 1916).
Interestingly, disjunctions are often treated as kind of “anomalies” in a suppos-
edly dominant pattern of continuous geographical relationships. In the case of the
Chilean vascular flora, it seems that disjunction is a common pattern and rather
the rule. Most of the generalized tracks that illustrate main distribution patterns of
85
65
24 12 29
58
86 41
30
4
Austral-antarctic Wide Neotropical and Wide Andean

Tropical Pacific Antitropical Pacific and Wide Antitropical
Circumaustral Central Andean and South-Amazonian
Circum-Pacific Panbiogeographic nodes
Fig. 3.17 Generalized tracks representing disjunct Chilean plant distributions. The zone of
confluence of different tracks can be interpreted as a composite biogeographic zone or a
panbiogeographic node (Heads 2004)
the Chilean genera connect territories separated by major physical barriers (oceans,
cordilleras) (Fig. 3.17). Only the wide neotropical track (65 genera) and the Andean
tracks (Wide Andean and Central Andean = 111 genera) encompass territories
on a terrestrial continuum, but many of these genera show indeed a disjunct dis-
tribution pattern (e.g. Ageratina, Bomarea, Dalea). The South-Amazonian track
comprises 39 genera that show a disjunct distribution between southern Chile and
south-eastern Brazil (e.g. Alstroemeria, Azara, Chloraea, Myrceugenia). Another
85 genera compose a wide antitropical (disjunct) track, and 58 genera comprise a
Pacific antitropical track, that connects disjunct genera from Chile/California, and
several more reaching Australasia (11) in a circum Pacific track. The australasi-
atic element is composed of three tracks: an austral-antarctic track (30 genera)
a South Pacific tropical track (24), and a circum-austral track (5 genera). Also
many of the pantropical (e.g. Cryptocarya, Cleome, Croton, Cissus, Passiflora)
and (sub)cosmopolitan genera (e.g. Sanicula, Coriaria) show a disjunction between
their Chilean and their global distribution.
3.2.1 Pacific-Atlantic Disjunctions

Landrum (1981), while analyzing the phylogeny and biogeography of Myrceugenia,
interpreted the disjunct distribution of this genus and other angiosperms between
SE Brazil and central/South Chile as the result of the former existence of a continu-
ous subtropical forest during Oligocene/Miocene times (see Sect. 1.2). According
a b c d
Fig. 3.18 Hypothesized range expansion and posterior fragmentation of the current disjunction
in Myrceugenia during the Cenozoic: a Palaeocene/Eocene; b Oligocene; c Miocene; d current
distribution (simplified from Landrum 1981)
to Landrum (1981), oceanic transgressions, the uplift of the Andes and climate
changes permitted the migration of this range from the south towards the north into
the configuration of the current distribution (Fig. 3.18).
Landrum (1981) noted that many other Chilean genera have similar disjunct dis-
tributions on the continent, such as Araucaria, Bomarea, Crinodendron, Lithrea,
Mutisia, Persea, Podocarpus, Perezia, Quillaja, and Viviania, and Weinmannia.
He interprets this patter as the result of the fragmentation of a continuous
Palaeogene/Neogene flora that occupied the whole subtropical belt before and dur-
ing the emergence of the Andes. The increasing aridity associated to the rise of the
Andes since the Miocene onwards seems to be the cause of the “Dry Diagonal”
and the origin of the isolation of the southern Chile/Argentina temperate forests
(Box 3.1). The hypothesis that the uplift of the Andes fragmented formerly contin-
uous, more mesic to hygric vegetation communities appears already in the earliest
plant geographical work of Reiche about the Chilean composites (see Chap. 8).
Stuessy and Taylor (1995) noticed that the uplift of the Andes is commonly
treated as a recent geological event, but it is a gradual process that has been active for
more than 65 million years from the Palaeogene onwards or even earlier (Riccardi
1988) (Box 1.5). This uplift process seems to be one cause for the high habitat
variability (geodiversity), and related high levels of endemism and plant diversity
in the Andes (Arroyo et al. 1988; Donato et al. 2003; Mutke and Barthlott 2005).
Several recent phylogenetic studies further support a direct association between the
diversification of Andean plants (e.g. Hughes and Eastwood 2006; Scherson et al.
2008) and the major episodes of Andean uplift, from the early Miocene (about 20
mya) to the Pliocene (about 3 mya) (Hooghiemstra and van der Hammen 1998)
(Box 1.5). The Andean uplift must have had a great influence on climate alter-
ations which directly influenced: (a) adaptive processes, (b) plant migrations, and
(c) speciation and extinction rates (Stuessy and Taylor 1995). Vicariant processes
also must have been very important in shaping the evolution and distribution of
South American plant groups (Landrum 1981) (Sect. 3.2).
Box 3.1 The Andean Dry Diagonal and the Fray Jorge
Forest
The “Andean Dry Diagonal” or “Dry Diagonal of South America” is one

of the main features of South American geography (De Martonne 1934;
Messerli et al. 1998). The core of hyperaridity lies in the Atacama Desert,
extending northwards along the Peruvian coast (Fig. 3.19), while semidesertic
conditions dominate in Argentinean ecosystems like el Monte and Patagonia
(Abraham et al. 2009). The origin of the “Andean Dry Diagonal” is close
related to the uplift of the cordillera since the Miocene into the Pliocene,
around 17 mya (Box 1.5).
5°
15°
Hyperarid
Arid
25°
Fray Jorge forest
35°
45°
55°
Fig. 3.19 a Arid Dry Diagonal (adapted from de Martonne 1934; Houston and Hartley
2003); b Fray Jorge Forest (original illustration by E. Sierra in Heusser (1971), by
permission of The University of Arizona Press)
During the Pliocene/Pleistocene, the “Dry Diagonal” established under a

global and regional cooling associated with the onset of Antarctic glacia-
tion, the development of the Circumantarctic and Humboldt currents and the
final Andean uplift (Sect. 1.2). This had at least three main effects on the
north/central Chilean ecosystems (Villagrán et al. 2004):
(a) Fragmentation of the subtropical forests of the southern Cone and their
restriction to the actual Pacific range (southern Chile/Argentina) and the
Atlantic range (southeastern Brazil) (Landrum 1981).
(b) Expansion of the subtropical sclerophyllous forests in central Chile,
adapted to a mediterranean climate.
(c) Fragmentation of the Chilean forests at their northern limit (30◦ S) due
to the increase of aridity in the Atacama, and consequently the relict
character of the forests which now depend on fog, as in the case of Fray
Jorge.
The Fray Jorge Forest (30◦ 40′ S) persists today thanks to the capitation
of fog water. It is surrounded by an arid territory and has long been
of interest to botanists (e.g. Philippi 1884; Muñoz Pizarro and Pisano
1947; Skottsberg 1948; Croizat 1962; Troncoso et al. 1980). This for-
est is considered as the northernmost advance of a plant community
that resembles the southern Valdivian forest. Its main component is the
“olivillo” (Aextoxicon punctatum), a Chilean/Argentinian endemic that is
the sole representative of the Aextoxicaceae, that has its main distribution
in southern Chile (Núñez-Avila and Armesto 2006). Other representatives
of the southern forests that have disjunct populations in Fray Jorge sep-
arated by hundreds of kilometres of the main distribution area to the
South are: Azara microphylla (Salicaceae), Lapageria rosea (Philesiaceae),
Griselinia scandens (Griseliniaceae), Mitraria coccinea and Sarmienta repens
(Gesneriaceae), Nertera granadensis (Rubiaceae), and several species of
ferns (e.g. Hymenophyllum peltatum, Asplenium dareoides, Rumohra adi-
antiformis), mosses and liverworts (Villagrán et al. 2004). According to these
authors, the forest components of the Fray Jorge flora were already present in
the mixed palaeofloras at the beginning of the Cenozoic (Palaeogene). They
correspond to tropical and australasiatic lineages that occupied the Gondwana
continent prior its fragmentation. They developed under warmer and more
humid climatic conditions (Villagrán et al. 2004). The actual floristic char-
acteristics of subtropical forests, affecting Fray Jorge as well as the Central
Chilean sclerophyllous forests, established during the Miocene climatic opti-
mum, when the mixed palaeofloras lost part of the australantarctic element
and were enriched by new neotropical lineages (Sect. 1.2). At that time the
connection with the subtropical forests still existed, under a warmer climate
and a constant rainfall regime from the west and the east.
3.2.2 Antitropical (Pacific) Disjunctions

As revised by Wen and Ickert-Bond (2009), researchers have debated the causes of
the antitropical (=amphitropical) disjunctions for over a century (e.g. Engler 1882;
Bray 1900; Du Rietz 1940; Johnston 1940; Constance 1963; Raven and Axelrod
1974; Hunziker et al. 1972; Solbrig 1972; Werger 1973; Carlquist 1983).
Raven (1963) distinguished three classes of antitropical disjunctions: (a) the
bipolar pattern (e.g. Empetrum, Euphrasia, see Du Rietz 1940); (b) the pattern
encompassing temperate taxa disjunct between North and South America (e.g.
Collomia, Osmorhiza, Phacelia); (c) the pattern showed by desert plants (e.g.
Errazurizia, Larrea, Tiquilia).
Examples given above are at the genus level, but there are also disjunc-
tions between identical or closely related species such as Hoffmannseggia glauca
(Simpson et al. 2005), or Tiquilia nuttallii (Moore et al. 2006).
Werger (1973) revised the hypotheses that could explain these disjunctions:
(a) long-distance dispersal; (b) stepping-stone migration across arid or semi-arid
“islands” across the Andes; (c) land-bridges, according to van Steenis (1962)
(Chap. 10), including wider coastal areas to the west of the continent; (d) polytopic
origin of closely related species due to a widespread ancestor, i.e. parallel evolution
of near-identical arid-adapted taxa from widely distributed tropical ancestors (cfr
Johnston 1940; Barbour 1969).
Werger’s opinion was, taking account of the taxa analysed, that the most plausible
hypotheses were the polytopic speciation and the tropical origin of the ancestors of
extant taxa, followed by vicariant speciation (Werger 1973, p 16). These conclusions
were contrary to the dominant view, since from the 1960s on, most researchers
favoured the explanation of long-distance dispersal by birds (Raven 1963; Cruden
1966; Carlquist 1983; 1972; Simpson and Neff 1985). “Small seeds occasionally
adhere to birds and exceptionally may not fall off until the bird has reached a
favourable habitat on the other side of the tropics. Considering the millions of birds
that fly between temperate North and South America every year, some transport
might happen at least at the rate postulated for the colonization of Hawaii which lies
on no known migration route” (Raven 1963, p 153).
3.3 In the Search for Centres of Origin: Dispersal v/s Vicariance

in the Chilean Flora
Linné believed that all species dispersed from Mount Ararat after leaving Noah’s
arch (Llorente Bousquets et al. 2003). Biogeography surpassed the myth but could
not leave the eternal search for centres of origin, a paradigm that gained its
strength in Darwin’s The Origin of Species, and was further developed by the “New
York School of biogeography” by renowned researchers such as WD Matthew,
GG Simpson, PJ Darlington Jr and GS Myers (Nelson and Ladiges 2001).
Dispersalist biogeography is based on the assumption that taxa originated in rel-
atively small areas (centres of origin) and therefore tries to reconstruct the routes
that organisms covered to colonize known past or present ranges. On the other
hand, many authors saw problems in the search for the centres of origin and migra-
tion routes, like botanist Stanley Cain, who asserted that “. . . [centres of origin]
have been largely accepted without question, despite the lack of substantiating data
in some cases, and have been variously and somewhat loosely employed” (Cain
1944, p 185). Croizat called ironically dispersalism “the science of the curious, the
mysterious, the improbable” (Croizat 1958).
Only from the 1980s onwards, disjunct distribution patterns on the continent and
between continents began to be interpreted as the result of vicariance (e.g. Landrum
1981). This was contemporary to the appearance of a new biogeographic paradigm
of vicariance (cladistic) biogeography (Nelson and Platnick 1981; Humphries and
Parenti 1999) (Box 3.2).
3.3 In the Search for Centres of Origin: Dispersal v/s Vicariance 115
Box 3.2 Vicariance Biogeography

Gareth Nelson and Norman Platnick developed this approach based on the
work of León Croizat (Nelson and Platnick 1981), but vicariance as an expla-
nation of disjunct distributions is already found in the early works of von
Ihering (1893), in F. Ratzel’s biogeography (Ratzel 1901) and in writings
from J.D. Hooker (Turrill 1953). In vicariance biogeography distributions
of monophyletic groups of taxa over areas are explained by the reconstruc-
tion of area cladograms (Fig. 3.20). These area cladograms are hypotheses
of historical relationships between areas and are derived from phylogenetic
and distributional information of the monophyletic groups concerned. A first-
order explanation for correspondence between phylogenetic relationships of
taxa and historical relationships among areas is vicariance (Humphries and
Parenti 1999). Hovenkamp (1997) even proposed that the object of historical
biogeography may be specifically the vicariance events and not necessarily the
areas. According to Wiley (1988), vicariance biogeography arose from three
events: (1) the emergence of plate tectonics as the major geological paradigm;
(2) the emergence of phylogenetic systematics; and (3) the panbiogeography
of Croizat that abandoned the centres of origin approach in favor of vicari-
ance explanations. Vicariance biogeography have been also called cladistic
biogeography, but Parenti and Ebach (2009) suggests that this last term shall
be restricted to “those methods that cpnvert taxon cladograms into areagrams
(area cladograms) and that infer biotic area homologs from such areagrams“
(Parenti and Ebach 2009, p 121).
Sp. 1 ABCD
A A
Sp. 2 CD
C B
Sp. 3 BCD
B C
Sp. 4 CD
Sp. 5 D D D
Fig. 3.20 Two possible resolved area cladograms (areagrams) for five hypothetical species
distributed in four areas (adapted from Parenti and Ebach 2009)
Nelson and Platnick’s approach was much extended during the 1980s by many
researchers, announcing a change in the biogeographic paradigm from the
Darwinian dispersalist biogeography (Zunino and Zullini 2003).
Vicariance biogeography reached more satisfactory explanations for

the disjunct pattern of many traditional studied groups like Nothofagus
(Humphries 1981) (Chap. 9). With major emphasis on phylogenetic rela-
tionships the approach is related to vicariance biogeography (with several
divergences) and is called phylogenetic biogeography, as developed by Lars
Brundin inspired by Willi Hennig’s principles (Williams and Ebach 2008)
(Box 10.3). “Taking a survey of the biogeographical literature of the last
decades we are faced with a rich flora of concepts like ‘means of disper-
sal’, ‘spread potential’, ‘jumps’, ‘island hopping’ ‘sweep-stakes’, and ‘filter
bridges’, all intermingled in peripheral discussions overlooking the ever
present central problem: which is the sister group of the discussed group,
and where is it living?” (Brundin 1966, p 5). Several Chilean endemic genera
have been investigated in relation to their closest relatives. Renner (2004) pro-
posed that Peumus (Monimiacea) diverged from the Monimia/Palmeria line
about 76 mya, by the disruption of a formerly continuous range that stretched
from Chile across Antarctica and the Kerguelen plateau to Madagascar (see
Sect. 1.2). Another charismatic genus is Gomortega: its fossils are only
known from the Late Oligocene-Early Miocene (Gomortegoxylon, Nishida
et al. 1989), but molecular analysis suggests an age of 100 million years
for Gomortega (Renner 2004) (Fig. 3.21 and Fig. 3.14). Whether the
Gomortegaceae was more numerous in the past and suffered from extremely
high extinction rates or just had a very slow speciation rate (just one species
existing for 100 mya!) is a matter of speculation.
Nemuaron (N Caledonia)
Atherosperma (Australia / Tasmania)
Laurelia sempervirens (Chile endemic)
Dryadodaphne (E Australia / N Guinea)
Daphnandra (E Australia)
Doryphora (E Australia)
Laureliopsis (Chile / Argentina)
Laurelia novae-zelandiae (NZ)
Gomortega (Chile endemic)
Fig. 3.21 Phylogenetic and geographic relationships of Gomortega (according to Renner

2004)
From the Philesiaceae, Vinnersten and Bremer (2001) suggested that the
ancestor of Ripogonum, Lapageria (Chilean endemic), and Philesia (South-
temperate) was distributed in South America and New Zealand and possibly
also in Australia. The isolation of South America from Australia and New
Zealand corresponds to a split off of the South American Lapageria and
Philesia from the Australian–New Zealand Ripogonum, estimated at ca
47 mya (Figs. 3.22, 3.11 and 3.14). The same interpretation is possible for
the split between Alstroemeriaceae and Luzuriagaceae, so it may be that ter-

mination of the Antarctic link during the Eocene resulted in two vicariance
events within the Liliales (Vinnersten and Bremer 2001).
Ripogonum (Australia/N. Guinea/NZ)

Lapageria (endemic Chile)
Philesia (Chile/Argentina)
Fig. 3.22 Phylogenetic and geographic relationships of the Philesiaceae (according to

Vinnersten and Bremer 2001)
Finally, the southern Andes endemic conifer Austrocedrus seems to be

related to fossils found in the Early Oligocene of Tasmania (Paull and Hill
2008). The plausible explanation is that the original range was disrupted by
one or more vicanriant events with Austrocedrus chilensis the sole survivor of
the lineage.
3.3.1 Revitalizing Long-Distance Dispersal

Vicariance explanations were not to last, since they remained dominant only until
the recent advent of molecular systematic techniques, particularly molecular-based
dating of lineage divergences (de Queiroz 2005). In Moore et al.’s (2006) opin-
ion “. . . using these techniques, much recent scholarship has demonstrated that
numerous plant disjunctions are far too young to have resulted from vicariance,
leaving transoceanic dispersal as the only plausible alternative. . . The realization
that long-distance dispersal may have been far more frequent than previously sup-
posed has led plant biogeographers using modern molecular tools to reexamine
the relative importance of vicariance and dispersal in explaining the classic pat-
terns of worldwide plant disjunction”. Moore et al. (2006) consequently found that
the disjunct distribution of extant species of Tiquilia is the result of at least four
long-distance dispersal events from North America to South America (Fig. 3.23a).
Many more examples of long-distance dispersal are arising nowadays in systematic
papers: applying the same approach, Simpson et al. proposed four long-distance dis-
persal events in Hoffmannseggia, colonizing North America from South America
(Fig. 3.23b). The proposed explanatory mechanism is bird dispersal at different
times in the late Cenozoic (late Miocene to Pleistocene). Also Meudt and Simpson
(2006) proposed that Ourisia arose in the Andes of central Chile, then spread to
4x
a b c
Fig. 3.23 a Distribution of Tiquilia species, originated by long-distance dispersal (according to

Moore et al. (2006)); b disjunct populations of Hoffmannseggia glauca, originated by a long-
distance dispersal event (according to Simpson et al. (2005)); c long distance dispersal events that
caused the disjunct distribution of Ourisia species (according to Meudt and Simpson (2006))
southern Chile and Argentina, then dispersed to the north-central Andes, and finally
to Tasmania and New Zealand (Fig. 3.23c).
Also Chung et al. (2005) proposed that the ancestor of Oreomyrrhis was dis-
tributed in Eurasia and during the late Cenozoic to the Quaternary dispersed to
North America and the southern Pacific Rim. For Osmorhiza species, Wen et al.
(2002) suggested, due to the lack of ITS sequence divergence, recent LDD events
via birds from western North America to South America.
Chacón et al. (2006) asserted that the presence of Oreobolus in the Hawaiian
archipelago can only be explained as a result of LDD from Australasia or Malesia.
The arrival of Oreobolus in South America was according to these authors a rela-
tively recent event (approximately 5.5 mya) and interpreted therefore as LDD from
Australia. The rise of South American species by vicariance after the Gondwanan
breakup is discarded, since it would have only been possible through the Antarctic
connection, thus much earlier.
Calviño et al. (2008) found that Eryngium and its two subgenera originated from
western Mediterranean ancestors and that the present-day distribution of the genus is
explained by several dispersal events, including a long trans-Pacific dispersal event
from Chile to Australia. Kadereit et al. (2008) further found that the Eryngium
ancestors (tribe Saniculeae) have their origin in southern Africa; from there, the
tribe reached western Eurasia; then Eryngium entered the New World, and Australia
was reached, at its earliest, between 2.6 and 2.2 mya. They described this dispersal
pattern as “from south to north and south again”.
Also Tremetsberger et al. (2005) proposed dispersal from Northwest Africa
across the Atlantic Ocean for the origin of the South American taxa of Hypochaeris
(Asteraceae). In the Brassicaceae, Bleeker et al. (2002) suggested that long-distance
dispersal via migrating birds explains the antitropical disjunction between South
American Rorippa philippiana and North American R. curvisiliqua. Also for
Lepidium Mummenhoff et al. (2001) proposed long-distance dispersal from west-
ern North America to South America by birds in the Pleistocene. And von Hagen
and Kadereit (2001) suggested that Gentianella (Gentianaceae) arrived via dis-
persal from South America into Australia/New Zealand less than 2.7 mya. For
Coprosma (Rubiaceae, Anthospermeae) Anderson et al. (2001) proposed Africa
as the ancestral area, followed by long-distance dispersal into the Pacific; for
Tetrachondra Wagstaff et al. (2000) suggested LDD between New Zealand – South
America; for Fagonia (Zygophyllaceae) Beier et al. (2004) proposed dispersal
between South America and southern Africa.
But in spite of new molecular tools, not every group has yet allowed researchers
to unravel its complicated biogeographic history, and several authors are more
cautious: “Given the inherent methodological problems, absolute age of clade diver-
gences, relevant as evidence of long distance dispersal or vicariance, cannot yet be
determined with confidence” (Ladiges et al. 2005) (Box 3.3).
Box 3.3 Rocks Around the Clock: The New Paradigm

of Molecular Dating
The program of molecular dating of phylogenies has becoming very popular

due to the availability of better resolved phylogenies in combination with new
methods for estimating divergence times (e.g. Sanderson 1997; Rutschmann
2006; Parker and Markwith 2007). There is also an increasing number of tax-
onomically identifiable fossils suited for calibration purposes (Gandolfo et al.
2008). In spite of the fact that the datings have given widely different results,
more recent studies tend to converge on similar ages (see Box 2.4).
But in spite of Donoghue and Benton’s (2007) assertion that “rocks and
clocks together are an unbeatable combination”, the molecular dating program
is not free of problems (Pulquério and Nichols 2007; Heads 2005). Anderson
et al. (2005), after discussing the difficulties and uncertainties in obtaining
a stem group age for the eudicots, finished their paper asking if the results
could not be rather an artifact from constraints or method. Variation in rates
of nucleotide substitution along a lineage and between different lineages has
demonstrated to be pervasive, therefore the “clock” model of molecular evo-
lution has been changed to the relaxed or “sloppy clock”, that tries to address
the deviations from the clock-like model (Drummond et al. 2006).
Sanderson and Doyle (2001) pointed out that “non-clocklike behaviour of
evolutionary rates might lead to significant deviations among results obtained
with different dating methods”. Different methods may introduce systematic
biases, which are generally hard to detect. Near and Sanderson (2004) sug-
gested that the accurate divergence time estimates might require multiple
reliable calibrations, but “Ironically, the systems in which divergence time
estimation from sequence data is needed most critically are the ones with few
or no good calibrations. . . Perhaps we should learn to walk in the context of
these systems before learning to run in the real world” (Near and Sanderson
2004). Returning to examples mentioned before, many works do not mention
the calibration point used for the analysis (e.g. Simpson et al. 2005; Meudt
and Simpson 2006).
The mechanisms responsible for long-distance dispersal were experimentally

analysed already by Darwin (1859). Ridley (1930) and Carlquist (1983), among oth-
ers, emphasized the huge possibilities of dispersal in plants related to the immense
diversity of seed types. More recently, Muñoz et al. (2004) modelled wind connec-
tivity that could explain floristic similarities in the southern hemisphere, especially
for groups known as “good dispersers” (=lichens, mosses, ferns). The authors sug-
gested that this could apply also to angiosperms with tiny seeds, like the orchids,
but no empirical observation supports this hypothesis. On the contrary, Van Steenis
(1962) rejected long ago this possibility: “Although dispersal of orchids may seem
easy by the large amount of dust seed, successful establishment may depend on
presence of its mycorrhizal fungus and insects for pollination. That the three of
them, fungus spores, seeds, and insects, will travel together over long distances by
chance is utterly unlikely”. Other genera that superficially appear as good dispersers,
like the representatives of the Asteraceae, are not necessarily good dispersers (see
Chap. 8).
3.3.2 Relativising Long-Distance Dispersal

Cain already warned us about the error in convenient dispersal stories: “Long-
distance dispersal operates for some organisms, and it is especially applicable
to littoral species and a portion of the biota of oceanic islands. The hypothesis,
however, is much too widely used; in most cases of wide disjunction, a care-
ful investigation shows that the dispersal mechanisms, agents, and establishment
requirements of the species rule out this explanation. All too frequently the assump-
tion of long-distance dispersal is merely a careless and easy way out of a difficult
problem and it leads to fanciful and even ridiculous conclusions” (Cain 1944,
pp 305–306).
But researchers tend to ignore or minimize empirical evidence: “Hoffmannseggia
fruits and seeds have no obvious adaptations for external animal dispersal and no
one has ever recorded their being eaten by birds. Nevertheless, we believe that bird
dispersal is the most likely explanation for the repeated pattern of long-distance
dispersal from South to North America” (Simpson et al. 2005). The same applies for
Prosopis: “Although there is no record of Prosopis pods being eaten by birds. . . bird
dispersal appears to be the most likely explanation for the [antitropical] relationships
we have observed” (Bessega et al. 2006).
Taking the dispersalist universe as a framework, Moore et al. (2006) recently
avoided mentioning alternative palaeogeographic hypothesis, as listed by Constance
(1963), and they ruled out the possibility of ancient vicariance events. These alter-
native hypotheses are categorized as “grotesque hypotheses that have been proposed
to avoid the bugaboo of long-distance dispersal” (Raven 1963, p 153). But whether
or not grotesque is better than bugaboo is just a matter of belief and not a matter of
evidence.
Early biogeographers were much more skeptic regarding the real possibility of
long-distance dispersal: “If migratory birds really have a high importance in the
dispersal of plants, the common floristic pattern would resemble continuous faecal
roads, and could be reconstructed as the prehistoric trading routes. But this is not
the case. . .” (von Ihering 1893).
Also the eminent geographer Friedrich Ratzel (1901) wrote down his thoughts
about the subject:
“If one inspects the real distribution of organisms, one cannot confirm the effec-
tivity of all the passive means of migration, which gained so much attention since
Darwin and Wallace”. By the way, he wrote also: “Since lands and oceans have
moved uninterrupted and are shifting nowadays under our eyes on the Earth’s sur-
face, the ‘Lebensraum’ for the terrestrial and aquatic organisms likewise changes
continuously”. Ratzel’s opinion is not just anecdotic for biogeographers, since aside
from his reputation as a human geographer, he is considered as the founder of the
term “biogeography” (Parenti and Ebach 2009, see also Sect. 10.6).
Aditionally, Du Rietz wrote: “My own experience of species differentiation on
isolated mountains or mountain chains does not make me inclined to believe that
long distance dispersal has played any greater role there than in oceanic islands [. . .]
Our present knowledge of bipolar plant distribution and of plant distribution in gen-
eral does not speak in favour of the assumption that the facts of bipolar distribution
could be explained by long distance dispersal” (Du Rietz 1940, pp 239–240). His
own experience was greatly influenced by Carl Skottsberg’s impressions on oceanic
islands such as Juan Fernández (see Chap. 5).
More recently, some authors are appealing to more complex solutions of
vicariance and long-distance dispersal events working together: for Raukaua
(=Pseudopanax), Mitchell and Wagstaff (2000) proposed a Gondwanan vicari-
ance between Australasia-South America and long-distance dispersal to Hawaii; for
Laurelia (Atherospermataceae), Renner et al. (2000) proposed an initial diversifi-
cation at 100–140 mya, probably in West Gondwana, early entry into Antarctica,
and long-distance dispersal to New Zealand and New Caledonia; Wanntorp and
Wanntorp (2003) proposed early vicariance followed by recent dispersals in
Gunnera. For Caltha (Ranunculaceae), Schuettpelz and Hoot (2004) proposed a
Northern hemisphere origin, followed by dispersal to the Southern Hemisphere
(Gondwanaland). For the southern family Proteaceae, Barker et al. (2007) found
that four of eight trans-continental disjunctions of sister groups fit vicariance scenar-
ios, and the other half can be only explained by long-distance dispersal. Calibration
rests on a few fossils from the subfamily Grevilleoideae. Results provide an age of
118.5 mya for the crown group of the Proteaceae, in agreement with results provided
by Anderson et al. (2005). In Barker et al.’s (2007) analysis, all Chilean genera
(Embothrium, Gevuina, Lomatia, Orites) appear to be congruent with a vicariant
history.
Finally, several authors (e.g. Sluys 1994) argued that the contrast of vicariance
versus dispersal is an artifice of poorly defined concepts. Williams and Ebach (2008,
p 240) called this “another false war”. Michaux (2001) suggested instead of this
simple contrast, the recognition of five processes that are not logically equivalent as
they operate at different time scales: modification, movement, mixing, splitting and
juxtaposition.
Modern analytical tools like DIVA-analysis are designed to find the optimal dis-
tributions of ancestral species by minimizing the number of dispersal and extinction
events (Ronquist 1997; Sanmartín and Ronquist 2004). Applying this approach,
Sanmartín et al. (2007) tested the directional dispersal in the Southern Hemisphere.
The direction of circumpolar currents predicts predominantly eastward dispersal
from New Zealand to South America, but contrary to the expectations, dispersal
between New Zealand and South America was more frequently inferred to be west-
ward. The authors concluded that there is still a need of a better understanding
of dispersal processes for applying realistic estimates of biogeographical recon-
structions (Sanmartín et al. 2007). This better understanding of dispersal processes
should not exclude a need for a better understanding of vicariance processes. These
last have been frequently oversimplified as a break-up model of several steps, obvi-
ating the complex geological and biotic nature of any region composed of geological
and biogeographical composite areas (e.g. Heads 1999, 2002, see discussion in
Sect. 10.2).
. . . the apparent conflict between those who advocate dispersal and those who advocate
vicariance is of little consequence for progress in biogeography. To understand biogeog-
raphy as a battle for unification between vicariance and dispersal processes is, in short,
meaningless. The real task in biogeography is the unification between classification and
explanation – a unification that has obsessed biologists since Haeckel and his discussion of
chorological processes (Williams and Ebach 2008, p 241).
References
Abraham E, del Valle HF, Roig F, Torres L, Ares JO, Coronato F, Godagnone R (2009) Overview
of the geography of the Monte Desert biome (Argentina). J Arid Environ, 73:144–153
Anderson CL, Rova JHE, Andersson L (2001) Molecular phylogeny of the tribe Anthospermeae
(Rubiaceae): systematic and biogeographic implications. Aust Syst Bot, 14:231–244
Anderson CL, Bremer K, Friis EM (2005) Dating phylogenetically basal eudicots using rbcL
sequences and multiple fossil reference points. Am J Bot, 92:1737–1748
Arroyo MTK, Squeo FA, Armesto JJ, Villagrán C (1988) Effects of aridity on plant diversity in the
northern Chilean Andes: results of a natural experiment. Ann Mo Bot Gard, 75:55–78
Barbour M (1969) Patterns of genetic similarity between Larrea divaricata of North and South
America. Am Midl Nat, 81:54–67
Barker NP, Weston PH, Rutschmann F, Sauquet H (2007) Molecular dating of the “Gondwanan”
plant familt Proteaceae is only partially congruent with the timing of the break-up of
Gondwana. J Biogeogr, 34:2012–2027
Barlow BA (1981) The Australian flora: its origin and evolution. In: Bureau of Flora and Fauna
(ed) Flora of Australia, vol. 1. AGPS, Canberra, pp. 25–75
Beier BA, Nylander JAA, Chase MW, Thulin M (2004) Phylogenetic relationships and biogeogra-
phy of the desert plant genus Fagonia (Zygophyllaceae), inferred by parsimony and Bayesian
model averaging. Mol Phylogenet Evol, 33:91–108
Bessega C, Vilardi JC, Saidman BO (2006) Genetic relationships among American species of
the genus Prosopis (Mimosoideae, Leguminosae) inferred from ITS sequences: evidence for
long-distance dispersal. J Biogeogr, 33:1905–1915
Bleeker W, Weber-Sparenberg C, Hurka H (2002) Chloroplast DNA variation and biogeography
in the genus Rorippa Scop. (Brassicaceae). Plant Biol, 4:104–111
References 123
Bray WL (1900) The relations of the North American flora to that of South America. Science,
12:709–716
Brundin L (1966) Transantartic relationships and their significance, as evidenced by chironomid
midges. Kungl. Svenska Vetenskapsakademiens Handlingar, 11:1–472
Cain SA (1944) Foundations of plant geography. Harper and Brothers, New York, NY
Calviño CI, Martinez SG, Downie SR (2008) The evolutionary history of Eryngium (Apiaceae,
Saniculoideae): Rapid radiations, long distance dispersals, and hybridizations. Mol Phylogenet
Evol, 46:1129–1150
Carlquist S (1983) Intercontinental dispersal. In: Kubitzki K, (ed) Dispersal and distribution:
an international symposium. Sonderbände des Naturwissenschaftlichen Vereins in Hamburg,
vol 7. Verlag Paul Parey, Hamburg/Berlin, 37–47
Chacón J, Madriñán S, Chase MW, Bruhl JJ (2006) Molecular phylogenetics of Oreobolus
(Cyperaceae) and the origin and diversification of the American species. Taxon, 55:359–36
Christ H (1867) Über die Verbreitung der Pflanzen der alpinen Region der europäischen
Alpenkette. Denkschr. Schweiz. Nat. Ges. 22
Chung KF, Peng CI, Downie SR, Spalik K, Schaal BA (2005) Molecular systematics of the
trans-Pacific alpine genus Oreomyrrhis (Apiaceae): phylogenetic affinities and biogeographic
implications. Am J Bot, 92:2054–2071
Constance L (1963) Amphitropical relationships in the herbaceous flora of the Pacific Coast of
North and South America: a symposium. Introduction and historical review. Q Rev Biol,
38:109–116
Cox CB (1990) New geological theories and old biogeographical problems. J Biogeogr 17(11):
7–130
Craw RC, Grehan JR, Heads MJ (1999) Panbiogeography: tracking the history of life. Oxford
Biogeography Series 11. Oxford University Press, New York, NY
Crisp MD, West JG, Linder HP (1999) Biogeography of the terrestrial flora. In: Orchard AE,
(ed) Flora of Australia, vol. 1, 2nd edn. Australian Biological Resources Study, Canberra,
pp 321–367
Croizat L (1952) Manual of Phytogeography, or an account of plant dispersal throughout the world.
Junk, The Hague
Croizat L (1958) Panbiogeography or an introductory synthesis of zoogeography, phytogeography,
and geology; with notes on evolution, systematics, ecology, anthropology, etc., 3 vol. Published
by the author, Caracas, Venezuela
Croizat L (1962) On the age of Fray Jorge and Talinay in Chile. Rev Univ (Chile), 47:57–61
Cruden RW (1966) Birds as agents of long-distance dispersal for disjunct plant groups of the
temperate Western Hemisphere. Evolution, 20:517–532
Darwin C (1859) The origin of species. Oxford World’s Classics (1996). Oxford University Press,
Cambridge
De Martonne E (1934) The Andes of the North-West Argentine. Geogr J, 84:1–14
de Queiroz A (2005) The resurrection of oceanic dispersal in historical biogeography. Trends Ecol
Evol, 20:68–73
Donato M, Posadas P, Miranda-Esquivel DR, Jaureguizar EO, Cladera G (2003) Historical bio-
geography of the Andean region: evidence from Listroderina (Coleoptera: Curculionidae:
Rhytirrhinini) in the context of the South American geobiotic scenario. Biol J Linn Soc,
80:339–352
Donoghue PCJ, Benton MJ (2007) Rocks and clocks: calibrating the Tree of Life using fossils and
molecules. Trends Ecol Evol, 22:424–431
Drummond AJ, Ho SYW, Phillips MJ, Rambaut A (2006) Relaxed phylogenetics and dating with
confidence. PLoS Biol, 4:e88
Du Rietz GE (1940) Problems of bipolar plant distribution. Acta Phytogeogr Suecica, 13:215–282
Engler A (1882) Versuch einer Entwicklungsgeschichte der Pflanzenwelt, insbesondere der
Florengebiete seit der Tertiärperiode, Vol. 2. Die Extratropischen Gebiete der Südlichen
Hemisphäre und die Tropischen Gebiete. Verlag von W. Engelmann, Leipzig
Gandolfo MA, Nixon KC, Crepet WL (2008) Selection of fossils for calibration of molecular
dating models. Ann Mo Bot Gard, 95:34–42
Glasby CJ (2005) Polychaete distribution patterns revisited: an historical explanation. Mar Ecol,
26:235–245
Good R (1947) The geography of the flowering plants. 4th edition 1974. Longman, London
Hannig E, Winkler H, (eds) (1926–1940) Die Pflanzenareale: Sammlung kartographischer
Darstellungen von Verbreitungsbezirken der lebenden und fossilen Pflanzen-Familien,
-Gattungen und–Arten. Gustav Fischer Verlag, Jena.
Heads M (1999) Vicariance biogeography and terrane tectonics in the South Pacific: analysis of
the genus Abrotanella (Compositae). Biol J Linn Soc, 67:391–432
Heads M (2002) Birds of paradise, vicariance biogeography and terrane tectonics in New Guinea.
J Biogeogr, 29:261–283
Heads M (2004) What is a node? J Biogeogr, 31:1883–1891
Heads M (2005) Dating nodes on molecular phylogenies: a critique of molecular biogeography.
Cladistics, 21:62–78
Heusser CJ (1971) Pollen and Spores of Chile. The University of Arizona Press, Tucson, Arizona
Hinojosa LF (2005) Cambios climáticos y vegetacionales inferidos a partir de paleofloras
Cenozoicas del sur de Sudamérica. Rev Geol Chil, 32:95–115
Hofsten N (1916) Zur älteren Geschichte des Diskontinuitätsproblems in der Biogeographie.
Zoologische Annalen, 7:197–353
Hooghiemstra H, van der Hammen T (1998) Neogene and Quaternary development of the
neotropical rain forest: the forest refugia hypothesis, and a literature overview. Earth Sci Rev,
44:147–183
Houston J, Hartley AJ (2003) The central Andean west-slope rainshadow and its potential
contribution to the origin of hyper-aridity in the Atacama Desert. Int J Climatol, 23:1453–1464
Hovenkamp P (1997) Vicariance events, not areas, should be used in biogeographical analysis.
Cladistics, 13:67–79
Hughes C, Eastwood R (2006) Island radiation on a continental scale: exceptional rates of plant
diversification after uplift of the Andes. Proc Natl Acad Sci Usa, 103:10334–10339
Humphries CJ (1981) Biogeographic methods and the southern beeches. In: Forey PL, (ed) The
evolving biosphere. Cambridge University Press, Cambridge, pp 283–297
Humphries CJ, Parenti LR (1999) Cladistic biogeography, 2nd edn. Oxford monographs on
biogeography N◦ 2. Clarendon Press, Oxford.
Hunziker JH, Palacios RA, de Valesi AG, Poggio L (1972) Species disjunctions in Larrea: evidence
from morphology, cytogenetics, phenolic compounds, and seed albumins. Ann Mo Bot Gard,
59:224–233
Johnston IM (1940) The floristic significance of shrubs common to North and South American
deserts. J Arnold Arbor, 21:356–363
Kadereit JW, Repplinger M, Schmalz N, Uhink CH, Worz A (2008) The phylogeny and biogeog-
raphy of Apiaceae subf. Saniculoideae tribe Saniculeae: from south to north and south again.
Taxon, 57:365–382
Katinas L, Morrone JJ, Crisci JV (1999) Track analysis reveals the composite nature of the Andean
biota. Aust J Bot, 47:111–130
Katinas L, Crisci JV, Wagner W, Hoch PC (2004) Geographical diversification of tribes Epilobieae,
Gongylocarpeae, and Onagreae (Onagraceae) in North America, based on parsimony analysis
of endemicity and track compatibility analysis. Ann Mo Bot Gard, 91:159–185
Ladiges PY, Kellermann J, Nelson G, Humphries CJ, Udovicic F (2005) Historical biogeography
of Australian Rhamnaceae, tribe Pomaderreae. J Biogeogr, 32:1909–1919
Landrum L (1981) The phylogeny and geography of Myrceugenia (Myrtaceae). Brittonia, 33:
105–129
Llorente Bousquets J, Papavero N, Bueno A (2003) Síntesis histórica de la biogeografía. In:
Llorente Bousquets J, Morrone JJ, (eds) Introducción a la Biogeografía en Latinoamérica:
Teorías, Conceptos, Métodos y Aplicaciones. Las Prensas de Ciencias, Facultad de Ciencias,
UNAM, México, pp 1–14
References 125
Luna Vega I, Alcántara Ayala O, Morrone JJ, Espinosa Organista D (2000) Track analysis and
conservation priorities in the cloud forests of Hidalgo (Mexico). Divers Distrib, 6:137–143
López RP (2003) Phytogeographical relations of the Andean dry valleys of Bolivia. J Biogeogr,
30:1659–1668
Mabberley DJ (1997) The plant-book, 2nd ed. (3rd edn 2008). Cambridge University Press,
Cambridge.
Messerli B, Ammann C, Geyh M, Grosjean M, Jenny B, Kammer K, Vuille M (1998) The problem
of the “Andean Dry Diagonal”: Current precipitation, late Pleistocene snow line, and lake level
changes in the Atacama Altiplano (18ºS–28º/29ºS). Bamberger Geogr Schr, 15:17–34
Meudt HM, Simpson BB (2006) The biogeography of the austral, subalpine genus Ourisia
(Plantaginaceae) based on molecular phylogenetic evidence South American origin and
dispersal to New Zealand and Tasmania. Biol J Linn Soc, 87:479–513
Meusel H (1943) Vergleichende Arealkunde. Einführung in die Lehre von der Verbreitung der
Gewächse mit besonderer Berücksichtigung der mitteleuropäischen Flora, 2 vols. Verlag
Borntraeger, Berlin-Zehlendorf.
Meusel H, Jäger E, (eds) (1992) Vergleichende Chorologie der Zentraleuropäischen Flora, vol. 3.
Gustav Fischer Verlag, Jena
Meusel H, Jäger E, Rauschert S, Weinert E, (eds) (1978) Vergleichende Chorologie der
Zentraleuropäischen Flora, vol. 2. Gustav Fischer Verlag, Jena
Michaux B (2001) Dispersal versus vicariance, artifice rather than contest. In: Metcalfe I, Smith
JMB, et al.(eds) Faunal and floral migrations and evolution in SE Asia-Australasia. AA
Balkema Publishers, Lisse, pp 311–318
Mihoc MAK, Morrone JJ, Negritto M, Cavieres LA (2006) Evolución de la serie Microphyllae
(Adesmia, Fabaceae) en la Cordillera de los Andes: una perspectiva biogeográfica. Rev Chil
Hist Nat, 79:389–404
Mitchell AD, Wagstaff SJ (2000) Phylogeny and biogeography of the Chilean Pseudopanax
laetevirens. New Zeal J Bot, 38:409–414
Moreira-Muñoz A (2007) Plant geography of Chile: an essay on postmodern biogeography.
Doctoral thesis, University Erlangen-Nürnberg, Germany
Moore MJ, Tye A, Jansen RK (2006) Patterns of long-distance dispersal in Tiquilia subg. Tiquilia
(Boraginaceae): implications for the origins of amphitropical disjuncts and Galapagos Islands
endemics. Am J Bot, 93:1163–1177
Mummenhoff K, Bruggemann H, Bowman JL (2001) Chloroplast DNA phylogeny and biogeog-
raphy of Lepidium (Brassicaceae). Am J Bot, 88:2051–2063
Muñoz J, Felicísimo AM, Cabezas F, Burgaz AR, Martínez I (2004) Wind as a long-distance
dispersal vehicle in the Southern Hemisphere. Science, 304:1144–1147
Muñoz-Schick M, Moreira-Muñoz A, León-Lobos P (2006) Nueva localidad en Chile para
Menodora linoides Phil. (Oleaceae), especie considerada extinta en el siglo XX. Gayana Bot,
63:175–179
Muñoz Pizarro C, Pisano E (1947) Estudio de la vegetación y flora de los Parques Nacionales Fray
Jorge y Talinay. Agric Téc (Chile), 7:71–190
Mutke J, Barthlott W (2005) Patterns of vascular plant diversity at continental to global scales.
Biol Skr, 55:521–531
Near TJ, Sanderson MJ (2004) Assessing the quality of molecular divergence time estimates
by fossil calibrations and fossil-based model selection. Philos Trans R Soc B Biol Sci, 359:
1477–1483
Nelson EC (1981) Phytogeography of southern Australia. In: Keast A, (ed) Ecological biogeogra-
phy of Australia. Dr W. Junk, The Hague, pp. 733–759
Nelson G, Platnick N (1981) Systematics and biogeography: Cladistics and vicariance. Columbia
University Press, New York, NY
Nelson G, Ladiges PY (2001) Gondwana, vicariance biogeography and the New York School
revisited. Aust J Bot, 49:389–409
Nishida M, Nishida H, Ohsawa T (1989) Comparison of the petrified woods from the Cretaceous
and Tertiary of Antarctica and Patagonia. Proc NIPR Symp Polar Biol, 2:198–212
Núñez-Avila MC, Armesto JJ (2006) Relict islands of the temperate rainforest tree Aextoxicon
punctatum (Aextoxicaceae) in semi-arid Chile: genetic diversity and biogeographic history.
Aust J Bot, 54:733–743
Parenti LR (2007) Common cause and historical biogeography. In: Ebach MC, Tangney RS, (eds)
Biogeography in a changing world. Systematics Association Special Volumes. CRC Press,
Boca Raton, FL, pp 61–82
Parenti LR, Ebach MC (2009) Comparative biogeography: discovering and classifying bio-
geographical patterns of a dynamic earth. University of California Press, San Francisco,
CA
Parker K, Markwith S (2007) Expanding biogeographic horizons with genetic approaches. Geogr
Compass, 1:246–274
Paull R, Hill RS (2008) Oligocene Austrocedrus from Tasmania (Australia): comparisons with
Austrocedrus chilensis. Int J Plant Sci, 169:315–330
Philippi F (1884) A visit to the northernmost forest of Chile. J Bot, 22:202–211
Pulquério MJF, Nichols RA (2007) Dates from the molecular clock: how wrong can we be? Trends
Ecol Evol, 22:180–184
Qian H (1999) Floristic analysis of vascular plant genera of North America north of Mexico:
characteristics of phytogeography. J Biogeogr, 26:1307–1321
Qian H, Song J-S, Krestov P, Guo Q, Wu Z, Shen X, Guo X (2003) Large-scale phytogeographical
patterns in East Asia in relation to latitudinal and climatic gradients. J Biogeogr, 30:129–141
Ratzel F (1901) Der Lebensraum: eine biogeographische Studie. Laupp’sche Buchhandlung,
Tübingen
Raven PH (1963) Amphitropical relationships in the floras of North and South America. Q Rev
Biol, 38:151–177
Raven PH (1972) Plant species disjunctions: a summary. Ann Mo Bot Gard, 59:234–246
Raven PH, Axelrod DI (1974) Angiosperm biogeography and past continental movements. Ann
Mo Bot Gard, 61:539–673
Reiche C (1905) Monotypische Gattungen der Chilenischen Flora. Verh Dtsch Wiss Vereins Stgo,
5:1–16
Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, Vol. VIII. Verlag von
Renner SS (2004) Variation in diversity among Laurales, early cretaceous to present. Biol Skr,
55:441–458
Renner SS, Murray D, Foreman D (2000) Timing transantarctic disjunctions in the
Atherospermataceae (Laurales): evidence from coding and noncoding chloroplast sequences.
Syst Biol, 49:579–591
Riccardi AC (1988) The Cretaceous system of southern South America. Geol Soc Am Memoir,
168:1–145
Ridley HN (1930) The dispersal of plants throughout the world. Lovell Reeve & Co, London
Ronquist F (1997) Dispersal-vicariance analysis: a new approach to the quantification of historical
biogeography. Syst Biol, 46:193–201
Rothmaler W (1955) Allgemeine Taxonomie und Chorologie der Pflanzen. Grundzüge der
speziellen Botanik. Libri Botanici vol. 3, IHW Verlang. Reprint 1992
Rutschmann F (2006) Molecular dating of phylogenetic trees: a brief review of current methods
that estimate divergence times. Divers Distrib, 12:35–48
Sanderson MJ (1997) A nonparametric approach to estimating divergence times in the absence of
rate constancy. Mol Biol Evol, 14:1218–1231
Sanderson MJ, Doyle JA (2001) Sources of error and confidence intervals in estimating the age of
angiosperms from rbcL and 18S rDNA data. Am J Bot, 88:1499–1516
Sanmartín I, Ronquist F (2004) Southern Hemisphere biogeography inferred by event-based
models: plant versus animal patterns. Syst Biol, 53:216–243
References 127
Sanmartín I, Wanntorp L, Winkworth RC (2007) West Wind Drift revisited: testing for direc-
tional dispersal in the Southern Hemisphere using event-based tree fitting. J Biogeogr, 34:
398–416
Scherson RA, Vidal R, Sanderson MJ (2008) Phylogeny, biogeography, and rates of diversification
of New World Astragalus (Leguminosae) with an emphasis on South American radiations. Am
J Bot, 95:1030–1039
Schuettpelz E, Hoot SB (2004) Phylogeny and biogeography of Caltha (Ranunculaceae) based on
chloroplast and nuclear DNA sequences. Am J Bot, 91:247–253
Simpson BB, Neff JL (1985) Plants, their pollinating bees, and the great American interchange.
In: Stehli FG, Webb SD, (eds) The great American biotic interchange. Plenum, New York, NY,
pp 427–452
Simpson BB, Tate JA, Weeks A (2005) The biogeography of Hoffmannseggia (Leguminosae,
Caesalpinioideae, Caesalpinieae): a tale of many travels. J Biogeogr, 32:15–27
Sklenar P, Balslev H (2007) Geographic flora elements in the Ecuadorian superpáramo. Flora,
202:50–61
Skottsberg C (1948) Apuntes sobre la flora y vegetación de Frai Jorge (Coquimbo, Chile). Acta
Horti Gotoburgensis, 18:91–184
Sluys R (1994) Explanations for biogeographic tracks across the Pacific Ocean: a challenge for
paleogeography and historical biogeography. Prog Phys Geog, 18:42–58
Solbrig OT (1972) New approaches to the study of disjunctions with special emphasis on the
American amphitropical desert disjunctions. In: Valentine DH, (ed) Taxonomy, phytogeogra-
phy and evolution. Academic Press, London, pp 85–100
Stuessy TF, Taylor C (1995) Evolución de la Flora Chilena. In: Marticorena C, Rodríguez R, (eds)
Flora de Chile, vol. 1. Universidad de Concepción, Concepción, pp 85–118
Stuessy TF, Tremetsberger K, Müllner AN, Jankowicz J, Guo Y-P, Baeza CM, Samuel RM (2003)
The melding of systematics and biogeography through investigations at the populational level:
examples from the genus Hypochaeris (Asteraceae). Basic Appl Ecol, 4:287–296
Takhtajan A (1986) Floristic regions of the world. Translated by Crovello TJ, edited by Cronquist
A from the original Russian edition, Leningrad (1978). University of California. Reprint 1988,
B. Singh, M. P. Singh, Dehra Dun, India
Tremetsberger K, Weiss-Schneeweiss H, Stuessy TF, Samuel R, Kadlec G, Ortiz MA, Talavera S
(2005) Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South
American Hypochaeris (Asteraceae, Cichorieae). Mol Phylogenet Evol, 35:102–116
Treviranus GR (1803) Biologie, oder Philosophie der lebenden Natur. Röwer, Göttingen
Troncoso A, Villagrán C, Muñoz-Schick M (1980) Una nueva hipótesis acerca del origen y edad
del bosque de Fray Jorge (Coquimbo, Chile). Bol Mus Nac Hist Nat (Chile), 37:117–152
Turrill WB (1953) Pioneer plant geography: the Phytogeographical Researches of Sir Joseph
Dalton Hooker. Contributions from the Int. Biohistorical Commission, N◦ 1, The Hague,
Netherlands
van Steenis CGGJ (1962) The theory of land bridges in botany, with particular references to tropical
plants. Blumea, 11:235–542
Villagrán C, Hinojosa LF (1997) Historia de los bosques del sur de Sudamérica, II: Análisis
fitogeográfico. Rev Chil Hist Nat, 70:241–267
Villagrán C, Armesto JJ, Hinojosa LF, Cuvertino J, Pérez C, Medina C (2004) El enigmático ori-
gen del bosque relicto de Fray Jorge. In: Squeo FA, Gutiérrez JR, Hernández IR, (eds) Historia
Natural del Parque Nacional Fray Jorge. Ediciones Universidad de La Serena, La Serena,
pp 3–43
Vinnersten A, Bremer K (2001) Age and biogeography of major clades in Liliales. Am J Bot,
88:1695–1703
von Hagen KB, Kadereit JW (2001) The phylogeny of Gentianella (Gentianaceae) and its col-
onization of the southern hemisphere as revealed by nuclear and chloroplast DNA sequence
variation. Org Divers Evol, 1:61–79
von Ihering H (1893) Das neotropische Florengebiet und seine Geschichte. Englers Bot Jahrb,
17:1–54
Wagstaff SJ, Martinsson K, Swenson U (2000) Divergence estimates of Tetrachondra hamiltonii
and T. patagonica (Tetrachondraceae) and their implications for austral biogeography. New
Zeal J Bot, 38:587–596
Wangerin W (1932) Florenelemente und Arealtypen. Beiträge zur Arealgeographie der deutschen
Flora. Beih. Z. Centralbl. (Drude Festschrift), vol 49
Wanntorp L, Wanntorp H-E (2003) The biogeography of Gunnera L.: Vicariance and dispersal.
J Biogeog, 30:979–987
Wen J, Ickert-Bond SM (2009) Evolution of the Madrean-Tethyan disjunctions and the North and
South American amphitropical disjunctions in plants. J Syst Evol, 47:331–348
Wen J, Lowry PP, Walck JL, Yoo K-O (2002) Phylogenetic and biogeographic diversification in
Osmorhiza (Apiaceae). Ann Mo Bot Gard, 89:414–428
Werger M (1973) Las disyunciones anfitrópicas en las floras xerofíticas norte y sudamericanas.
Darwiniana, 18:9–18
Wielgorskaya T (1995) Dictionary of generic names of seed plants. Columbia University Press,
New York, NY
Wiley EO (1988) Vicariance biogeography. Ann Rev Ecol Syst, 19:513–542
Williams DM (2007) Ernst Haeckel and Louis Agassiz: trees that bite and their geographical
dimension. In: Ebach MC, Tangney RS, (eds) Biogeography in a changing world. Systematics
Association Special Volumes. CRC Press, Boca Raton, FL, pp 1–59
Williams DM, Ebach MC (2008) Foundations of systematics and biogeography. Springer, New
York, NY
Wulff EV (1950) An introduction to historical plant geography. The Chronica Botanica Company,
Waltham, MA
Zunino M, Zullini A (2003) Biogeografía: la dimensión espacial de la evolución. Fondo de Cultura
Económica, México. From the 1995 Italian edition
Chapter 4
Biogeographic Regionalization
Abstract Biogeographic regionalization has been a main task from the beginnings
of biogeography with the early works of Gottfried Treviranus, August P. de
Candolle, or Frederick Schouw. Regionalization results in a hierarchy encompassing
several categories like the realm, the region, the provinces, and so on. Comparison
of available regionalization proposals for the Chilean territory is important but chal-
lenging, since each author puts the emphasis on a different level in this hierarchy. An
independent exercise of regionalization is undertaken by the analysis of the endemic
genera with the program NDM/VNDM. This yielded 3 areas of endemism located
in Central Chile.
Biogeographic regionalization is tied to the initials of biogeography, beginning with

the early works of G. Treviranus, A.P. de Candolle, and J.F. Schouw (Sect. 4.1).
Biogeographic regionalization rests upon the chorological tradition (e.g. Huxley
et al. 1998, Sect. 3.1), and it is still of crucial importance in modern biogeography
(Llorente Bousquets and Morrone 2005; McLaughlin 2007; Mackey et al. 2008;
Proches 2008; Escalante 2009).
Biogeograpic regionalization results in a hierarchical system – equivalent to a
taxonomic hierarchy – for categorizing geographic areas in terms of their biota
(Zunino and Zullini 2003; Escalante 2009). The existence of hierarchical systems
is a highly relevant topic in systematics and biogeography, although the nature of
such systems is still controversial. While several authors maintain that hierarchies
tend to be artificial human constructions (e.g. Grehan 2001), others maintain that
they are characteristic of nature’s organization (Eldredge 1985; McLaughlin 1992,
1994) (Fig. 4.1). Both visions seem to be complementary: it is a fact that hierar-
chies are essential for man’s understanding of the world: “The assessed patterns
of organismal relationship are used to construct hierarchical classifications of coor-
dinate and subordinate groups that are information-rich and have high predictive
efficacy; these are the taxonomic hypotheses that change with new information and
new modes of analysis” (Stuessy 2006). “The organization of nature is profoundly
hierarchical, because from its beginning, interactions between simple elements have
continuously created more complex systems, that themselves served as the basis for
still more complex systems” (Jagers op Akkerhuis 2008).

DOI 10.1007/978-90-481-8748-5_4, ⃝
130 4 Biogeographic Regionalization
Fig. 4.1 “Natural”

hierarchies; entities in the Taxonomic hierarchy Biogeographic hierarchy
hierarchies are not necessarily
equivalent Orders Realms
Families Regions
Tribes Dominions
Genera Provinces
Species Ditricts
A regionalization comprises from the general to the particular a hierarchy begin-

ning with the realm (=kingdom), the region, the dominion, and the province (e.g.
Engler 1882; Takhtajan 1986; Cabrera and Willink 1973) (Fig. 4.1). There could
be also intermediate categories, as subregions or subdominions. The basic unit is
the province, although several authors suggest the district (=Bezirk) as the finest
unit (e.g. Engler 1882). Each of the levels in the biogeographic hierarchy contains
the subsequent levels, on the base of the characteristics (taxa) they share; a realm
contains various regions, a region contains several dominions, and so on.
4.1 The Chilean Plants in the Global Concert

The Chilean flora appeared as early as the first attempts to geographically classify
the global flora. Hereafter a summary of the different ways in which the Chilean
flora has been classified is outlined (Table 4.1).
Gottfried R. Treviranus (1776–1837), one of the naturalists who coined the term
“biology” (Engels 2005), first intended a global floristic classification, organiz-
ing the world flora in 8 principal floras (Hauptfloren). In this early classification
Chile, (including Magallanes and Tierra del Fuego), and New Zealand composed
an Antarctic Flora (Antarktische Flor) (Treviranus 1803). Treviranus, based on the
early works of JI Molina (1740–1830), J Banks (1743–1820), and G Forster (1754–
1794), explicitly recognised the floristic relationship between southernmost South
America and Australasia, due to the similarities between New Zealand and Tierra
del Fuego. He also noticed the existence of an antitropical floristic element, such
as Pinguicula, Salix or Ribes (Sect. 3.1). He found this relation surprising since at
that time the floristic knowledge of Tierra del Fuego was limited to just 40 species
(Treviranus 1803, p 132).
4.1 The Chilean Plants in the Global Concert 131
Table 4.1 The geographical classification of the Chilean flora in the global context
Author N◦ realms N◦ regions Classification of the Chilean Flora
GR Treviranus (1803) 8 Flor – Antarktische Flor

August Pyramus de – 20 le Chili and les terres Magellaniques
Candolle (1820)
Schouw (1823) 25 – South of 42◦ = Antarktisches Florenreich;
42◦ –23◦ = Reich der Holzartigen
Synanthereen; North of 23◦ = Reich der
Cactus und Piper
Alphonse de Candolle – 45 Region 35, Le Chili, and region 36, la Patagonie,
(1835) la terre de Feu et les iles Malouines (Falkland)
Grisebach (1872) – 24 North to 23◦ = Tropischen Anden; 23◦ –34◦ =
Chilenisches Übergangsgebiet; 34◦ –56◦ =
Antarktisches Waldgebiet
Engler (1879–1882) 4 31 Südamerikanische Florenreich = to 41◦ ;
Altoceanisches Florenreich = south from 41◦
Drude (1884) 14 55 South of 41◦ = Antarktisches Florenreich; North
of 41◦ = Andines Florenreich
Drude (1890) 14 55 Andines Florenreich and Antarktisches
Florenreich
Diels (1908) 6 7 Antarktisches Florenreich and Neotropisches
Florenreich
Christ (1910) – 12 On the base of fern species, Chile was classified
onto an Andine Flora and a Süd Chilenische
Flora, with the limit at around 39 ◦ S
Good 1947 (1974) 6 37 Antarctic kingdom, Patagonian region, South of
41◦ ; Neotropical kingdom, Andean region,
North of 41◦
Mattick (1964) 6 43 South = Antarktisches Florenreich; North =
Neotropisches Florenreich
Takhtajan (1961) 6 37 Antarctic kingdom, Patagonian region, South of
40◦ ; Neotropical kingdom, Andean region,
North of 40◦
Takhtajan (1986) 6 35 Holantarctic kingdom, Chile-Patagonian region,
South of 25◦ to Antarctic peninsula and
Malvinas Islands (Falkland Is.)
Cox (2001) 5 – South American kingdom
Morrone (2002) 3 12 Austral kingdom, Andean region
Adapted from Moreira-Muñoz (2007).
At that time biogeographic representation by means of biogeographic maps

was just at the beginning, and in the hands of Jean B. Lamarck and Augustin-
Pyramus de Candolle. In 1805 they published the “first biogeographical map”
for the third edition of the Flore française (Ebach and Goujet 2006). A-P de
Candolle, in his Géographie botanique, further classified the world in 20 floris-
tic regions: Chile fitted into two regions, le Chili and les terres Magellaniques
(de Candolle 1820). De Candolle’s world classification still lacked a map. Three
Fig. 4.2 Scheme of

Schouw’s early classification
regarding South America: a
cinchona realm; b Cactus and
Piper realm; c palms and b
Melastomataceae realm; d a
arboreal Compositae realm; e
Antarctic realm (primarily
Nothofagus) c
years later, the Danish botanist Joakim Frederik Schouw (1789–1852) published
the first world map (Schouw 1823; Mennema 1985). Schouw proposed 25 floris-
tic realms (Florenreiche). Southern South America was classified into two realms:
the Reich der Holzartigen Synanthereen (Compositae), and the Antarktisches Reich,
from 40◦ S to the South (Fig. 4.2).
Applying the concept of “endemism” as first proposed by A-P de Candolle,
Schouw stated explicitly the criteria for classifying and delimiting the floristic
realms (Drude 1884, p. 13):
1. half of the known plants have to be native to the territory in question;

2. 1/4 of the genera have to be endemic or have their maximal distribution there;
3. a plant realm has to have some endemic families.
De Candolle’s son Alphonse de Candolle (1835) critized Schouw’s criteria and

proposed 45 botanical regions, but for Chile he maintained the division into two
regions. But soon de Candolle the younger rejected such schemes of regions and
turned to be the first critic of the task of floristic classification: “I thus hold divi-
sions of the sphere by areas, suggested until now, for artificial systems . . . . they just
harmed science” (A de Candolle 1855, pp. 1304–1305, as quoted by Nelson 1978).
Later he states, with reference to his Geographie Botanique: “my work remained
completely different from that what my father thought, because the documentation
had become more numerous, and my ideas had singularly moved away from those
which had reigned in science for twenty years” (A de Candolle Mémoires, p 395, as
quoted by Nelson 1978).
Is seems that as botanical information became overwhelming, the task of phy-
togeographical classification was getting more and more difficult. One of the most
important naturalists in this era of growing botanical knowledge was JD Hooker
(1817–1911) (Sect. 2.1). Joining James Cook on the Endeavour, he much improved
the floristic knowledge of the southern hemisphere. Hooker’s publications compiled
as The Botany of the Antarctic Voyage. . . (Hooker 1844–1860) were, accord-
ing to Thiselton-Dyer (1909), almost as epoch-making as Darwin’s Origin of
species.
By the second half of the nineteenth century a huge amount of floristic knowledge
had accumulated. This knowledge, coupled with the ecological principles devel-
oped since A. von Humboldt, permitted August Grisebach (1814–1879) to publish
his Vegetation der Erde, which related explicitly the plant world with the regional
climates (Grisebach 1872). In Grisebach’s view, Chile had to be classified into 3
regions (Gebiete): 1. a Chilean floristic core, Chilenisches Übergangsgebiet (tran-
sition zone) from 23◦ to 34◦ S, that holds a “unique flora”; 2. an Antarctic region
(Antarktisches Waldgebiet), ranging from 34◦ to 56◦ S, characterized especially by
the genus Nothofagus; and 3. a tropical Andean flora that ranges from Ecuador to
northern Chile (Table 4.1).
Adolf Engler (1844–1930), one of the greatest German botanists, was the first
one to try a synthesis of the evolution of the plant world on a global scale (Engler
1879, 1882). He classified the world flora into four realms and 31 regions, divid-
ing each region also into provinces and districts, thus designing a very detailed
hierarchical system that turned into the platform for all coming classification sys-
tems. Chile was classified into the Südamerikanische Florenreich (recognizing the
Nordchilenische Provinz as the Chilenische Übergangsgebiet from Grisebach) and
in the Altoceanisches Florenreich, South of 36◦ S. This Altoceanisches Florenreich
grouped southern Chile together with New Zealand’s South Island, the sub-Antarctic
islands, most of Australia and the Cape region of South Africa (Table 4.1), (Fig. 4.3).
“Engler was surprisingly perceptive in realizing that, scattered over the islands and
lands of the southernmost part of the world, lay the remains of a single flora, which
he called ‘the Ancient Ocean’ flora. It was over 80 years before acceptance of the
movement and splitting of continents at last explained this very surprising pattern
of distribution” (Cox and Moore 2005, p 26).
Later, Engler suggested that Australe Florenreich would be a better name as it
was characterized by the Austral-antarktischen Florenelement (Engler 1899, p 149).
He also added a fifth kingdom, the Ozeanisches Florenreich, which was composed
of the aquatic plants from the vast oceans.
Oscar Drude (1852–1933) worked closely with A. Engler for the publication of
the series “Die Vegetation der Erde”. Being a student from Grisebach, Drude found
several difficulties in synthesizing the floristic knowledge of his predecessors with
the growing ecological knowledge systematized by Grisebach (1872). Drude first
published his work Die Florenreiche der Erde based on a floristic approach (Drude
1884). He defined 14 floristic kingdoms and 55 floristic regions =Gebiete. Northern
b
Fig. 4.3 Floristic realms: a according to Engler (1882); b Drude (1884)
Chile to 41◦ S corresponded to the Andines Florenreich and southern Chile to the
Antarktisches Florenreich (Table 4.1) (Fig. 4.3).
Drude’s concern about the floristic and ecological differences led him to publish
separate maps for a floristic classification and for a vegetation classification (Drude
1887). Three years later he abandoned the floristic classification altogether: “The
maps published in the geographical reports of 1884 about my floristic classifica-
tion show the uncertainty of the boundary lines due to numerous migration routes
and directions of dispersal, which overlap from one to the other realm; it is a long
Fig. 4.4 Floristic realms according to Diels (1908)
known fact that each attempt to draw strict floristic boundaries, is itself ruinous”
(Drude 1890, p. 329). Drude decided to join the more physiognomic approach from
his mentor Grisebach, and he modified the classification on the basis of the new
climatological basis provided by Wladimir Köppen (1884). The floristic kingdoms
still numbered 14, and South America remained unmodified (Drude 1887).
Ludwig Diels (1874–1945), successor of Engler in Berlin, synthesized Drude’s
classification into six floristic realms, following the early proposal by Engler (1882)
but obviating the oceanic realm (Ozeanisches Florenreich), and dividing Engler’s
Altoceanisches Florenreich into an Antarktis, an Australis and a Capensis (Fig. 4.4).
Diels (1908) was the first to raise the African Cape region to the category of a
realm. He considered the Australisches Florenreich (Australis) as comprising only
Australia and Tasmania – and considered Malesia and New Zealand as part of
the Paläotropis. South America and Central America including Mexico and Baja
California was part of the Neotropis, but southernmost South America retained
its designation as a realm, the Antarktis. Diels (1908) little book was reprinted
five times until 1958, and his realm classification was retained adding only more
details at the regional scale by Mattick (1964) and later popular authors like Good
(1947) and Takhtajan (1986). Diels’ proposal modified by the mentioned authors is
still preferred in all modern German phytogeography textbooks (e.g. Richter 1997;
Schroeder 1998; Frey and Lösch 2004).
Almost contemporaneous to Diels (1908) is Christ’s (1910) specific proposal for
the world classification of ferns. As regards this plant group, Chile shared an Andean
flora with the other Andean territories, but harboured its own Chilean fern flora west
of the Andes and south of 39 ◦ S (Christ 1910).
The English botanist Ronald Good (1896–1992) published the first edition of
The Geography of the Flowering Plants in 1947. The work became one of the most
popular books in the field, reaching four editions and two reprints between 1947
and 1974. He followed Diels with the 6 realms, dividing them into 37 regions.
Chile south of 41◦ was classified into the “Antarctic kingdom”, and the north as
the “Neotropical kingdom”. The scheme is very similar to that of Russian botanist
Armen Takhtajan (1910–2009), which became very popular after its translation into
English. In a first version he maintained the 6 realms of Diels and the 37 regions of
Good (Takhtajan 1961) but in his later proposal he reduced the regions to 35. Diels’
scheme of 6 realms stayed unchanged (Takhtajan 1986). In his first classification he
considered Central Chile as part of the Neotropical region (Takhtajan 1961), but in
his later work he classified all the southern cone South of 25◦ into the Holantarctic
kingdom (Takhtajan 1986).
The basic scheme of 6 floristic realms proposed by Diels (1908) stayed
unchanged during the twentieth century. Only more recently Cox (2001) reanal-
ysed both long-standing floristic and faunistic schemes.1 For the global flora his
proposal was a rearrangement of the six former floristic realms into five, more or
less consistent with the continental shapes: the Holartic, South American, African,
Indo-Pacific, and Australian. This proposal has been disputed by Morrone (2002),
and is worth of being revised hereafter.
4.2 The Austral v/s the Neotropical Floristic Realm

We have known for more than a hundred years that southern South America, Tasmania-
Australia, and New Zealand are inhabited by numerous groups of plants and animals which
are more closely related to another than to any other group (Brundin 1966, p 8)
As described in Sect. 4.1, the Chilean flora has been alternatively classified under the
neotropical and/or the austral floristic realm. We can mention at least three visions
that are in conflict:
(a) the older view of Engler (1882), Diels (1908) and Skottsberg (1910), that draw
a boundary between the neotropical and the austral kingdoms at around 47◦ S;
(b) Takhtajan’s (1986) later proposal setting this limit at around 23◦ S;
(c) the more recent proposal of Cox (2001) that aligns all of the South American
flora under an “American kingdom”.
Regarding the former Antarctic kingdom Cox wrote: “. . . the consistency of the
plant geographical system is better served by transferring some of the regions of the
Antarctic Kingdom to the South American Kingdom and the rest to the Australian
Kingdom, in each case noting their individual historical and ecological characteris-
tics” (Cox 2001). The author was critical toward Takhtajan’s Holantarctic kingdom
because Takhtajan’s Holantarctic is based at the family level mainly on American
endemic families (e.g. Thyrsopteridaceae, Lactoridaceae, Gomortegaceae,
Aextoxicaceae). The dominance of American families seems to give reason to Cox
for transferring the areas of their occurrence to the American kingdom.
Cox’s vision is contested by Morrone (2002) due to neglecting the austral floris-
tic and faunistic relationships. Morrone (2002) related the classification to the his-
1 The faunistic regions date back to Sclater (1858) and Wallace (1876).
4.2 The Austral v/s the Neotropical Floristic Realm 137
tory of these biotas, coming to a scheme of only 3 biotic realms: the Holarctic
kingdom, the Tropical kingdom (=East Gondwana), and the Austral kingdom
(=West Gondwana). This was Engler’s early intention (1879, 1882); in fact the
result is remarkably similar to Engler’s, but it groups the paleotropis and neotropis
into one tropical realm.
Specifically reanalysing floristic relationships, recently Moreira-Muñoz (2007)
found support for the existence of an austral floristic realm, composed of more than
60 genera and 15 families which uniquely share the southern territories of South
America, Australasia and to a minor extent, by the Cape Floristic Region of South
Africa (Moreira-Muñoz 2007) (Table 4.2).
Table 4.2 Families composing the Austral floristic realm
Families Distribution Genera/species
Araucariaceae SE Asia, western Pacific, South America (disjunct 3/38

southern Chile/Argentina and SE Brazil)
Asteliaceae New Zealand, Tasmania, SE Australia, New Guinea, 2–4/36
Pacific Islands, Hawaii, Chile, Mascarenes
Atherospermataceae Chile/Argentina, Tasmania, Australia, New Zealand, 6/11
New Caledonia, New Guinea
Berberidopsidaceae Chile, E Australia 1/2
Calceolariaceae Tropical and W temperate South America, Brazil, 2/240–270
New Zealand
Centrolepidaceae SE Asia, Malesia, New Zealand, southern South 3/35
America
Corsiaceae Southern South America, SE Asia, 3/30
Papuasia-Australia
Escalloniaceae Central and South America, SE and SW Australia, 8/68
New Zealand, Réunion Is
Griseliniaceae New Zealand (2 spp.), Chile (5), one to Argentina 1/7
and a variety disjunct between Chile/SE Brazil
Luzuriagaceae Southern Argentina and Chile, Falkland Islands, 2/5
New Zealand and Australia (New South Wales to
Tasmania)
Nothofagaceae New Guinea, New Caledonia, E Australia, New 1/36
Zealand, South America
Proteaceae Australasia, Africa, South America 70/1,000
Restionaceae Australasia, South Africa, Madagascar, South 58/520
America
Stylidiaceae = Scattered in South East Asia to Australia and New 6/157
Donatiaceae Zealand, southern South America
Source: Moreira-Muñoz (2007), by permission of John Wiley and Sons.
4.2.1 Floristic Elements in the Latitudinal Profile

Moreira-Muñoz (2007) provided support for the further recognition of an austral
and a neotropical floristic realm in South America, but the question of the boundary
between both still remains.
After grouping the Chilean plants into seven floristic elements (Kontingente),
Reiche (1907, p 319) (Table 3.1) proposed that the neotropical and antitrop-
ical (Reiche’s Californian) elements occur in Chile due to a migration route
(Wanderungslinie) from the north to the south along the Andes down to Magallanes,
until this migration was stopped at the border of the “Antarctic realm” (situated
sensu Reiche from ~ 40◦ to the south along the coast).
Schmithüsen (1956) plotted the floristic elements along the latitudinal gradient.
He defined therefore eight elements from the northern neotropical to the southern-
most evergreen subantarctic forests. He could not define a sharp limit and proposed
spatial imbrications of the different elements in Central Chile between 30◦ and
40◦ S (Fig. 4.5). Setting sharp limits between floristic/biogeographic units always
has been problematic (e.g. Drude 1890, Sect. 4.1), but the analysis of the range
extension of taxa has been ever informative in this respect (e.g. Godley 1963; Reiche
1907).
The seven floristic elements distinguished here (Sect. 3.1) contain genera with a
diversity of geographic range sizes. In every element, with exception of the endemic
element, there are some genera distributed over the whole country, from the south-
ernmost Cabo de Hornos to the northern Parinacota province. Every element also
30°
x 1. General floristic area of

x neotropical origin
2. Neotropical endemic flora
x from La Serena
x x 3. Neotropical sclerophyllous
element
x
x x 4. Laurophyllous neotropical
x x x and south-hemispheric origin
x x x
x x 5. Forest flora of south-
x x x hemispheric
x x
subtropical origin
x x
x x 6. Temperate rainforest of
40°
neotropical and south-
x hemispheric origin
x x 7. Deciduous forest of sub-
x antarctic origin
Fig. 4.5 Floristic elements in x
Central Chile (adapted from 8. Evergreen forest of sub-
x antarctic origin
Schmithüsen 1956)
Table 4.3 Average of

geographic range of floristic Floristic element Average (◦ S)
elements
3. Neotropical 29.6
1. Pantropical 32.2
6. Endemic 32.2
4. Antitropical 35.1
7. Cosmopolitan 36.0
2. Australasiatic 41.5
5. South-temperate 42.3
contains a number of genera known for their very restricted range or known only
from a couple of localities. One method of assessing the main massing of the genera
is to calculate for every element the average distribution of the genera that compose
them [northern limit + southern limit/2]. Results are shown in Table 4.3.
The neotropical element has the northernmost average (29.6◦ S), while the
south-temperate element has the southernmost average (42.3◦ S). The australasiatic
element has an average at 41.5◦ S. The pantropical, antitropical, cosmopolitan and
endemic elements show average latitudes between 32◦ and 36◦ S. The fact that most
elements have their average in Central Chile tends to reinforce the early view of
Grisebach (1872), Engler (1882), and Schmithüsen (1956) to consider this region a
transition zone with different, converging elements. Central Chile is still considered
as a key region for the understanding of the biogeographical history of several South
American plant groups, like the Bromeliaceae (Schmidt Jabaily and Sytsma 2010).
The southernmost average reached by south-temperate and australasiatic genera
show the consistency of the relationship between southern Chile and other austral
territories like New Zealand (Moreira-Muñoz 2007).
4.2.2 Similarity Along the Latitudinal Gradient

Since the latitudinal ranges of the genera does not account for possible distribu-
tion or collection gaps, four regional floras were selected for a similarity analysis:
Antofagasta (ANT), Coquimbo (COQ), Biobío (BIO), and Magallanes (MAG)
(Fig. 4.6). The regions included in the analysis have a floristic checklist (Table 4.4),
that has been revised and homologized. Regions analysed are dissimilar in area, but
some of them harbour similar numbers of native genera (e.g. COQ=457, BIO=465),
MAG showing the lowest generic richness: MAG=252.
The Jaccard similarity index was calculated for the data sets (Zunino and Zullini
2003; Cox and Moore 2005; Lomolino et al. 2006). The highest floristic similarity
is between COQ and BIO, sharing 323 genera (Table 4.5). The lowest similarity is
shown by ANT and MAG, which share 97 genera. There seems to be a relationship
between the similarity and the geographic distance, and in Fig. 4.6 both variables
are represented, showing this trend of decreasing similarity as the regions are further
apart.
72° 68°
1.0
18°
22°
0.8
ANT
26°
Jaccard similarity
0.6
30°
COQ
COQ/BIO
34°
ANT/COQ
0.4
BIO
38°
BIO/MAG
ANT/BIO COQ/MAG
42°
0.2
ANT/MAG
46°
0.0
50°
0 2000 4000 6000

54°
Distance (km) MAG
Fig. 4.6 a Similarity versus geographic distance between Chilean regions; b location of compared
regions
Table 4.4 Regions used in the floristic similarity analysis
N◦ native vascular
Abbreviation Regions Area (km2 ) plant genera Source
ANT Antofagasta 126,049 316 Marticorena et al. (1998)

COQ Coquimbo 40,580 457 Marticorena et al. (2001)
BIO Biobío 37,063 465 CONC herbarium,
R. Rodríguez pers.
comm.
MAG Magallanes 132,033 252 Henríquez et al. (1995)
Table 4.5 Floristic similarity and geographic distance of different regions in Chile: Antofagasta
(ANT), Biobío (BIO), Coquimbo (COQ) and Magallanes (MAG). Geographic distance has been
calculated as the latitudinal difference between the respective geographic centroid of each region
Compared regions Shared genera Similarity Jaccard Distance (km)
ANT/COQ 244 0.46 777

ANT/BIO 174 0.29 1, 527
ANT/MAG 97 0.21 3, 227
COQ/BIO 323 0.54 750
COQ/MAG 150 0.27 2, 450
BIO/MAG 178 0.33 1, 700
Table 4.6 Floristic elements in each of the compared regions
South-
Pantropical Australasiatic Neotropical Antitropical temperate Endemic Cosmopolitan
REG n % n % n % n % n % n % n %
ANT 28 8.9 8 2.5 115 36.5 45 14.3 17 5.4 15 4.8 82 26.0

COQ 39 8.6 22 4.9 114 25.1 73 16.1 42 9.3 38 8.4 117 25.8
BIO 42 9.1 41 8.9 100 21.6 72 15.6 51 11.0 27 5.8 123 26.6
MAG 11 4.4 38 15.1 29 11.5 44 17.5 42 16.7 1 0.4 84 33.5
These regions have been further compared regarding the floristic elements com-
posing these regional floras. Results indicate a trend along the latitudinal gradient
in Chile: the cosmopolitan, antitropical, south-temperate, and australasiatic genera
show a proportional increase towards the south, while the proportion of neotropical,
pantropical and endemic genera decrease towards the south (Table 4.6, Fig. 4.7).
Highly remarkable is the replacement of the neotropical by the australasiatic genera
between BIO and MAG (arrow in Fig. 4.7).
The replacement of neotropical genera by australasiatic genera in southernmost
Chile (Magallanes) clearly shows the consistency of the inclusion of subantarc-
tic Chile south of 47◦ in an austral floristic realm, as earlier proposed by Engler
(1882), Drude (1884), Reiche (1907, p 282), Diels (1908), and Skottsberg (1916).
In Skottsberg’s Plant geographical map of South America south of 41◦ latitude
(Skottsberg 1910), the author proposed a limit at 47◦ –48◦ S, between a more
species-rich temperate rainforest to the north, and a “subantarctic” species-poor
temperate rainforest to the south. In his more complete account of the southern
flora, he explicitly proposed a limit between an “Andean floristic region” and a
40
35
30
1. Pantropical
25 2. Australasiatic
3. Neotropical
% 20 4. Antitropical
5. South-temperate
15 6. Endemic
7. Cosmopolitan
10
5
0
ANT COQ BIO MAG
regions
Fig. 4.7 Floristic elements present in four regional floras (percentage). Arrow showing relative
replacement of neotropical genera by australasiatic ones
“Subantarctic region” from 48◦ S to the south (including the Falkland [Malvinas]
Islands) (Skottsberg 1916, pp 344–345).
4.3 Regions and Provinces

Below the realm level, several schemes have been proposed for the classification of
the Chilean territory into regions, subregions and provinces, mostly by means of the
distribution of faunal groups (Artigas 1975; Morrone et al. 1997; Hernández et al.
2005; Casagranda et al. 2009).
Biogeographic schemes for Chile based on floristic analysis are sparse. The first
plant geographical map for Chile accompanied Reiche’s analysis of the distribution
of the Compositae (Asteraceae) family (Chap. 8). He defined, based on his experi-
ence, ten floristic regions and subregions for the Chilean Compositae (Chap. 8 and
Fig. 8.7). On a second map he also proposed possible migration routes for these taxa
(Fig. 8.7).
The first attempt towards a synthetical cartography for the Chilean flora is found
in the two maps that accompanied Reiche’s Plant geography (Reiche 1907) at a
scale 1:7,500,000 (Fig. 4.8). In the first map Reiche proposed distribution ranges
and limits for some key taxa. In the second map he divided the country in several
floristic units, integrating floristic and physiognomic knowledge (Fig. 4.8).
Reiche proposed the limit between the Antarctic and Neotropical realms at
around 40◦ S (Table 4.7). Although the resulting picture looks rather simple, this
nevertheless is the first attempt of a floristic cartography for Chile. After Reiche,
attempts turned into a more physiognomic approach (see Sect. 1.3).
In the search for a synthetic view that integrates floristic and faunistic infor-
mation, the most long-standing scheme for South America is the one by Cabrera
and Willink (1973). They classified Chile into two regions (Neotropical and
Antarctic) and dominions (Andino-Patagónico and Subantártico). They further sub-
divided these areas in four provinces: provincia del Desierto, provincia Puneña
(high Andes), provincia Chilena, and provincia Subantártica. They subdivided the
provinces into smaller districts (not mapped). This was a remarkable attempt to inte-
grate historical and ecological information into one classification scheme, although
the approach still remained physiognomic rather than floristic/faunistic (see a revi-
sion by Ribichich 2002). Also some confusion arose since Cabrera and Willink
(1973) translated the traditional concept of the realm as región biogeográfica and the
traditional concept of the region as dominio biogeográfico (Fig. 4.1). Interestingly,
their provinces indeed correspond to a third level in a hierarchy, and have been more
recently retrieved by Morrone (2001) (Fig. 4.9a).
Morrone (2001) applied individual tracks for the definition of the provinces and
emphasises the importance biogeographic homology, i.e. a common evolutionary
history in the definition of the provinces. He also argues that one of the main prob-
lems is to define the limits between the units. In fact, several authors have focused
in the transition zones rather than in the cores of the units (Fig. 4.5) (Ruggiero and
Ezcurra 2003; Morrone 2006; see Drude’s early concerns in Sect. 4.1).
4.3 Regions and Provinces 143
a b
Fig. 4.8 Maps accompanying Reiche’s Plant Geography (1907): a floristic regions, see Table 4.7;
b distribution limits for selected taxa
Table 4.7 Floristic units from the “Plant Geography” of Reiche (1907)
Regions and subregions Extension
I North Chile 18◦ –30◦ 40′ S

(A) Coast Arica to Antofagasta 18◦ –23◦ 30′ S, characteristic Euphorbia
lactiflua
(B) Coast Antofagasta to Caldera 23◦ 30′ –27◦ S
(C) Coast Caldera to Fray Jorge (río Limarí) 27◦ –30◦ 40′ S, the southern limit is at Fray
Jorge, the “northernmost advance of the
hygrophilous forest”
(D) Interior Cordilleras 18◦ –30◦ 40′ S
II Central Chile 30◦ 40′ S–37◦ , Río Limarí (Fray Jorge
forest) to Concepción
(A) Coast Fray Jorge to Valparaíso 30◦ 40′ –33◦ 30′ S
(B) Coast Valparaíso to Maule 33◦ 30′ –35◦ S
(C) Maule to Concepción 35◦ –37◦ S
(A′ ) Cordillera Ovalle to Santiago 30◦ 40–33◦ 30′ S
(B′ ) Cordillera Santiago to Rancagua 33◦ 30′ –35◦
(C′ ) Cordillera Rancagua to Chillán 35◦ –37◦
III South Chile 37◦ –54◦ S
(A) Coast Concepción to Río Imperial 37◦ –39◦ S
(B) Coast Río Imperial to Chiloé 39◦ –43◦ 30′ S (At 40◦ , northern limit of the
Antarctic floristic realm)
(C) Coast Chiloé to Taitao 43◦ 30′ –47◦ S
(D) Coast Taitao peninsula to Cabo de Hornos 47◦ –56◦ S
(A′ ) Cordillera Chillán to Panguipulli 37◦ –39◦ 30′ S
(B′ ) Cordillera Panguipulli to Campo de Hielo Sur 39◦ 30′ –48◦ S
(C′ ) Campo de Hielo Sur to Magallanes 48◦ –55◦ S, Cordillera and Magallanes
steppe
4.3.1 Endemism as the Base for Regionalization

Most regionalizations rest upon endemic taxa. Since Augustin Pyramus de Candolle
(1820) coined the term, endemism has turned out to be one of the most appealing
concepts in historical biogeography. A geographic area that contains two or more
non-related endemic taxa is formally defined as an area of endemism, a concept of
vital importance in modern historical biogeography (Harold and Mooi 1994; Linder
2001). In the words of Nelson and Platnick (1981) “the most elementary questions
of historical biogeography concern areas of endemism and their relationships”.
Several methods could be applied nowadays to the problem of regionalization
of Chile: methods in common use are Parsimony Analysis of Endemicity (PAE),
and the optimality method (NDM/VNDM). Both methods have been widely used in
several geographic areas and taxa (Nihei 2006; Escalante 2009).
An exercise is presented here, done with the Chilean endemic plant genera
(Sect. 3.1). 65 genera endemic to continental Chile were considered for the analysis,
since remaining genera are known from just one or two localities. Several individual
distribution maps of endemic genera were presented in Fig. 3.13.
4.3 Regions and Provinces 145
20!
58
59
61
30!
63
62
64
40!
Area 0
Area 1
65
Area 2
69
70
50!
66
67
Fig. 4.9 Biogeographic provinces for Chile: a according to Morrone (2001): Puna (58), Atacama
(59), Coquimbo (61), Santiago (62), Juan Fernández (63), Maule (64), Valdivian Forest (65),
Magellanic Forest (66), Magellanic Páramo (67), Central Patagonia (69), Subandean Patagonia
(70); b according to consensus areas of endemism obtained by NDM/VNDM on the base of the
endemic genera
PAE. This method classifies areas using shared taxa using parsimony (Rosen
1988; Nihei 2006). In a PAE cladogram, groups of areas sharing exclusive taxa
(based at least on two synapomorphies) are identified as areas of endemism. Since
PAE does not perform satisfactorily with fewer characters (taxa) than species
(quadrats), it was not applied for the analysis of the endemic genera. It was used
for the regionalization of Chilean Cactaceae species (Sect. 7.2).
NDM/VNDM. The optimality algorithm is a biogeographic method to identify
areas of endemism that calculates an endemicity index for a set of areas based on the
adjustment of the distributions of two or more species (Szumik et al. 2002; Szumik
and Goloboff 2004). The algorithm is implemented in the software NDM/VNDM
(Goloboff 2005). Recent results are showing that the use of this algorithm can
improve the number of areas of endemism obtained by other methods (Casagranda
et al. 2009; Carine et al. 2009; Escalante et al. 2009).
The input for the program NDM is the presence/absence of taxa in grid cells.
In this case, 59 quadrats of 1 × 1 degree lat/long cover the area of mainland Chile
occupied by endemic genera. Collection localities are entered as xy data. The opti-
mality method was carried out with NDM/VNDM v. 2.6 (Goloboff 2005) with the
next options: saving sets with two or more endemic genera, and saving sets with
a score above 2.0. The type of swapping was of one cell at a time, with 0% of
unique species in overlapping subsets. The search was performed until the number
of sets was showed stable in 20 repetitions with different random seeds, using edge
proportions. With these parameters, 17 areas of endemism were obtained.
Consensus of 30% of similarity was applied to the obtained sets. With the
option “against any of the other areas in the consensus” three consensus areas were
obtained, to some extent overlapping. Genera contributing to the final score are those
that reach more than 0.5 (Escalante pers. comm., with 1 the highest value possible
for a single genus) (Fig. 4.9a and b).
Consensus Area n◦ 0 is built upon eight sets. It is supported by 14 endemic gen-
era reaching a score of 4.3. It includes 18 quadrats in Central Chile between at
28◦ (coast) and 37◦ S. Its endemic genera are Adenopeltis, Avellanita, Epipetrum,
Jubaea, Miersia, Moscharia, Ochagavia, Placea, Pleocarphus, Speea, Tecophilaea,
Tetilla, Traubia and Trevoa. The consensus includes two genera with scores below
0.5 (Cyssatobryon and Neoporteria).
Consensus Area n◦ 1 was obtained from two sets, supported by the presence of
five endemic genera and a score of 3.71. It includes 26 quadrats in Central-South
Chile between 31◦ (coast) and 42◦ S. Its endemic genera are Acrisione, Fascicularia,
Gymnachne, Lapageria and Vestia. The consensus includes three genera with scores
below 0.5 (Francoa, Peumus and Nothanthera).
Consensus Area n◦ 2 is built by seven sets, formed by 10 endemic genera and
a score of 3.52. It includes 18 quadrats in North-Central Chile between 24◦ and
33◦ S. Its endemic genera are Bakerolimon, Balsamocarpon, Bridgesia, Copiapoa,
Dinemagonum, Eriosyce, Leontochir, Phrodus, Pintoa and Thelocephala. The
consensus includes one genus with a score below 0.5 (Microphyes).
Comparing the areas of endemism recognized by NDM with previous biogeo-
graphic schemes, it appears noteworthy that Area 2 is perfectly coincident with
Morrone’s (2001) provincia de Coquimbo and to some extent with the distrito
Coquimbano from Cabrera and Willink (1973).2 On the contrary, Area 1 encloses
part of Morrone’s (2001) provincia de Coquimbo, the provincia de Santiago and
provincia del Maule, and the northern part of the provincia del Bosque Valdiviano.
The northern limit of NDM’s Area 1 is coincident with the location of the Fray Jorge
forest, a long recognized northern advance of the Valdivian forest (Box 3.1). The Río
Limarí and Fray Jorge forest was already proposed by Reiche as the limit between
2 Compared with the description in Cabrera and Willink’s text, not the map!
References 147
the floristic regions of North Chile and Central Chile (Table 4.7). Finally, NDM’s
Area 0 encompasses most genera strictly endemic (14) to Central Chile between
28◦ and 37◦ . This area is highly coincident, especially in its southern limit, with the
region of Central Chile from Reiche (1907).
The three consensus areas can be interpreted as having each an evolutionary his-
tory. It is interesting that consensus area 0 has a degree of overlap with the one
to the north (Area 2) and the one to the south (Area 1), showing even a range of
overlap between the three from 31◦ (coast) to 33◦ S. This range can be interpreted
as a panbiogeographic node, i.e. an area where different evolutionary lines coincide
(Heads 2004). Indeed, at 33◦ S, La Campana National Park has been long recognized
as a mosaic of elements of Neotropical and Gondwanic origin (Luebert et al. 2002),
showing also the extant northern distribution limit of the genus Nothofagus in the
Southern Cone of America (Chap. 9).
The results of this exercise further show that the units of a biogeographic region-
alization rarely have sharp limits and, on the contrary, tend to superpose in the form
of transition areas. Still, results seem to be highly dependent on the grid size, the
origin assigned to the grid, the data base strength, the number of species analysed
and the search parameters (Carine et al. 2009). More exercises on this topic have
been done with the Chilean Cactaceae and Asteraceae in Chaps. 7 and 8.
Finally, the naming of biogeographic units resulting from a regionalization is not
a trivial thing, and a formal system has been recently proposed by Ebach et al. (2008)
as an International Code of Area Nomenclature, ICAN. Further developments in
this field of research should take account of this systematization proposal of the
Systematic and Evolutionary Biogeographical Association, SEBA (http://www.uac.
pt/~seba/).
References
Artigas J (1975) Introducción al estudio por computación de las areas zoogeográficas de Chile
continental basado en la distribución de 903 especies terrestres. Gayana (Chile) 4:1–25
Brundin L (1966) Transantarctic relationships and their significance, as evidenced by chironomid
midges: with a monograph of the subfamilies Podonominae and Aphroteniinae and the austral
Heptagyiae. K Sven Vetenskapsakad Handl 11:1–472
Cabrera AL, Willink A (1973) Biogeografía de América Latina. Monografía 13. Serie de Biología.
(2nd Revised edn 1980). Secretaría General de la Organización de los Estados Americanos,
Washington, DC
Carine MA, Humphries CJ, Guma R, Reyes-Betancort JA, Santos Guerra A (2009) Area and
algorithms: evaluating numerical approaches for the delimitation of areas of endemism in the
Cabary Islands archipelago. J Biogeogr 36:593–611
Casagranda MD, Roig-Juñent S, Szumik C (2009) Endemismo a diferentes escalas espaciales: un
ejemplo con Carabidae (Coleóptera: Insecta) de América del Sur austral. Rev Chil Hist Nat
82:17–42
Christ H (1910) Die Geographie der Farne. Gustav Fischer Verlag, Jena
Cox CB (2001) The biogeographic regions reconsidered. J Biogeogr 28:511–523
Cox CB, Moore PD (2005) Biogeography: an ecological and evolutionary approach, 7th edn.
Blackwell Publishing, Oxford
de Candolle A (1835) Introduction à l’étude de la Botanique I, II. Librairie Encyclopédique de

Roret, Paris
de Candolle A (1855) Géographie botanique raisonnée, 2. vol. Paris
de Candolle, A-P (1820) Geographie botanique. In: Dictionnaire des sciences naturelles, XVIII.
Flevrault, Strasbourg, pp 359–422
Diels L (1908) Pflanzengeographie. Göschen’sche Verlagshandlung, Leipzig
Drude O (1884) Die Florenreiche der Erde: Darstellung der gegenwärtigen Verbreitung
sverhältnisse der Pflanzen: Ein Beitrag zur vergleichenden Erdkunde. Ergänzungsheft N◦ 74,
Petermanns Mitteilungen, Gotha, Justus Perthes
Drude O (1887) Atlas der Pflanzenverbreitung. Berghaus” Physikalischer Atlas, Abteilung V,
Gotha, Justus Perthes
Drude O (1890) Handbuch der pflanzengeographie. Bibliothek Geographischer Handbücher (ed.
by F Ratzel). Verlag J. Engelhorn, Stuttgart
Ebach MC, Goujet DF (2006) The first biogeographical map. J Biogeogr 33:761–769
Ebach MC, Morrone JJ, Parenti LR, Viloria AL (2008) International code of area nomenclature.
J Biogeogr 35:1153–1157
Eldredge N (1985) Unfinished synthesis: biological hierarchies and modern evolutionary thought.
Oxford University Press, New York, NY
Engels E-M (2005) Ethik in den biowissenschaften. In: Maring M, (ed) Ethisch-Philosophisches
Grundlagenstudium 2. Ein Projektbuch. Münster, LIT Verlag, pp. 135–166
Florengebiete seit der Tertiärperiode. Vol 1. Die Extratropischen Gebiete der Nördlichen
Hemisphäre. Verlag von W. Engelmann, Leipzig.
Florengebiete seit der Tertiärperiode. Vol. 2. Die Extratropischen Gebiete der Südlichen
Hemisphäre und die Tropischen Gebiete. Verlag von W. Engelmann, Leipzig.
Engler A (1899) Die Entwickelung der Pflanzengeographie in den letzten hundert Jahren und
weitere Aufgaben derselben. (ed by Gesellschaft für Erdkunde zu Berlin) pp 1–247
Escalante T (2009) Un ensayo sobre regionalización biogeográfica. Rev Mex Biodivers 80:
551–560
Escalante T, Szumik C, Morrone JJ (2009) Areas of endemism of Mexican mammals: reanalysis
applying the optimality criterion. Biol J Linn Soc 98:468–478
Frey W, Lösch R (2004) Lehrbuch der Geobotanik, 2nd edn. Elsevier-Spektrum Akad. Verlag,
Munich
Godley EJ (1963) Contributions to the plant geography of southern Chile. Rev Univ (Chile)
48:31–39
Goloboff P (2005) NDM/VNDM ver. 2.6. Programs for identification of areas of endemism. http://
www.zmuc.dk/public/phylogeny/endemism
Good R (1947) The geography of the flowering plants. 4th edition 1974. Longman, London
Grehan JR (2001) Panbiogeography from tracks to ocean basins: evolving perspectives. J Biogeogr
28:413–429
Grisebach A (1872) Die vegetation der Erde nach ihrer klimatischen Anordnung: ein Abriss der
vergleichenden Geographie der Pflanzen, 2 vols, 1st edn. Engelmann Verlag, Leipzig. (2d
revised ed. 1884)
Harold AS, Mooi RD (1994) Areas of endemism – definition and recognition criteria. Syst Biol
43:261–266
Heads M (2004) What is a node? J Biogeogr 31:1883–1891
Henríquez JM, Pisano E, Marticorena C (1995) Catálogo de la flora vascular de Magallanes (XII
Región), Chile. An Inst Pat Ser Cs Nat (Chile) 23:5–30
Hernández CE, Moreno RA, Rozbaczylo N (2005) Biogeographical patterns and Rapoport’s rule
in southeastern Pacific benthic polychaetes of the Chilean coast. Ecography 28:363–373
Huxley CR, Lock JM, Cutler DF, (eds) (1998) Chorology, taxonomy and ecology of the floras of
Africa and Madagascar. Royal Botanic Gardens, Kew
References 149
Jagers op Akkerhuis GAJM (2008) Analysing hierarchy in the organization of biological and
physical systems. Biol Rev 83:1–12
Köppen W (1884) Die Wärmezonen der Erde, nach der Dauer der heissen, gemässigten and kaltern
Zeit and nach der Wirkung der Wärme auf die organische Welt betrachtet. Meteor Z 1:215–226
Linder HP (2001) On areas of endemism, with an example from the African Restionaceae. Syst
Biol 50:892–912
Llorente Bousquets J, Morrone JJ, (eds) (2005) Regionalización biogeográfica en Iberoamérica y
tópicos afines. Las Prensas de Ciencias, Facultad de Ciencias, UNAM, México DF
Lomolino MV, Riddle BR, Brown JH (2006) Biogeography, 3rd edn. Sinauer Associates,
Sunderland, MA
Luebert F, Muñoz-Schick M, Moreira-Muñoz A (2002) Vegetación y Flora de La Campana. In:
Elórtegui S, Moreira-Muñoz A, (eds) Parque Nacional La Campana: origen de una Reserva de
la Biosfera en Chile Central. Taller La Era, Santiago
Mackey B, Berry S, Brown T (2008) Reconciling approaches to biogeographical regionalization:
a systematic and generic framework examined with a case study of the Australian continent.
J Biogeogr 35:213–229
Marticorena C, Matthei O, Rodríguez R, Arroyo MTK, Muñoz-Schick M, Squeo FA, Arancio G
(1998) Catálogo de la Flora Vascular de la Segunda Región (Región de Antofagasta), Chile.
Gayana Bot 55:23–83
Marticorena C, Squeo FA, Arancio G, Muñoz-Schick M (2001) Catálogo de la flora vascular de
la IV Región de Coquimbo. In: Squeo FA, Arancio G, Gutiérrez JR, (eds) Libro Rojo de la
Flora Nativa y de los Sitios Prioritarios para su Conservación: Región de Coquimbo. Ediciones
Universidad de La Serena, La Serena, pp 105–142
Mattick F (1964) Übersicht über die Florenreiche und Florengebiete der Erde. In: Melchior H,
(ed) A. Engler’s Syllabus der Pflanzenfamilien, vol. 2, 12th edn. Angiospermen, Gerbrüder
Borntraeger, Berlin, pp 626–629
McLaughlin SP (1992) Are floristic areas hierarchically arrange? J Biogeogr 19:21–32
McLaughlin SP (1994) Floristic plant geography: the classification of floristic areas and floristic
elements. Prog Phys Geogr 18:185–208
McLaughlin SP (2007) Tundra to tropics: the floristic plant geography of North America. Sida Bot
Misc 30:1–58. Botanical Research Institute of Texas
Mennema J (1985) The first plant distribution map. Taxon 34:115–117
Moreira-Muñoz A (2007) The Austral floristic realm revisited. J Biogeogr 34:1649–1660
Morrone JJ (2001) Biogeografía de América Latina y el Caribe. Manuales y Tesis SEA 3, Zaragoza
Morrone JJ (2002) Biogeographical regions under track and cladistic scrutiny. A comment on CB
Cox (2001). J Biogeogr 29:149–152
Morrone JJ (2006) Biogeographic areas and transition zones of Latin America and the Caribbean
Islands based on panbiogeographic and cladistic analyses of the entomofauna. Annu Rev
Entomol 51:467–494
Morrone JJ, Katinas L, Crisci JV (1997) A cladistic biogeographic analysis of Central Chile.
J Comp Biol 2:25–42
Nelson G (1978) From Candolle to Croizat: comments on the history of biogeography. J Hist Biol
11:289–305
Nelson G, Platnick N (1981) Systematics and biogeography: Cladistics and vicariance. Columbia
University Press, New York, NY
Nihei SS (2006) Misconceptions about parsimony analysis of endemicity. J Biogeogr 33:2099–
2106
Proches S (2008) Three ways to split a continent. J Biogeogr 35:195–196
Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, vol. VIII. Verlag von
Ribichich AM (2002) El modelo clásico de la fitogeografía de Argentina: un análisis crítico.
Interciencia 27:669–675
Richter M (1997) Allgemeine Pflanzengeographie. Teubner Studienbücher der Geographie,

Stuttgart
Rosen BR (1988) From fossils to earth history: applied historical biogeography. In: Myers AA,
Giller PS, (eds) Analytical biogeography. Chapman and Hall, London, pp 437–481
Ruggiero A, Ezcurra C (2003) Regiones y transiciones biogeográficas: complementariedad de los
análisis en biogeografía histórica y ecológica. In: Morrone JJ, Llorente J, (eds) Una perspectiva
latinoamericana de la biogeografía. Las Prensas de Ciencias, UNAM, Mexico DF, pp 141–154
Schmidt Jabaily R, Sytsma KJ (2010) Phylogenetics of Puya (Bromeliaceae): placement, major
lineages, and evolution of Chilean species. Am J Bot 97:337–356
Schmithüsen J (1956) Die räumliche Ordnung der chilenischen Vegetation. Bonn Geogr Abh
17:1–86
Schouw JF (1823) Grundzüge einer allgemeinen Pflanzengeographie (+ Atlas). Reimer, Berlin
Schroeder F-G (1998) Lehrbuch der Pflanzengeographie. Quelle and Meyer, Wiesbaden
Sclater PL (1858) On the general geographical distribution of the members of the class Aves. Zool
J Linn Soc 2:130–145
Skottsberg C (1910) Botanische Ergebnisse der schwedischen Expedition nach Patagonien
unde dem Feuerlande 1907–1909. I. Übersicht über die wichtigsten Pflanzenformationen
Südamerikas S von 41◦ , ihre geographische Verbreitung und Beziehungen zum Klima. K Sven
Vetenskapakad Handl 46:1–28
Skottsberg C (1916) Botanische Ergebnisse der schwedischen Expedition nach Patagonien und
dem Feuerlande 1907–1909. V. Die Vegetationsverhältnisse längs der Cordillera de los Andes
südlich von 41 Grad südlicher Breite. K Sven Vetenskapsakad Handl 56:1–366. Stockholm,
Almqvist and Wiksell
Stuessy TF (2006) Principles and practice of plant taxonomy. In: Leadlay E, Jury S, (eds)
Taxonomy and plant conservation. Cambridge University Press, Cambridge, pp 31–44
Szumik CA, Cuezzo F, Goloboff P, Chalup AE (2002) An optimality criterion to determine areas
of endemism. Syst Biol 51:806–816
Szumik C, Goloboff P (2004) Areas of endemism: an improved optimality criterion. Syst Biol
53:968–977
Takhtajan A (1961) The origin of angiospermous plants, 2nd edn. original Russian edition, Gos.
izd. Vysšaja Škola, Moscow
Takhtajan A (1986) Floristic regions of the world. University of California (translated from the
original Russian edition 1978, Leningrad). Reprint (1988), B Singh, MP Singh, Dehra Dun,
India
Thiselton-Dyer WT (1909) Geographical distribution of plants. In: Seward AC (ed) Darwin and
modern science. Cambridge University Press, Cambridge, pp 298–318
Treviranus GR (1803) Biologie, oder Philosophie der lebenden Natur. Röwer, Göttingen
Wallace AR (1876) The geographical distribution of animals. Macmillan and Co, London
Zunino M, Zullini A (2003) Biogeografía: la dimensión espacial de la evolución. Fondo de Cultura
Económica, México. From the 1995 Italian edition
Part III
Islands Biogeography
Chapter 5
Pacific Offshore Chile
Abstract The study of island biotas has been one of the most productive
issues in biogeography. Indeed, one of the most interesting and challenging
aspects of Chilean plant geography correspond to the Pacific islands offshore the
American continent. This applies especially for the Juan Fernández and the Islas
Desventuradas archipelagos that are analyzed regarding their geographical relation-
ships. The flora of Juan Fernández is especially attractive for biogeography due to
the presence of many locally endemic taxa, and a primitive endemic family: the
Lactoridaceae, represented by the only species Lactoris fernandeziana. Finally, the
threatened status of most of the Fernandezian flora is discussed with attention for
the possibilities of conservation and restoration.
The Pacific islands offshore the continent have been called “the portion of Chile that
is not in America”1 (Fig. 1.1). These are three separate clusters of islands that are
emerged seamounts on the Nazca Plate, formed by intraplate volcanoes that occur
along linear chains (Stern et al. 2007). The volcanic origin of these islands is most
evident in Rapa Nui, which harbors three coalescent volcanic centres (Rano-Kau,
Poike, and Maunga Terewaka), plus ca 70 secondary eruptive centres (IGM 2005).
5.1 Rapa Nui

Rapa Nui (= Isla de Pascua, Easter Island) (27◦ 05′ S, 109◦ 20′ W, 160 km2 ) (Fig. 5.1)
is known all over the world due to its peculiar culture and the moai sculptures.
Considered the most isolated inhabited island on the planet, it lies 3,760 km away
from the South American continent, and 3,800 km from Tahiti. It occurs at the west-
ern edge of a chain of volcanic seamounts that also includes Isla Salas y Gómez
(26◦ 27′ S, 105◦ 28 W) (see Fig. 1.1). In spite of the geographical isolation, Rapa Nui
has a long history of human occupation. The timing of the agricultural coloniza-
tion of Rapa Nui has been dated at ca 1200 AD, and was accompanied by rapid
deforestation, probably exacerbated by the primeval fires (Mann et al. 2008). Bork
1 Balcells (2005) called it “el Chile que no está en América”.

DOI 10.1007/978-90-481-8748-5_5, ⃝
154 5 Pacific Offshore Chile
Fig. 5.1 Rapa Nui (Easter Island) after Skottsberg (1920–1956)
(2006) proposed, on the base of soil analysis, that the extinct palm Paschalococos
disperta, once occupied the whole island. The isolated system was not able to sup-
port the human impact and the community collapsed (Bologna and Flores 2008).
Only recent decades have seen a rebirth of the community promoted by the national
and international interest on the ancient culture (Arancibia 2009).
From a botanical point of view, Isla de Pascua is the poorest of the Chilean Pacific
islands, showing only a 7.7 % of specific endemism (Marticorena 1990). There are
anyhow some interesting native genera and families (mostly ferns) not present in
continental Chile, such as Davallia, Psilotum, Vittaria, Doodia (Chap. 2). These are
pantropical genera, widely distributed on the Pacific Islands.
The long history of occupation by Polynesian folks has left a landscape and
a floristic poverty that seems to be very far from the original one (prior to the
arrival of man) (Zizka 1991; Bork 2006). Based on the works of Fuentes (1913)
(Fig. 5.2) and Skottsberg (1920–1956), more recently revised by Marticorena (1990)
and Zizka (1991), a checklist with a total of 29 native genera pertaining to 20
families was compiled (Table 5.1). A synopsis of the native flora is very difficult:
several taxa are treated as either native or alien by different authors (or idiochores
v/s anthropochores sensu Zizka 1991).
Several taxa that appeared with a question mark in Zizka’s revision and are con-
sidered as introduced by Marticorena (1990) have been left out from the analysis
(e.g. Caesalpinia, Calystegia), while some listed by Marticorena as aliens have
been retained due to the reasons given by Zizka, such as Triumfetta (Malvaceae), or
Kyllinga (Cyperaceae). The best represented family is the Poaceae, with 5 genera.
The resulting phytogeographic analysis shows a clear predominance of pantropical
(41%) and cosmopolitan genera (49%), with little presence of australasiatic (7%)
5.1 Rapa Nui 155
a b
c d
Fig. 5.2 Aspects of the botany of Isla de Pascua: a cover of the early review by F. Fuentes (1913);
b Doodia paschalis; c Sophora toromiro; d Axonopus paschalis (SGO collections)
(Doodia, Rytidosperma) and one antitropical genus (Agrostis) (Fig. 5.3). The extinct
palm Paschalococos disperta, known only from subfossil endocarps, is possible
related to Jubaea chilensis from the continent.
The natural history of the island was discussed by Skottsberg (1920–1956). He
emphasized the floristic relationships with the palaeotropics, and remarked that the
flora is so depauperate due to human influence, that it does not allow any further bio-
geographical conclusions regarding the origins of the island flora. Today the island’s
flora is dominated by alien species, but the presence of several endemic taxa like
Table 5.1 List of native plant genera and species of Rapa Nui
Floristic
Groups Family Genus N◦ species element
Ferns Aspleniaceae Asplenium 2 7

Blechnaceae Doodia 1 2
Davalliaceae Davallia 1 1
Dennstaedtiaceae Microlepia 1 1
Dryopteridaceae Diplazium 1 7
Dryopteris 1 7
Polystichum 1 7
Lomariopsidaceae Elaphoglossum 1 1
Ophiglossaceae Ophioglossum 2 7
Polypodiaceae Microsorum 1 1
Psilotaceae Psilotum 1 1
Thelypteridaceae Thelypteris 2 1
Vittariaceae Vittaria 1 1
Dicots Apiaceae Apium 1 7
Convolvulaceae Ipomoea 1 1
Fabaceae Sophora 1 7
Malvaceae Triumfetta 1 1
Samolaceae Samolus 1 7
Solanaceae Lycium 1 7
Solanum 1 7
Monocots Cyperaceae Kyllinga 1 1

Pycreus 1 1
Scirpus 1 7
Juncaceae Juncus 1 7
Poaceae Agrostis 1 4
Axonopus 1 1
Paspalum 1 7
Rytidosperma 1 2
Stipa 1 7
N°
Element genera
1. Pantropical 12 Cosmopolitan
2. Australasiatic 2 48% Pantropical
41%
3. Neotropical 0
4. Antitropical 1
5. South-temperate 0
6. Endemic 0
Antitropical Australasiatic
7. Cosmopolitan 14 4% 7%
Fig. 5.3 Floristic elements of Rapa Nui

5.2 Islas Desventuradas 157
“toromiro” (Sophora toromiro) and especially its archaeological and cultural rich-
ness, justify its status as a National Park since 1935 and the declaration as UNESCO
World Heritage Centre since 1995.
5.2 Islas Desventuradas

This relatively small volcanic archipelago is located approximately 850 km off
the Chilean coast (see Fig 1.1). The Desventuradas archipelago (i.e. “unfortunate
islands”) consists of two main islands: Isla San Félix (26◦ 17′ S, 80◦ 05′ W) and Isla
San Ambrosio (26◦ 21′ S, 79◦ 53′ W) (Fig. 5.4); plus several rocks and stacks: Islote
Gonzalez and Roca Catedral. Together, the Desventuradas Islands have a surface
area of only 10.3 km2 . The topography is very rugged, with peak elevations of 193
m asl on Isla San Félix, and 479 m asl on Isla San Ambrosio.
In spite of the relative sparse flora, these little islands have attracted natural-
ists due to their isolation and the presence of several interesting endemic taxa.
Botanical descriptions of the islands’ flora have been given by Philippi (1870, 1875),
Johnston (1935), Skottsberg (1937, 1951, 1963), Gunckel (1951), and Kuschel
(1962). More recent treatments are authored by Marticorena (1990) and Hoffmann
and Teillier (1991). According to these authors, the vascular flora of the islands
consists of 13 families, 18 genera and 25 native species, including several endemic
genera: Lycapsus and Thamnoseris (Asteraceae), Nesocaryum (Boraginaceae), and
Sanctambrosia (Caryophyllaceae). Teillier and Taylor (1997) add one native genus
to the list, Maireana (Amaranthaceae), with one species formerly known only
from Australia. Based on the works of these authors, a checklist with a total of
19 native genera pertaining to 13 families was compiled (Table 5.2). The best
represented family is the Amaranthaceae (4 genera formerly classified under the
Chenopodiaceae).
The floristic element that dominates in the islands is the cosmopolitan element
(52%), but the endemic element is also noteworthy, reaching a 21% (Fig. 5.5). This
Fig. 5.4 Islas Desventuradas: San Félix and San Ambrosio (reproduced with permission of the
Geographical Military Institute (IGM), Chile)
Table 5.2 List of native plant genera and species of Islas Desventuradas. Taxa endemic to the
archipelago in bold
Floristic
Groups Family Genus N◦ SP element
Dicots Aizoaceae Tetragonia 1 4

Amaranthaceae Atriplex 2 7
Chenopodium 1 7
Maireana 1 2
Suaeda 2 7
Asteraceae Lycapsus 1 6
Thamnoseris 1 6
Boraginaceae Nesocaryum 1 6
Brassicaceae Lepidium 1 7
Caryophyllaceae Sanctambrosia 1 6
Spergularia 1 7
Cucurbitaceae Sicyos 1 4
Frankeniaceae Frankenia 1 7
Malvaceae Cristaria 2 3
Fuertesimalva 1 3
Plantaginaceae/ Plantago 1 7
Veronicac.
Solanaceae Solanum 1 7
Urticaceae Parietaria 1 7
Monocots Poaceae Eragrostis 1 7
percentage is higher than the one for the continental flora or Juan Fernández at the
genus level. Furthermore, Marticorena (1990) reports a level of endemism of 60.6%
at the species level, the highest for a Chilean region.
The natural history of the archipelago has been analysed by Skottsberg (1937),
who argued that the Desventuradas flora shows a continental character rather than
an oceanic one. The existence of four endemic angiosperm genera and 20 endemic
species reinforces the view of an old floristic history not in accordance with rela-
tively recent migration events (discussion in Sect. 5.3). This view can be challenged
by the recent report of the genus Maireana, formerly known only from Australia
Element N° genera Australasiatic

5%
1. Pantropical 0 Neotropical
10%
2. Australasiatic 1
3. Neotropical 2
Cosmopolitan Antitropical
4. Antitropical 2 53% 11%
5. South-
temperate 0
6. Endemic 4
Endemic
21%
7. Cosmopolitan 10
Fig. 5.5 Floristic elements of Desventuradas Islands

5.3 Juan Fernández Archipelago 159
and hypothesized as the result of a recent long-distance dispersal event (Teillier and
Taylor 1997). The contemporary discovery of the genus on the coast of the Atacama
region (Marticorena 1997) tends to support this hypothesis, since it is unlikely
that such a shrub stayed unnoticed till the present. This could be one of the few
worldwide contemporaneous evidences for the effective operation of long-distance
dispersal (see Chap. 3).
5.3 Juan Fernández Archipelago

A prominent feature on the Nazca Plate is the Juan Fernández hot spot chain (see
Fig. 1.1). The Juan Fernández Ridge is a topographic swell crested by a series of
disconnected, large seamounts that first collided with the Chile margin in the north
at about 22 mya (Early Miocene) (Yáñez et al. 2001). The ridge has moved progres-
sively southwards to the current collision point located at roughly 32◦ –33◦ S. Today
the hot spot chain has two principal islands located between 667 and 850 km from
the continent (see Fig. 1.1):
– Alejandro Selkirk Island (33◦ 46′ S, 80◦ 47′ W) (also known as Isla Masafuera),
850 km from the American continent. Its highest elevation is Cerro Los Inocentes
(1,380 m asl) (Fig. 5.6a).
– Robinson Crusoe Island, (33◦ 38′ S, 78◦ 51′ W) (also known as Isla Masatierra),
located at around 667 km from the continent. Its highest peak is Cerro El Yunque
(915 m asl). Close to Robinson Crusoe there are two islets: Islote Juanango,
and Santa Clara, this latter located at 1 km southwest of Robinson Crusoe
(Fig. 5.6b).
The archipelago is worldwide known as having been the scenario for the real his-
tory that inspired the novel of Robinson Crusoe. The island Masafuera was renamed
in honour of the Scottish seaman Alejandro Selkirk, who survived 4 years and
4 months on Masatierra. Since its official discovery in 1574 by the pilot Juan
Fernández, Masatierra was an obligate anchor place for seamen and buccaneers
after trespassing Cape Horn, i.e. lost treasure histories are symptomatic on the
archipelago. But several botanists have long stated that the real treasure of this
archipelago lies in its unique plant world.
5.3.1 The Unique Plant World of Juan Fernández

The flora of the archipelago has been long a subject of interest for botanists (Gay
1832; Philippi 1856; Johow 1896 (Fig. 5.7), Skottsberg 1920–1956; Muñoz Pizarro
1969, Stuessy et al. 1984a, 1998b; Marticorena et al. 1998; Danton et al. 2000;
Danton and Perrier 2003). The native vascular flora comprises 203 species, 110
genera, and 55 families (Marticorena et al. 1998; Danton and Perrier 2006). The
best represented family are the Poaceae (10 genera) and the Asteraceae (9 gen-
era). The Fernandezian flora harbours two endemic families (Lactoridaceae and
160
a b
5
Fig. 5.6 Juan Fernández archipelago: a Isla Alejandro Selkirk (Masafuera); b Isla Robinson Crusoe (Masatierra) (from Skottsberg (1920–1956))
Pacific Offshore Chile
Fig. 5.7 a cover of Johow’s work; b illustration of (Juania australis) from the Flora de las Islas
de Juan Fernández
Thyrsopteridaceae) (see Sect. 2.2), 12 strictly endemic genera (plus 2 genera that
are endemic to continental Chile and the islands: Ochagavia and Notanthera)
(Table 5.3), and 132 endemic species (Marticorena et al. 1998) (Figs. 5.9 and 5.10).
The analysis of the floristic elements of the flora of Juan Fernández shows
that 14% are neotropical genera and 11% are australasiatic genera. Most impor-
tant is the cosmopolitan element (28%), with a somewhat smaller contribution of
the pantropical (17%), the endemic (14%) and the antitropical elements (12%)
(Fig. 5.8).
Floristic Element N° genera Cosmopolitan Pantropical

27% 17%
1. Pantropical 19
2. Australasiatic 12 Australasiatic
11%
3. Neotropical 16
4. Antitropical 13
5. South- Endemic
14% Neotropical
temperate 5 14%
6. Endemic 15 South-
temperate Antitropical
4% 12%
7. Cosmopolitan 30
Fig. 5.8 Floristic elements of Juan Fernández

Table 5.3 List of native plant genera and species of Juan Fernández Islands. Taxa endemic to the
archipelago in bold
Floristic
Ferns Aspleniaceae Asplenium 4 7

Blechnaceae Blechnum 7 7
Dennstaedtiaceae Histiopteris 1 1
Hypolepis 1 1
Dicksoniaceae Dicksonia 2 2
Lophosoria 1 3
Dryopteridaceae Elaphoglossum 1 1
Megalastrum 1 1
Polystichum 1 7
Rumohra 1 2
Gleicheniaceae Sticherus 3 1
Hymenophyllaceae Hymenoglossum 1 6
Hymenophyllum 10 7
Serpyllopsis 1 5
Trichomanes 3 1
Lycopodiaceae Lycopodium 2 7
Ophioglossaceae Ophioglossum 1 7
Polypodiaceae Grammitis 1 1
Pleopeltis 2 1
Synammia 1 5
Pteridaceae Adiantum 1 7
Notholaena 1 1
Pteris 3 7
Tectarinaceae Arthropteris 1 2
Thyrsopteridaceae Thyrsopteris 1 6
Woodsiaceae Cystopteris 1 4
Dicots Amaranthaceae Chenopodium 3 7
Sarcocornia 1 7
Apiaceae Apium 2 7
Centella 1 1
Eryngium 4 7
Asteraceae Abrotanella 1 2
Centaurodendron 2 6
Dendroseris 11 6
Erigeron 5 4
Gamochaeta 1 3
Lagenophora 1 2
Robinsonia 8 6
Taraxacum 2 4
Yunquea 1 6
Berberidaceae Berberis 2 4
Boraginaceae Selkirkia 1 6
Brassicaceae Cardamine 3 4
Campanulaceae Lobelia 1 1
Wahlenbergia 5 4
Caryophyllaceae Spergularia 2 7
Convolvulaceae Calystegia 1 7
Dichondra 1 1
Table 5.3 (continued)
Floristic
Ericaceae Empetrum 1 4
Gaultheria 1 4
Escalloniaceae Escallonia 1 3
Euphorbiaceae Dysopsis 1 3
Fabaceae Sophora 2 7
Gunneraceae Gunnera 4 1
Haloragaceae Haloragis 2 2
Lactoridaceae Lactoris 1 6
Lamiaceae Cuminia 1 6
Loranthaceae Notanthera 1 6
Myrtaceae Myrceugenia 2 3
Myrteola 1 3
Ugni 1 3
Orobanchaceae Euphrasia 1 4
Phrymaceae Mimulus 1 7
Piperaceae Peperomia 4 1
Ranunculaceae Ranunculus 1 7
Rhamnaceae Colletia 1 3
Rosaceae Acaena 1 4
Margyracaena 1 6
Margyricarpus 1 3
Rubus 1 7
Rubiaceae Coprosma 2 2
Galium 1 7
Oldenlandia 1 1
Nertera 1 2
Rutaceae Fagara 2 1
Salicaceae Azara 1 3
Santalaceae Santalum 1 2
Solanaceae Nicotiana 1 1
Solanum 2 7
Urticaceae Boehmeria 1 1
Parietaria 1 7
Urtica 3 7
Verbenaceae Rhaphithamnus 1 5
Plantaginaceae/ Plantago 3 7
Veronicac.
Winteraceae Drimys 1 3
Monocots Arecaceae Juania 1 6

Bromeliaceae Greigia 1 3
Ochagavia 1 6
Cyperaceae Carex 2 7
Cyperus 2 7
Eleocharis 1 7
Machaerina 1 1
Oreobolus 1 2
Scirpus 2 7
Uncinia 4 2
Floristic
Iridaceae Herbertia 1 3
Libertia 1 2
Juncaceae Juncus 5 7
Luzula 1 7
Orchidaceae Gavilea 1 5
Poaceae Agrostis 1 4
Bromus 1 4
Chusquea 1 3
Danthonia 2 4
Leptophyllochloa 1 5
Megalachne 2 6
Nassella 2 3
Piptochaetium 1 3
Podophorus 1 6
Trisetum 1 7
a b
Fig. 5.9 Juan Fernández

ferns: a Arthropteris
altescandens; b Dicksonia
berteroana; c Thyrsopteris
elegans (photo credits: a–c
S. Elórtegui Francioli) c
a b
c d e
Fig. 5.10 Juan Fernández angiosperms: a Lactoris fernandeziana; b Sophora fernandeziana;

c Rhaphithamnus venustus; d Escallonia callcottiae; e Gunnera peltata (photo credits: a–e S.
Elórtegui Francioli)
5.3.2 Floristic Similarity of Juan Fernández

Including Juan Fernández (JF) into the similarity analysis done between Chilean
regions (Sect. 4.2) results in a closer floristic relationship between JF/MAG than
between JF and the other Chilean regions (Table 5.4, Fig. 5.11). JF shares more
native genera with BIO (81) but Jaccard’s similarity is higher between JF/MAG,
taking account of the much richer generic flora of BIO as compared to MAG or JF.
In Fig. 5.11, geographic distance is plotted against Jaccard’s distance. Two
expected trends are recovered, one for the continental regions, and the decreasing
similarity between JF and each of the continental regions according to their geo-
Table 5.4 Floristic similarity between regions including Juan Fernández
Compared regions Similarity Jaccard Distance (km) Shared genera
ANT/COQ 0.46 777 244

ANT/BIO 0.29 1,527 174
ANT/MAG 0.21 3,227 97
COQ/BIO 0.54 750 323
COQ/MAG 0.27 2,450 150
BIO/MAG 0.33 1,700 178
JF/MAG 0.18 2,261 56
JF/BIO 0.16 792 81
JF/COQ 0.12 895 64
JF/ANT 0.08 1,439 33
1.0
0.8
Jaccard similarity
0.6
COQ/BIO
ANT/COQ
0.4
BIO/MAG
ANT/BIO COQ/MAG
0.2
JF/BIO JF/MAG ANT/MAG

JF/COQ
JF/ANT
0.0
0 2000 4000 6000

Distance [km]
Fig. 5.11 Floristic similarity between different regions in Chile: Antofagasta (ANT), Biobío
(BIO), Coquimbo (COQ) and Magallanes (MAG). Jaccard similarity is plotted against geograph-
ical distance within Chile (white squares), and between Juan Fernández and Chilean continental
regions (black squares). The lines represent trends as exponential curves. The relation JF/MAG is
considered as an outlier. The arrow and question mark show the possible position of the relation
JF/MAG following the trend (see discussion in the text)
graphic distances. But the relationship MAG/JF is noteworthy and escapes from this
trend (outlier). According to geographic distance, these two regions should have a
much lower similarity (see arrow). This closer floristic relationship between MAG
and JF appeals to two different explanations:
(a) The Magallanes biota had once a more northward distribution, till central Chile,
and from there it reached the islands via long-distance dispersal. The presence
of south-temperate elements in the Fray Jorge fog forest at 30◦ 40′ long has
been suggested as evidence of the more widespread occurrence of temperate
forests during the Cenozoic (Box 3.1). But the remnants of these forests are
still represented along the coast of BIO and there is no apparent reason why the
relation BIO/JF stays within the trend of the similarity/distance curve, contrary
to MAG/JF.
(b) An alternative explanation for the floristic similarity between JF and MAG is a
direct connection of the two land masses. This explanation is contrary to current
geological scenarios, since the islands seem to be geologically too young for
this type of explanation. The islands are located on top of the Juan Fernández
ridge, which controls the tectonic and geological evolution of the southern
Andes at 33◦ –34◦ S since the Tertiary (Yáñez et al. 2002). The seafloor age
assigned by Müller et al. (1997) to the seafloor offshore Chile between 18◦ and
40◦ S, based on magnetic anomalies and relative plate reconstructions, ranges
from 20 to 48 mya. Juan Fernández rests on seafloor dated at around 20–33 mya.
Several early geologists believed in a former land west of South America occupy-
ing the whole Pacific (Burckhardt 1902; Beloussov 1968), or at least a portion of it
as a Transandiner Kontinent close to the South American coast (Illies 1967; Cañas
1966). Brüggen (1950, p 59) proposed the name Tierra de Juan Fernández for this
larger continental land mass west of current South America. Miller (1970) anal-
ysed different possibilities for the disappearance of this land and concluded that the
Ozeanisierung of the Juan Fernández Land occurred in the Late Cenozoic, and that
this land was not at all at the same location of today’s Juan Fernández archipelago
(Miller 1970, p 934). Based on the floristic similarities as analysed in the present
work, this land should have existed much more to the South. To fit in the trend
line of the similarity/distance relation shown by other Chilean regions, the Juan
Fernández islands (Juan Fernández Land) could be located at the same longitude
(80◦ W), but at 48◦ S instead of 33◦ , i.e. 15 degrees latitude or around 1,650 km
towards the South. (Fig. 5.12, see also Sect. 10.4).
Biogeographic connections between Juan Fernández, Magallanes and the
Falkland (Malvinas) Islands have been also identified by Morrone (1992), as
a generalized track composed by plants as well as insects, Crustaceans, and
Oligochaeta.
Fig. 5.12 Hypothetical

closer geographic position of
Juan Fernández to southern
South America (Magallanes),
according to floristic
similarity in Fig. 5.11 (grey
dotted line); and possible
position of the Juan
Fernández Land or the
Pacifica continent, according
to Dickins et al. (1992) (black
dotted line)
5.3.3 Origins of the Fernandezian Flora

The origin of the Fernandezian flora has been the subject of considerable study
and debate (e.g. Skottsberg 1925, 1936, 1956; van Balgooy 1971). Traditionally the
origin of an island flora has been explained in direct relationship with its nearest
continental mass by means of long-distance dispersal (e.g. Carlquist 1974). Oceanic
islands are often explained as geologically new territories and Juan Fernández is not
an exception: Masatierra has been dated at ca. 4 mya, Masafuera at 1–2 mya, and
Isla Santa Clara at 5.8 mya (Stuessy et al. 1984b). The islands are supposed to be the
products of isolated intraplate volcanism associated with the volcanic hotspot (Stern
et al. 2007). The relative young age of the archipelago seems to leave no doubt for
a recent origin of its flora, as revised by Bernardello et al. (2006).
But the high level of endemism, the variety of floristic relationships and the lim-
ited methods of dispersal put some unresolved questions on this theme. Bernardello
et al. (2006) report that 80% of the island’s species have dry fruits, and fleshy fruits
are comparatively uncommon, challenging the supposed ability for long-distance
dispersal by birds. In fact, the dispersal syndrome that prevails in the flora is
autochory (i.e. autonomous passive dispersal). Bernardello et al. (2006) therefore
suggest that the principal dispersal processes are anemochorous dispersal (air flota-
tion) and epizoochory (carried by birds attached to their feathers). But since the
extant bird fauna is scarce, the real opportunities for dispersal are relatively few.
This was early recognized by Skottsberg (1925, 1956) and he therefore suggested
that the origin of the island flora should be found in alternative palaeoscenarios.
Skottsberg revised and discussed all available evidence in floristic and faunistic ele-
ments, in geotectonics of the Pacific, and in the continental Cenozoic flora and
came to the conclusion that the Fernandezian flora is not of an oceanic but of
a continental nature. This is in conflict with the early classification of Wallace
(1880) that considered all oceanic islands as having an oceanic biota (see also
Cowie and Holland 2006). Skottsberg proposed a “tentative sketch” on the his-
tory of the Fernandezian flora that is in concordance with Brüggen’s Tierra de
Juan Fernández, an old submerged landmass west of today’s South America that
could have been the source for the unique Fernandezian flora (Brüggen 1950;
Skottsberg 1956, p 394) (Fig. 5.12).
Skottsberg’s critical vision has been systematically oversimplified by modern
authors, while emphasizing that he noted the close relationship with the American
flora (e.g. Crawford et al. 1990; Bernardello et al. 2006). Skottsberg certainly rec-
ognized the floristic relationship between the islands and the American continent,
but he also noted the closest relationship with the western Pacific and especially
with Australasia. Indeed, several genera not found in continental Chile show a
wider distribution in Australasia, such as Coprosma (Rubiaceae), Arthropteris
(Tectarinaceae) (Fig. 5.9), Haloragis (Haloragaceae) or Santalum (Santalaceae).
“In traditional dispersalist interpretations of oceanic island biotas, a volcanic
island just ‘pops up’ and its location is not considered relevant. Islands are assumed
to be populated from the nearest mainland and endemism is explained by isola-
tion from the mainland However, since volcanism recurs in certain areas, volcanic
islands might instead ‘inherit’ flora and fauna from prior volcanic islands in the
same region” (Heads 2009, p 236). Heads (2009) exemplifies his proposal with
Fitchia (Asteraceae) a small tree endemic to montane forests in SE Polynesia.
The genus has “presumably survived as a metapopulation on the volcanic islands
and atolls which have come and gone around the Cook Islands/Tokelau and other
localities in SE Polynesia. Its ancestors may date back to the origin of the Pacific
plate and the Cretaceous plateaus” (Heads 2009, p 236). A similar history can be
hypothesized for the small trees Robinsonia, Dendroseris, Yunquea, endemic to Juan
Fernández, and Thamnoseris and Lycapsus endemic to the Desventuradas Islands.
These Asteraceous genera could be the remnants of a more “ancient Asteraceous
world” related to the origins of the Pacific plate.
The sole presence of the endemic family Lactoridaceae with its single species
puts a big question mark on the supposed oceanic character of the Fernandezian
flora. Lactoris has been the subject of many studies because of its systematic
placement among basal angiosperms (e.g. Stuessy et al. 1998a; González and
Rudall 2001). The cladistic analysis by Lammers et al. (1986) suggests that the
Lactoridaceae diverged sometime prior to the Maastrichtian (69 mya). This has been
corroborated recently by the analysis of Wikström et al. (2001): Lactoris appears
as a very ancient taxon at the base of the angiosperm phylogenetic tree: the split
between Lactoris and Aristolochia has been dated at around 85 mya.
In Lammers et al. (1986) opinion “it seems unlikely that the Lactoridaceae
evolved autochthonously in the Juan Fernandez Islands. A more plausible hypoth-
esis is that the plants on Masatierra are relicts of a once more extensive conti-
nental distribution in South America and possibly other portions of the Southern
Hemisphere, perhaps originating from the western Gondwanaland flora”. Indeed,
microfossils related to Lactoris (Fig. 5.13) and referred to the fossil genus
Lactoripollenites have been found in South Africa and Australia (Macphail et al.
1999), thus suggesting that the Lactoridaceae were widespread across the Southern
Hemisphere during the Late Cretaceous (Lammers et al. 1986). “Differences
between Lactoripollenites and Lactoris pollen imply that these represent different
clades within the Lactoridaceae or that the former evolved into the latter genus else-
where in the Southwest Pacific region prior to its migration onto Masatierra in the
Plio-Pleistocene” (Macphail et al. 1999).
Recently, a new fossil find of lactoridaceous tetrads from the early Miocene of
eastern Patagonia has been reported by Gamerro and Barreda (2008). These authors
compiled all the fossil evidence for the Lactoridaceae concluding that it probably
has a South African origin, with the oldest records in the Cretaceous (Turonian–
Campanian), reaching its widest palaeogeographical distribution (Australia, India,
Antarctica, and North and South America) by the Maastrichtian. According to this
new finding “the family may have migrated into South America, either via Africa
(through the Atlantic Ocean) or Antarctica, by the Maastrichtian, growing in eastern
Patagonia up to the early Miocene. Arid conditions established in this region by the
middle–late Miocene onwards would have determined the restriction of forests to
the western lands. The Lactoridaceae may have followed a similar migration pat-
tern towards the Pacific coast of South America” (Gamerro and Barreda 2008). The
Fig. 5.13 Lactoris fernandeziana, as illustrated in Skottsberg (1920–1956): a top of branch in

August; b terminal inflorescence; c female flower; d hermaphrodite flower. See also Fig. 5.10
last part of its history is already known: “The shifting of Lactoridaceae towards
Masatierra Island would have occurred during the last 4 my by long-distance
dispersal events (perhaps by birds)” (Gamerro and Barreda 2008).
Crawford et al. (2001) also suggested that the species or its ancestors could have
reached Masatierra by means of long-distance dispersal, but may have been car-
ried by the wind, due to its small seeds. However, the authors recognized that “the
plants occur primarily in the forest understory, which would seemingly minimize
the effectiveness of wind as a means of long-distance dispersal” (Crawford et al.
2001, p 189).
Other endemic taxa might be also the remnants of older groups that have evolved
in a completely different palaeogeographic scenario. As an example, the fern species
Thyrsopteris elegans is the sole representative of the Thyrsopteridaceae (Smith et al.
2006) (Sect. 2.2). This fern is clearly related to the tree ferns, but of uncertain posi-
tion within the group (Korall et al. 2006). Fossil representatives of the genus (or
related taxa) are widespread; its distributional restriction to Juan Fernández is obvi-
ously a relict situation (Kramer and Green 1990). Also the shrubby Fernandezian
Wahlenbergia species, together with the species from New Zealand and St. Helena,
are considered as the more basal members of the wahlenbergioid group, suggesting
a Gondwanic origin (Eddie et al. 2003).
The splitting between Drimys confertifolia from the continental sister species
(clade D. andina + D. winteri), has been dated at 9–10.8 mya by the phylogenetic
analyses from Marquínez et al. (2009). These authors are well aware of the fact that
this age notably predates the formation of the Juan Fernández islands.
Stuessy et al. (1984b) knew the earlier view from Brüggen (1950) and Skottsberg
(1956), but they gave much value to the Potassium-Argon dating. Thus, most of the
papers dealing with the evolution of the Fernandezian flora in the last two decades
start from the 4 mya date. The dating of Santa Clara puts another question on the
problem, since this little islet seems to be almost two million years (5.8 +-2.1 mya)
a b c
Fig. 5.14 Reconstruction of the geomorphological history of Masatierra + Santa Clara: a putative
emerged shape of the islands at 4 mya; b erosional patterns showing amphitheater valleys; c present
configuration showing bathymetric contours (adapted from Stuessy et al. 1984a)
older than Robinson Crusoe. Stuessy et al. (1984b), in spite of their high confidence
in the dating, do not rule out other models of Pacific aseismic ridges (e.g. Nur and
Ben-Avraham 1981). Also, Stuessy et al. (1984a) recognized that the islands could
have been much more extended and may have been rapidly eroded during the last
4 million years (Fig. 5.14). All Chilean Pacific islands show indeed advanced ero-
sion stages, related to volcanic sector-collapse processes (Araya-Vergara and Vieira
2005; Stern et al. 2007).
Thus, advancing definitive conclusions on this topic can be very risky; biogeo-
graphical islands’ enigmas are not exclusive to Juan Fernández: the moss species
Hedenasiastrum percurrens, which is endemic to the Madeiran archipelago, most
probably diverged about 40 mya. But the emergence of Madeira has been dated at
only 5.2 mya (Aigoin et al. 2009).
5.3.4 Conservation of Juan Fernández Plants

The archipelago is one of the most dramatically threatened territories worldwide
(Stuessy et al. 1998c), an actual tendency on oceanic islands (Sax and Gaines 2008).
The combined introduction of browsing animals and highly invading continental
plants place the fernandezian native flora at a competitive disadvantage (Dirnböck
et al. 2003). Today at least 75% of the endemic flora is highly threatened (Swenson
et al. 1997; Cuevas and van Leersum 2001). Danton and Perrier (2005, 2006) listed
eight species that went extinct (EX) in historical times, mostly during the last
decades: Podophorus bromoides, Santalum fernandezianum, Chenopodium neso-
dendron, Empetrum rubrum, Eryngium sarcophyllum, Notanthera heterophylla,
Robinsonia berteroi, and R. macrocephala. Applying IUCN criteria, the authors
further classified one more species as “Extinct in the wild” (Margyracaena skotts-
bergii, still conserved ex-situ at the National Botanical Garden in Viña del
Mar). Another 28 species were classified as “Critically endangered” (CR), 79 as
“Endangered” (EN), and 65 as “Vulnerable” (VU). Finally, 32 species were classi-
fied as “Least Concern” (LC) or “Data Deficient” (DD). That means that from the
a
20
m Canelo (Drimys comfertifolia) Pteris berteroana
Luma (Myrceugenia fernandeziana) Dysopsis hirsuta
Dicksonia berteroana Thyrsopteris elegans
Blechnum schottii
15
Rumohra berteroana
10
5
Blechnum cordatum
Polystichum tetragonum Hymenophyllum fuciforme
Blechnum cycadifolium Robinsonia evenia
0
0 5 10 15 m 20
b
20
m Canelo (Drimys confertifolia)
Luma (Myrceugenia fernandeziana)
Arthropteris altescandens
Maqui (Aristotelia chilensis)
15
Pteris berteroana
Rubus ulmifolius
Steine
10
0
0 5 10 15 m 20
Fig. 5.15 Natural v/s invaded vegetation stands in Masatierra: a lower montane forest near
Plazoleta del Yunque; b native Drimys confertifolia and Myrceugenia fernandeziana oppressed
by the continental aliens Aristotelia chilensis and Rubus ulmifolius (original illustrations by Rudi
Rössler (Erlangen University) based on schemes by Prof. Michael Richter, December 2005)
213 native species evaluated, nine are already extinct in the archipelago and 172
(80%)! are critically endangered to vulnerable.
The native flora is being directly attacked by invading animals (e.g. rabbits, rats)
and plants, and the alien flora is continuously expanding: Johow (1896) listed 95
introduced species, whereas Marticorena et al. (1998) listed a total of 212 natu-
ralized species in the archipelago (against 211 native). This number is increasingly
growing, and species that are presently considered naturalized and cultivated (poten-
tially invaders), bring the number at 503 species (Danton and Perrier 2006). That
means that the alien flora is already double as large as the native flora!
There have been continuous efforts to eradicate the plant and animal pests (rab-
bits, goats, rats), but they still do not show much success (Cuevas and van Leersum
2001). Even after cattle exclusion, vegetation recovery seems to be very slow
(Cuevas and Le Quesne 2006). Presently the high number of alien species does
not seem to be the worst problem: most effective invaders are several continen-
tal natives like “maqui” (Aristotelia chilensis) and “murtilla” (Ugni molinae), plus
the European weed “zarzamora” (Rubus ulmifolius) (Fig. 5.15b). These three alien
species are virtually replacing the native vegetation on Masatierra. Modelling the
invasion’s extent for these highly invasive species since their introduction 80 years
ago, suggests that 50% of the remaining montane forest could be totally replaced in
just another 80 years. . . (Greimler et al. 2002; Dirnböck et al. 2003).
Even of the iconic plant Lactoris fernandeziana, the last survey could find no
more than 1,000 plants (Ricci 2001). Conservation efforts in situ as well as ex situ
at the National Botanical Garden in Viña del Mar continue, but most botanists are
pragmatically pessimistic about the future of this unique flora, if definite actions
are not taken. The great botanist Carl Skottsberg was very impressed to see the last
exemplar of the species Santalum fernandezianum (Box 5.1, Fig. 5.16)
Box 5.1 Skottsberg’s Impressions on the Encounter

with the Last Exemplar of the Extinct Santalum
fernandezianum
“The discovery of the sandal-wood, famous
since the days of Solomon, on Juan Fernández
most surely attracted notice. We have no reports
of it previous to 1624, when, according to
Burney, L’Heremite reported sandal-trees in
great number. According to another authority
ships used to visit the place as early as 1664
to bring the valuable wood to the coast, where
it was highly appreciated. One did not think
of preserving anything; a hundred years later it
was hardly possible to find a living tree, and in
the beginning of last century it was regarded as
extinct. No botanist had ever seen the leaves or flowers. Suddenly F. Philippi
in Santiago got some fresh twigs brought to him in 1888; he found them to
belong to the genus Santalum; the species being new, it received the name of
S. fernandezianum. The general interest in the tree was increased, but nobody
told where the branches came from; a living tree was still unknown. Only in
1892 did Johow get news of one; a colonist had found it in Puerto Inglés, high
up in the valley. He was the first botanist who saw this plant. It is easily under-
stood that I was anxious to become the second. How many people had looked
for other specimens! All their efforts were fruitless; as far as we knew Johow’s
tree was the very last. If it were still there! The man who brought Johow to
the spot still lived, and after we had explained our purely scientific interest he
promised to send his son with us. It would have been more than uncertain for
us alone to look for a single tree in a valley clad with virgin forest. [. . .] We
walked up the valley and made an ascent of the western side; the place is so
steep that one is forced to grasp the trees and shrubs to get a foothold. Our
guide stopped, looked round for a minute, down a few hundred yards, and we
had reached our destination. The last sandal-tree. Absolutely the last descen-
dant of Santalum fernandezianum. It is so queer to stand at the death-bed of a
species; probably we were the last scientists who saw it living. We look at the
old tree with a religious respect, touch the stem and the firm, dark green leaves
– it is not only an individual, it is a species that is dying. It cannot last very
long. There is only one little branch left fresh and green; the others are dead.
We cut a piece to get specimens of the peculiar red, strongly scented wood. A
photo was taken, I made some observations on the place, and we said good-
bye. Should I happen to go there once more I shall not see the sandal-tree; it
will be dead and its body cut up into precious pieces – curiosities taken away
by every stranger” (Skottsberg 1911).
Fig. 5.16 Herbarium exemplar of Santalum fernandezianum from SGO

b d
Fig. 5.17 Juan Fernández landscapes: a view towards the west from Mirador de Selkirk; b Puerto
Inglés; c volcanic scarps seen from the sea; d intensive erosion towards Puerto Francés (photo
credits: (a–d) S. Elórtegui Francioli)
With the support of several botanists like Carl Skottsberg and the Chilean state,
Juan Fernández has the status of a National Park since 1935, and it has been declared
also as a Biosphere Reserve in 1977 (see Sect. 6.1). These formal protection statuses
so far did not succeed to prevent the extinction process that is currently affecting the
Fernandezian flora. There is still much work to be done on the control of pests
and invasive species, in erosion control and ecological restoration. Encouraged staff
and personal efforts done for decades by the personal from CONAF (Corporación
Nacional Forestal) could not stop the threats. Personal engagement exists, but it
must be supported by national and international investment in conservation and
restoration. If international conservation agencies are looking for highly threatened
territories to invest their efforts, here we have a territory that has too long been
waiting for a coordinated action to definitely stop the natural depletion and floral
extinctions. “If we do not stop the invaders now, the saddest aspects of Robinson
Crusoe’s parable will succeed: nature will have been definitely subjugated by man,
transforming a marvelous floristic and evolutionary space into a human-induced
desert” (Danton and Perrier 2006, p 420) (Fig. 5.17).
References
Aigoin DA, Devos N, Huttunen S, Ignatov MS, Gonzalez-Mancebo JM, Vanderpoorten A (2009)
And if Engler was not completely wrong? Evidence for multiple evolutionary origins in the
moss flora of Macaronesia. Evolution 63:3248–3257
Arancibia V, (ed) (2009) Rapa Nui: iorana te ma’ohi: dilemas estratégicos. Eds. Universidad
Católica de Chile, Santiago
Araya-Vergara JF, Vieira R (2005) Formas costeras y submarinas de destrucción en los sistemas
volcánicos San Félix y Robinson Crusoe, Pacífico Suroriental. Ciencia y Tecnología del Mar
28:5–23
Balcells I (2005) El Chile que no está en América. In: Badal G, Balcells F, Montanari M, et al.
(eds) Chile: País Oceánico (2nd edn). Ocho Libros Editores, Santiago, pp 96–97
Beloussov VV (1968) Some general aspects of development of the Tectonosphere. Rep. 28. Int
Geol Congr 1:9–17
Bernardello G, Anderson GJ, Stuessy TF, Crawford DJ (2006) The angiosperm flora of the
Archipelago Juan Fernández (Chile): origin and dispersal. Can J Bot 84:1266–1281
Bologna M, Flores JC (2008) A simple mathematical model of society collapse applied to Easter
Island. EPL 81: 48006 (4 pp) doi: 10.1209/0295–5075/81/48006
Bork H-R (2006) Landschaften der Erde unter dem Einfluss des Menschen. Wissenschaftliche
Buchgesellschaft (WGB), Darmstadt
Brüggen J (1950) Fundamentos de la Geología de Chile. Instituto Geográfico Militar, Santiago
Burckhardt C (1902) Traces géologiques d’un ancient continent Pacifique. Rev. Mus. La Plata
10:179–192
Cañas J (1966) Tectónica y sedimentación de la Formación Los Molles, Prov. de Aconcagua.
Comun. Escuela De Geol. (Santiago) 11:22
Carlquist S (1974) Island biology. Columbia University Press, New York, NY
Cowie RH, Holland BS (2006) Dispersal is fundamental to biogeography and the evolution of
biodiversity on oceanic islands. J Biogeogr 33:193–198
Crawford DJ, Stuessy TF, Lammers TG, Silva M, Pacheco P (1990) Allozyme variation and evo-
lutionary relationships among three species of Wahlenbergia (Campanulaceae) in the Juan
Fernandez Islands. Bot Gaz 151:119–124
References 177
Crawford D, Tago-Nakazawa M, Stuessy TF, Anderson GJ, Bernardello G, Ruiz E, Jensen RJ,
Baeza CM, Wolfe AD, Silva M (2001) Intersimple sequence repeat (ISSR) variation in Lactoris
fernandeziana (Lactoridaceae), a rare endemic of the Juan Fernández Archipelago, Chile. Plant
Spec Biol 16:185–192
Cuevas JG, Le Quesne C (2006) Low vegetation recovery after short-term cattle exclusion on
Robinson Crusoe Island. Plant Ecol 183:105–124
Cuevas JG, van Leersum G (2001) Proyecto conservación, restauración y desarrollo de las islas
Juan Fernández, Chile. Rev Chil Hist Nat 74:899–910
Danton P, Breteau E, Baffray M (2000) Les îles de Robinson: Trésor vivant des mers du Sud, entre
légende et réalité. Ed. Nathan.
Danton P, Perrier C (2003) Liste de la flore vasculaire de l’île Robinson Crusoe, archipel Juan
Fernández, Chili. J Bot Soc Bot France 24:67–78
Danton P, Perrier C (2005) Notes sur la disparition d’une espèce emblématique: Robinsonia bert-
eroi (DC.) Sanders, Stuessy & Martic. (Asteraceae), dans l’île Robinson Crusoe, archipel Juan
Fernández (Chili). J Bot Soc Bot France 31:3–8
Danton P, Perrier C (2006) Nouveau catalogue de la flore vasculaire de l’archipel Juan Fernández
(Chili). Acta Bot Gall 153:399–587
Dickins JM, Choi DR, Yeates AT (1992) Past distribution of oceans and continents. In:
Chatterjee S, Hotton N III, (eds) New concepts in global tectonics. Texas Tech. University
Press, Lubbock, TX, pp. 193–199
Dirnböck T, Greimler J, Lopez P, Stuessy TF (2003) Predicting future threats to the native
vegetation of Robinson Crusoe Island, Juan Fernandez Archipelago, Chile. Conserv Biol
17:1650–1659
Eddie WMM, Shulkina T, Gaskin J, Haberle RC, Jansen RK (2003) Phylogeny of Campanulaceae
s. str. inferred from ITS sequences of nuclear ribosomal DNA. Ann Mo Bot Gard 90:
554–575
Fuentes F (1913) Reseña botánica sobre la Isla de Pascua. Bol Mus Nac Hist Nat 5:320–337
Gamerro JC, Barreda V (2008) New fossil record of lactoridaceae in southern South America: a
palaeobiogeographical approach. Bot J Linn Soc 158:41–50
Gay C (1832) La Isla de Juan Fernández. Reprint 1974. Mus Nac Hist Nat (Santiago) Ser Educativa
9:5–16
González F, Rudall PJ (2001) The questionable affinities of Lactoris: evidence from
branching pattern, inflorescence morphology and stipule development. Am J Bot 88:
2143–2150
Greimler J, López P, Stuessy TF, Dirnböck T (2002) The vegetation of Robinson Crusoe Island
(Isla Masatierra), Juan Fernández Archipelago, Chile. Pac Sci 56:263–284
Gunckel H (1951) Algunas plantas vasculares de la Isla de San Ambrosio. Rev Univ (Santiago)
36:291–314
Heads M (2009) Globally basal centres of endemism: the Tasman-Coral Sea region (south-west
Pacific), Latin America and Madagascar/South Africa. Biol J Linn Soc 96:222–245
Hoffmann AJ, Teillier S (1991) La flora de la Isla San Félix (Archipiélago de las Desventuradas,
Chile). Gayana Bot 48:89–99
IGM (2005) Atlas de Chile. Instituto Geográfico Militar, Santiago
Illies H (1967) Randpazifische Tektonik und Vulkanismus im südlichen Chile. Geol Rundsch
57:81–101
Johnston IM (1935) The flora of San Félix Island. J Arnold Arbor 16:440–447
Johow F (1896) Estudios sobre la flora de las Islas de Juan Fernández. Imprenta Cervantes,
Santiago
Korall P, Pryer KA, Metzgar JS, Schneider H, Conant DS (2006) Tree ferns: monophyletic groups
and their relationships as revealed by four protein-coding plastid loci. Mol Phylogenet Evol
39:830–845
Kramer KU, Green PS (1990) Pteridophytes and gymnosperms. In: Kubitzki K, (ed) The families
and genera of vascular plants, vol 1. Springer, Berlin
Kuschel G (1962) Zur Naturgeschichte der Insel San Ambrosio (Islas Desventuradas, Chile). 1.
Reisebericht, geographische Verhältnisse und Pfalzenverbreitung. Arkiv Bot 4:413–419
Lammers TG, Stuessy TF, Silva M (1986) Systematic relationships of the Lactoridaceae, an
endemic family of the Juan Fernandez Islands, Chile. Plant Syst Evol 152:243–266
Macphail MK, Partridge AD, Truswell EM (1999) Fossil pollen records of the problematical
primitive angiosperm family Lactoridaceae in Australia. Plant Syst Evol 214:199–210
Mann D, James E, Chase J, Beck W, Reanier R, Mass M, Finney B, Loret J (2008) Drought,
vegetation change, and human history on Rapa Nui (Isla de Pascua, Easter Island). Quat Res
69:16–28
Marquínez X, Lohmann LG, Faria Salatino ML, Salatino A, González F (2009) Generic relation-
ships and dating of lineages in Winteraceae based on nuclear (ITS) and plastid (rpS16 and
psbA-trnH) sequence data. Mol Phylogenet Evol 53:435–449
Marticorena C (1990) Contribución a la estadística de la flora vascular de Chile. Gayana Bot 47:
85–113
Marticorena C (1997) La presencia de Maireana brevifolia (R.Br.) Paul G. Wilson
(Chenopodiaceae), en Chile continental. Gayana Bot 54:193–194
Marticorena C, Stuessy TF, Baeza CM (1998) Catalogue of the vascular flora of the Robinson
Crusoe or Juan Fernández Islands, Chile. Gayana Bot 55:187–211
Miller H (1970) Das problem des hypothetischen „Pazifischen Kontinentes“ gesehen von der
chilenischen Pazifikküste. Int J Earth Sci 59:927–938
Morrone JJ (1992) Revisión sistemática, análisis cladístico y biogeografía histórica de los géneros
Falklandius Enderlein y Lanteriella gen. Nov. (Coleoptera: Curculionidae). Acta Entomol Chil
17:157–174
Müller RD, Roest WR, Royer J-Y, Gahagan LM, Sclater JG (1997) Digital isochrones of the
world’s ocean floor. J Geophys Res 102:3211–3214
Muñoz Pizarro C (1969) El Archipiélago de Juan Fernández y la conservación de sus recursos
naturales renovables. Reprint 1974. Mus Nac Hist Nat (Santiago) Ser Educativa 9:17–47
Nur A, Ben-Avraham Z (1981) Volcanic gaps and the consumption of aseismic ridges in South
America. In: Kulm LVD, Dymond J, Dasch EJ et al. (eds) Nazca Plate: Crustal Formation and
Andean Convergence. Geol Soc Am Mem 154:729–740
Philippi RA (1856) Observaciones sobre la flora de Juan Fernández. An Univ Chile 13:157–169
Philippi RA (1870) Vegetation der Inseln San Ambrosio und S. Felix. Bot. Zeitung (Berlin)
28:496–502
Philippi F (1875) La flora de las islas San Ambrosio i San Félix. An Univ Chile 47:185–194
Ricci M (2001) Evaluation of conservation status of Lactoris fernandeziana Philippi
(Lactoridaceae) in Chile. Biodivers Conserv 10:2129–2138
Sax DF, Gaines SD (2008) Species invasions and extinction: the future of native biodiversity on
islands. Proc Natl Acad Sci Usa 105:11490–11497
Skottsberg C (1911) The wilds of Patagonia: a narrative of the Swedish expedition to Patagonia,
Tierra del Fuego and the Falkland Islands in 1907–1909. Macmillan, Edward Arnold, NY,
Londres
Skottsberg C, (ed) (1920–1956) Natural history of Juan Fernandez and Easter Island, 3 vol.
Almqvist & Wiksell, Uppsala
Skottsberg C (1925) Juan Fernandez and Hawaii. A phytogeographical discussion. Bernice P.
Bishop Mus. Spec. Publ. 16:1–47
Skottsberg C (1936) La población botánica y zoológica de las islas chilenas del Pacífico. Imprenta
Universitaria, Santiago. Translated by G. Looser from the French original: Berland L, Berlioz J,
Bryan EH et al. (eds) Le peuplement des îles pacifiques du Chili. Contribution à l’étude du
peuplement zoologique et botanique des îles du Pacifique (1934), Société de Biogéographie,
Paris, pp 271–280
Skottsberg C (1937) Die flora der Desventuradas-Inseln (San Félix und San Ambrosio) nach
Sammlungen F. Johows Göteborgs Kungl Vetensk. -Vitterh -Samh. Handl. ser 5 B 5:1–88
Skottsberg C (1951) Weitere Beiträge zur Flora der Insel San Ambrosio. Arkiv Bot 2:12
References 179
Skottsberg C (1956) Derivation of the flora and fauna of Juan Fernandez and Easter Island. In:
Skottsberg C, (ed) The natural history of the Juan Fernandez and Easter Islands, vol 1 (Botany).
Almqvist & Wiksell, Uppsala, pp 193–405
Skottsberg C (1963) Zur Naturgeschichte Der Inseln San Ambrosio (Islas Desventuradas, Chile)
2. Blütenpflanzen. Arkiv Bot 4:464–490
Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG (2006) A classification for
extant ferns. Taxon 55:705–731
Stern C, Moreno H, López-Escobar L, Clavero J, Lara LE, Naranjo JA, Parada MA, Skewes MA
(2007) Chilean volcanoes. In: Moreno T, Gibson W, (eds) The geology of Chile. The Geological
Society of London, London, pp 147–178
Stuessy TF, Sanders RW, Silva M (1984a) Phytogeography and evolution of the flora of the
Juan Fernández Islands: a progress report. In: Redovsky FJ, Raven PH, Sohmer SH, (eds)
Biogeography of the Tropical Pacific. Assoc Syst Coll and B.P. Bishop Museum, Lawrence,
KS, pp 55–69
Stuessy TF, Foland KA, Sutter JF, Sanders RW, Silva M (1984b) Botanical and geological
significance of Potassium-Argon dates from the Juan Fernandez Islands. Science 225:49–51
Stuessy TF, Crawford DJ, Anderson GJ, Jenner RJ (1998a) Systematics, biogeography and
conservation of Lactoridaceae. Perspect Plant Evol Syst 1–2:267–290
Stuessy TF, Crawford DJ, Marticorena C, Silva M (1998b) Isolating mechanisms and modes of
speciation in endemic angiosperms of the Juan Fernández Islands. In: Stuessy TF, Ono M,
(eds) Evolution and speciation of island plants. Cambridge University Press, Cambridge,
pp 79–95
Stuessy TF, Swenson U, Marticorena C, Matthei O, Crawford DJ (1998c) Loss of plant diversity
and extinction on Robinson Crusoe Islands, Chile. In: Peng CI, Lowry PP (eds) Rare, threatened
and endangered floras of Asia and the Pacific Rim. Acad Sin Monogr 16:243–257
Swenson U, Stuessy TF, Baeza M, Crawford DJ (1997) New and historical plant introductions, and
potential pests in the Juan Fernández Islands, Chile. Pac Sci 51:23–253
Teillier S, Taylor C (1997) Maireana Moq., un nuevo género para Chile (Islas Desventuradas).
Gayana Bot 54:15–17
van Balgooy MMJ (1971) Plant-geography of the pacific. Blumea 6(Suppl):1–22
Wallace AR (1880) Island life (or The phenomena and causes of insular faunas and floras: including
a revision and attempted solution of the problem of geological climates). Macmillan, London
Wikström N, Savolainen V, Chase MW (2001) Evolution of the angiosperms: calibrating the family
tree. Proc R Soc Lond B 268:2211–2220
Yáñez GA, Cembrano J, Pardo M, Ranero C, Selles D (2002) The challenger-Juan Fernández-
Maipo major tectonic transition of the Nazca-Andean subduction system at 33–34◦ S: geody-
namic evidence and implications. J South Am Earth Sci 15:23–38
Yáñez GA, Ranero CR, von Huene R, Díaz J (2001) Magnetic anomaly interpretation across the
southern central Andes (32◦ –34◦ S): the role of the Juan Fernández Ridge in the late Tertiary
evolution of the margin. J Geophys Res Solid Earth 106:6325–6345
Zizka G (1991) Flowering plants of Easter Island. Palmarum Hortus Francofurt 3:1–108
Chapter 6
Islands on the Continent:
Conservation Biogeography in Changing
Ecosystems
Abstract Conservation biogeography meets many challenges around the world

associated with increasing human pressure on ecosystems and processes like global
change. Threats as well as conservation opportunities are differentially distributed
over the Chilean territory, and are here explored in relation to modern approaches
such as systematic conservation planning.
During the seventies researchers found that concepts theoretical and empirical con-
cepts developed under the mainframe of island biogeography, as extinction and
colonization, could be adapted to the analysis of other isolated territories on the
mainland like mountains, deserts, water pods, or forest fragments surrounded by
the human-altered matrix (MacArthur and Wilson 1967; Marzluff 2005; Whittaker
et al. 2008). In practice, concepts of island biogeography have been increasingly
applied in conservation planning and the design of reserves networks (Wilson and
Willis 1975; Shafer 1990; Pisano 1996; Lomolino et al. 2006). The explicit link
between conservation and biogeography has been systematized as a new scientific
field, namely “conservation biogeography” (Whittaker et al. 2005).
6.1 Fragmentation v/s Conservation on Chilean

Landscapes
Chile has a long history of landscape modification and forests substitution, since
the native people inhabiting the land, through the Spanish conquerors’ settlement
and the successive waves of European immigration (Lauer 1961; Golte 1973). The
country’s rapid economic development during the last decades of the twentieth cen-
tury has only increased the expansion of the cultural landscape and the urban limits
(Pauchard et al. 2006). Dramatic is the widespread substitution of sclerophyllous
and deciduous forests in Central Chile by exotic plantations and agriculture (Torres-
Mura et al. 2008), and the progressive fragmentation, by logging and clearance, of

DOI 10.1007/978-90-481-8748-5_6, ⃝
182 6 Islands on the Continent
the southern temperate forests (Grez et al. 2006; Echeverría et al. 2006) (Figs. 6.1
and 6.2).
The impacts of human activities already affect most ecosystems, from the north-
ern “bofedales” (cushion bogs) and tamarugales through the mediterranean matorral
to the coastal southern forests (Hoffmann 1998; Smith-Ramírez 2004; Torres-Mura
et al. 2008) (see Sect. 1.3). Activities that have a big impact on Chilean ecosystems
are related to all industrial sectors from forestry to mining, including agriculture and
even non-regulated tourism. Fires are not a natural aspect of the Chilean territory,
but induced by colonos affected ca 3 million hectares in the north patagonian forests
between the 1936 and 1956 (Quintanilla 2008) and continue affecting thousands
of hectares yearly in Mediterranean Chile (Quintanilla and Castro 1998). Mining
is the main economic activity of the country, and the environmental impacts of
the industry are huge, especially in the North where the sparse water resource has
been constantly captured by the mining projects in detriment of the fragile Andean
ecosystems and indigenous communities (see also Sect. 7.3). Mining projects are
even affecting other physical features like glaciers and rock glaciers (Brenning
2008).
Calls for a reduction of the effects of human impact on Chilean ecosystems were
already made by Gay (1838), Albert (1903), Garaventa (1936), Elizalde-McClure
(1970), Cunill (1970), and Muñoz Pizarro (1973), among others. Federico Albert
was the principal promoter of the forest and conservation legislation (Camus 2006).
Carl Skottsberg was one of the promoters of the creation of Juan Fernández National
Park (Chap. 5). Agustín Garaventa was the promoter of the creation of iconic La
Campana National Park (Garaventa 1964). Carlos Muñoz Pizarro (Chap. 2) pro-
vided the first national list of conservation priority sites (1969), and the first national
account of endangered plant species (1973). Following scientific recommendations,
the reaction of the state was a strong impulse towards the creation of protected areas
during the 1960s and 1970s according to the “pristine” or “wilderness” concept
(Pauchard and Villarroel 2002) (Box 6.1).
a b
Fig. 6.1 Examples of human impacts on native ecosystems: a substitution of matorral for plan-
tations; b exploitation of alerce (Fitzroya cupressoides) inside the temperate forest (photos by
A. Moreira-Muñoz)
6.1 Fragmentation v/s Conservation on Chilean Landscapes 183
b c
Fig. 6.2 Cultural landscapes: a intensively cultivated Central Depression, Río Aconcagua; b
advancement of the city upon natural ecosystems as the case of Concón palaeodunes; c map of
Central Chile showing remnants of native vegetation and cultural land uses; d intensive substi-
tution of natural vegetation for Pinus plantations, Maule region (photo/maps credits: a, b, d A.
Moreira-Muñoz; c National Vegetation Survey (1997) CONAF/CONAMA/BIRF)
Box 6.1 The First Protected Area and the Evolution of the
SNASPE
The first Chilean protected area was created in 1907, namely the Malleco
National Forest Reserve established in southern Chile (Cabeza 1988). In
1925 the first National Park, Benjamín Vicuña Mackennna, was created and
a few years later its limits were modified. The first protected area which
still exists is Vicente Pérez Rosales National Park, designed in 1926 (Fig. 6.3).
Arica y Biobío
Parinacota
Araucanía
20°
Tarapacá
Los Ríos 40°
Los Lagos
Antofagasta
25°
45°
I. de Pascua
Aysén
Atacama
0 250 500 km
30°
Coquimbo 50°
Magallanes
Juan Fernández Valparaíso

Metropolitana
O`Higgins
35°
Maule
55°
Fig. 6.3 Distribution of protected areas in Chilean administrative regions: arrow shows the
location of Vicente Pérez Rosales National Park; dots represent reduced areas; stars show
the presence of Biosphere Reserves
The creation of protected areas in the early 1900s was more a response to
the interest of visionary naturalists acting against deforestation, rather than a
national conservation policy (Pauchard and Villarroel 2002). During the 1960s
and 1970s the protected surface grew exponentially, especially in southern
Chile, and this was more related to aesthetic considerations and the low value
6.2 Global Change Biogeography: A Science of Uncertainties . . . and Possibilities 185
of the land than to “biodiversity”, a concept that at that time still did not
existed. Only in 1984 the “Sistema Nacional de Áreas Silvestres Protegidas
del Estado” (SNASPE) was officially created. The SNASPE adopted the
framework of the 1978 IUCN protected area categories to comply with inter-
national agreements. Four categories of protected areas were defined: Virgin
Region Reserves (with no unit in Chile), National Parks, Natural Monuments,
and National Reserves. The management of the system was assigned to the
“Corporación Nacional Forestal” (CONAF), a private corporation dependent
of the Agriculture Ministry.
At present (December 2009) the system is composed of 97 units occu-
pying more than 14 million ha, or 19% of the national territory (Fig. 6.3)
This includes 33 National Parks, 49 National Reserves, and 15 Natural
Monuments.
The 19% of national protected territory is a good proportion at global standards.

Nevertheless, the system shows a weakness as it is not representative: most of the
protected areas are located in the south, and the Central Chilean ecosystems are
absolutely underrepresented (Mardones and Henríquez 1996; Luebert and Becerra
1998; Luebert and Pliscoff 2006). This last assessment shows that the protected
areas system still does not protect 25 of 127 ecosystems, and 47 of the 127 ecosys-
tems have less than 10% of their remnant surface under protection (Luebert and
Pliscoff 2006). The ecosystem conservation approach is still under development
in Chile, since most conservation efforts have been concentrated on several iconic
target faunal or floral species, like alerce (Fitzroya cupressoides).
Efforts to improve the representativeness of the system include the recognition
of priority sites at the national and regional levels, coupled with species red lists
(Benoit 1989; Muñoz-Schick et al. 1996; Squeo et al. 2001, 2008; Serey et al.
2007; CONAMA 2008), but the real incorporation of new units is a slow process.
A newer system should incorporate, under different protocols of use and manage-
ment, natural sanctuaries, private parks, and other categories. Complementary to
the in-situ conservation are efforts catalogued as ex-situ, like the National Botanic
Garden (www.jardin-botanico.cl), the Metropolitan Botanic Garden “Chagual”
(www.chagual.cl) and the Millennium Seedbank with its centre located in Vicuña,
Coquimbo region (León-Lobos et al. 2003).
6.2 Global Change Biogeography: A Science of Uncertainties . . .

and Possibilities
Range shifts of species promoted by global change have enormous implications for
biodiversity planning and management (Root et al. 2007). This means but working
within scenarios of crescent uncertainty (Regan et al. 2005; Pearson et al. 2006;
Guilhaumon et al. 2008; McDonald-Madden et al. 2008). “Our uncertainty regard-

ing the consequences of losing species can seem overwhelming, even paralyzing”
(Kareiva and Levin 2003, p. 345). This might not be stimulating thoughts for plan-
ners and decision-makers, and serve to maintain skepticism towards global change.
Alternatively, these people need certainties in order to act, and therefore they need to
tergiversate scientific models into “realities” (Lehmkuhl 2008; Egner 2008). “While
the climate skeptics have sought to refute climate change science by exposing the
socially negotiated assumptions and uncertainties of the climate models, advocates
of greenhouse gases reduction have responded by denying them altogether. Neither
response is very helpful. What is needed instead is a more reflexive understanding
of science as a social practice” (Demeritt 2001, p. 329).
In this sense, conservation biogeography seeks a better understanding of the
impacts of global change on ecosystems and species. Species range dynamics have
long been of crucial interest for ecologists and biogeographers as illustrated by the
classical example from Peters and Lovejoy (1992) (Fig. 6.4).
Ecological responses to recent climate change are already evident (Channell and
Lomolino 2000; Walther et al. 2002; Araújo et al. 2004). Global climate change has
already begun affecting species’ geographic ranges by means of poleward shifts in
range limits correlated with a warming climate and changes in precipitation, and
phenological changes portend poleward shifts (Parmesan and Yohe 2003; Parmesan
2007). Upslope shifts in range boundaries along temperate elevational gradients also
have been documented (Wilson et al. 2007; Lenoir et al. 2008). Even in tropical
lowland areas, biotas may face a level of contraction due to global warming (Colwell
et al. 2008).
The effects of global change on Chilean geosystems are increasingly attract-
ing attention and research (e.g. Andrade and Peña 1993; Lauenroth et al. 2004).
Widespread retreat of glaciers in the Central Andes during the nineteenth and twen-
tieth century has been extensively documented (Rivera et al. 2006; Le Quesne et al.
2009) (Fig. 6.5). Scenarios of increasing temperatures and decreasing precipitation,
spatially differentiated, predict important effects on the Chilean ecosystems (see
Arroyo et al. 1993; DGF 2006; Searle and Rovira 2008).
To improve biodiversity conservation across the landscape, a possibility is to
abandon the “wilderness ideal” (Jax and Rozzi 2004) and move towards a more
Planed Actual Actual

Reserve Reserve Reserve
a b c
Fig. 6.4 a represents a reserve created today for the conservation of a targeted endangered taxon
(black area). b It is very plausible that in several decades the geographic range of this species
will move (arrow) in response to anthropogenic pressure and climate changes. c The core of the
distribution area would rapidly disappear and the species would be left at the edge. The reserve,
at least for the conservation of this taxon, has become useless (redrawn from Peters and Lovejoy
(1992))
Fig. 6.5 Glaciar retreat in Cajón de Cipreses (35ºS) from 1840 to the present; a illustration from
Pissis’ Atlas (1975) (Chap. 2); b glacier retreat as shown in an aerial photograph (from Le Quesne
et al. 2009, by permission of Elsevier)
realistic one, in which human activities are incorporated into conservation objec-
tives in vast territories, i.e. a model of conservation “in the real world” (Prendergast
et al. 1999; Moreira-Muñoz 2005; Branquart et al. 2008). This is certainly a very
compelling option for a rapidly urbanizing country like Chile. The actual tendency
runs towards a new protected areas systems (a kind of SNASPE 2.0) that explicitly
incorporated Natural Sanctuaries, RAMSAR-sites, marine reserves (Rovira et al.
2008), and privately protected areas into a new spatially and operationally integrated
system. This should include peri-urban parks at the edge of the city (Elórtegui
2005) and novel concepts like geoparks (Castro and Zúñiga 2007) that follow new
international standards of protected areas systems (e.g. Boitani et al. 2008).
In the same direction is the reconsideration of the old but always valid biosphere
reserve model: “Biosphere reserves celebrate the cultural relationship between peo-
ple and places” (Berghofer et al. 2008) and are indeed the best suited places for
promoting the environmental education of the community (Fraser and Jamieson
2003). The spatial arrangement of transition zones, buffers and corridors has to
be realized with the participation of local stakeholders in a joint process with the
scientific community, as in the case of La Campana-Peñuelas Biosphere Reserve
(Elórtegui and Moreira-Muñoz 2002). This implies the rescue of ancient local
knowledge, which opens a better opportunity for learning, communication, and
acting. Indeed, the association with indigenous and rural communities towards com-
mon conservation goals is one of the biggest challenges for the protected areas
system in Chile (see Oltremari and Jackson 2006; Castro and Romo 2008).
Biosphere reserves in Chile are still lacking support to cover the multiple chal-
lenges that includes this nomination, but a slow but constant local impulse is
generating management plants (e.g. for La Campana) and the creation of new
reserves, like Cabo de Hornos (Rozzi et al. 2004) and Bosques Templados, this lat-
ter in a bi-national effort with Argentina. Currently there are nine reserves in Chile
on the more diverse ecosystems (Fig. 6.3).
The biosphere reserves concept is an attempt to integrate the protected areas
with the surrounding human matrix, trying to couple biodiversity conservation with
human interests (Naughton-Treves et al. 2005). Therefore, conservation planning
problems can be formulated as optimization problems (Sarkar et al. 2006; Margules
and Sarkar 2007) (Box 6.2).
Box 6.2 Systematic Conservation Planning and Priority Site

Selection
Different algorithms have been developed in the search for the “best” solu-
tion for conservation, and planning tools are typically used to satisfy this
while dealing with multiple criteria (Sarkar et al. 2006). This approach got
the name of “systematic conservation planning”, defined as a structured
approach with feedback and reiteration at any stage (Margules and Sarkar
2007). These stages include engagement of stakeholders, choices about bio-
diversity measurements, GIS mapping, setting of surrogates, the application
of complementarity as a main concept for the identification of new conser-

vation areas, and monitoring management actions, among others (Margules
and Pressey 2000). A key concept is complementarity: the marginal contri-
bution of an area to represent the full set of attributes defined as biodiversity
surrogates (Margules and Sarkar 2007).
Conservation biogeography is specifically focusing on inter-related fac-
tors like scale dependency (both spatial and temporal), and inadequacies in
taxonomic and distributional data (the so-called Linnean and Wallacean short-
falls): the Linnean shortfall means the lack of appropriate knowledge on the
taxonomy of the species; and the Wallaceaen shortfall, approaching the lack
of adequate knowledge on the distribution of the species (Whittaker et al.
2005). As expressed by Myers and Giller (1988): “Knowledge of the distribu-
tion of most organisms is weak, as is knowledge of the faunas and floras of
many regions of the globe. In this respect, current biogeographic hypotheses
are built around very incomplete basic data and we are perhaps trying to run
before we can walk” (Myers and Giller 1988, p. 21)
The systematic conservation planning approach has been recently applied
in Chile by Ramírez de Arellano (2007) and Tognelli et al. (2009). As an
example from Central Chile, currently protected areas and priority sites have
been assessed by a set of surrogates related to available biological informa-
tion at the regional scale (Fig. 6.6). The map shows some of the cartographic
results expressed as complementary sites composing a regional connected
reserves network, concentrated towards the coastal Cordillera, which harbours
most of endemics and taxa with restricted ranges.
71°
32°
33°
Protected areas
Priority sites
Complementary sites
0 25 50 km
34°
Fig. 6.6 Proposal of a network of reserves for the conservation of the vascular flora of the
Valparaíso region (Morales and Moreira-Muñoz 2009)
Cartographic proposals are highly dependent on available biological data: most

models rely on species richness, but there are other aspects to be considered like
“phylogenetic diversity” (Rodrigues and Gaston 2002; Forest et al. 2007). Specific
assessment for southern South America showed that Central Chile is one of the high-
est priority areas for conservation (Posadas et al. 2001). Also Heads (2009) empha-
sized the importance of South American basal angiosperm groups like Lactoris,
Gomortega, Berberidopsis, and Aextoxicon, as components of one of the “glob-
ally basal centres of endemism”. All these relevant species are in serious danger of
extinction due to human activities (Hechenleitner et al. 2005; Alarcón et al. 2007),
together with other iconic species like Nothofagus alessandrii (Bustamante and
Castor 1998; Olivares et al. 2005), Nothofagus glauca (Burgos et al. 2008), Fitzroya
cupressoides, Pitavia punctata or Legrandia concinna (Altamirano et al. 2007).
References
Alarcón D, Smith-Ramírez C, Hechenleitner P, Oliva M, Pinto M (2007) Nuevas poblaciones de
la enredadera Berberidopsis corallina Hook. F. en la Región del Bío Bío, Chile. Ubicación,
hábitat y grado de conservación. Gayana Bot 64:216–229
Albert F (1903) Informe de Federico Albert sobre sus estudios en las provincias de Malleco, Cautín
y Valdivia. Memoria del Ministerio de Industria y Obras Públicas, Santiago
Altamirano A, Echeverría C, Lara A (2007) Effect of forest fragmentation on vegetation structure
of Legrandia concinna (Myrtaceae) threatened populations in south-central Chile. Rev Chil
Hist Nat 80:27–42
Andrade B, Peña H (1993) Chilean geomorphology and hydrology: response to global change. In:
Mooney HA, Fuentes ER, Kronberg BI, (eds) Earth system responses to global change: contrast
between North and South America. Academic Press, New York, NY, pp 101–114
Araújo MB, Cabeza M, Thuiller W, Hannah L, Williams PH (2004) Would climate change drive
species out of reserves? An assessment of existing reserve-selection methods. Glob Change
Biol 10:1618–1626
Arroyo MTK, Armesto JJ, Squeo FA, Gutiérrez J (1993) Global change: Flora and Vegetation
of Chile. In: Mooney HA, Fuentes ER, Kronberg BI, (eds) Earth system responses to
global change: contrast between North and South America. Academic Press, New York, NY,
pp 239–263
Benoit I, (ed) (1989) Libro Rojo de la Flora Terrestre de Chile. Ministerio de Agricultura,
Corporación Nacional Forestal, Santiago
Berghofer U, Rozzi R, Jax K (2008) Local versus global knowledge: diverse perspectives on nature
in the Cape Horn Biosphere Reserve. Environ Ethics 30:273–294
Boitani L, Cowling RM, Dublin HT, Mace GM, Parrish J, Possingham HP, Pressey RL,
Rondinini C, Wilson KA (2008) Change the IUCN protected area categories to reflect
biodiversity outcomes. PLoS Biol 6:e66. doi:10.1371/journal.pbio.0060066
Branquart E, Verheyen K, Latham J (2008) Selection criteria of protected forest areas in Europe:
the theory and the real world. Biol Conserv 141:2795–2806
Brenning A (2008) The impact of mining on rock glaciers and glaciers: examples from Central
Chile. In: Orlove BS, Wiegandt E, Luckman B, (eds) Darkening peaks: glacier retreat, science,
and society. University of California Press, Berkeley, CA, pp 196–205
Burgos A, Grez AA, Bustamante RO (2008) Seed production, pre-dispersal seed predation and
germination of Nothofagus glauca (Nothofagaceae) in a temperate fragmented forest in Chile.
For Ecol Manag 255:1226–1233
Bustamante RO, Castor C (1998) The decline of an endangered temperate ecosystem: the ruil
(Nothofagus alessandrii) forest in central Chile. Biodivers Conserv 7:1607–1626
References 191
Cabeza A (1988) Aspectos históricos de la legislación forestal vinculada a la conservación, la

evolución de las áreas silvestres protegidas de la zona de Villarrica y la creación del primer
Parque Nacional en Chile. Documento de Trabajo No. 101, CONAF, Santiago
Camus P (2006) Ambiente, bosques y gestión forestal en Chile. 1541–2005. Colección Sociedad y
Cultura, Central de Investigaciones Diego Barros Arana, LOM ediciones, Santiago
Castro C, Zúñiga A (2007) Geomorfología del AMCP – MU Isla Grande Atacama. Instituto de
Geografía, Pontificia Universidad Católica de Chile, Santiago
Castro V, Romo M (2008) Tradiciones Culturales y Biodiversidad. In: CONAMA (ed)
Biodiversidad de Chile: patrimonio y desafíos, 2nd edn. CONAMA, Santiago, pp 468–493
Channell R, Lomolino MV (2000) Trajectories to extinction: spatial dynamics of the contraction
of geographical ranges. J Biogeogr 27:169–179
Colwell RK, Brehm G, Cardelus CL, Gilman AC, Longino JT (2008) Global warming, elevational
range shifts, and lowland biotic attrition in the wet tropics. Science 322:258–261
CONAMA (2008) Biodiversidad de Chile: Patrimonio y Desafíos. 2nd edn. Comisión Nacional de
Medio Ambiente, Chile. www.conama.cl
Cunill P (1970) Factores en la destrucción del paisaje chileno: recolección caza y tala coloniales.
Inf Geogr 20:235–264
Demeritt D (2001) The construction of global warming and the politics of science. Ann Assoc Am
Geogr 91:307–337
DGF (2006) Estudio de la Variabilidad Climática en Chile para el siglo XXI: Informe Final.
Departamento de Geofísica, Facultad de Ciencias Físicas y Matemáticas, Universidad de Chile,
CONAMA, Santiago
Echeverría C, Coomes D, Salas J, Rey-Benayas JM, Lara A, Newton AC (2006) Rapid deforesta-
tion and fragmentation of Chilean Temperate Forests. Biol Conserv 130:481–494
Egner H (2008) See what you can’t see – on “truth” and the credibility of science in the climate
discourse. GAIA 17:337–338
Elizalde-McClure R (1970) La sobrevivencia de Chile. Editorial Universitaria, Santiago
Elórtegui S, (ed) (2005) Las Dunas de Concón: El desafío de los espacios silvestres urbanos. Taller
La Era, Valparaíso
Elórtegui S, Moreira-Muñoz A, (eds) (2002) Parque Nacional la campana: Origen de una reserva
de la Biósfera en Chile Central. Taller la Era, Santiago
Forest F, Grenyer R, Rouget M, Jonathan Davies T, Cowling RM, Faith DP et al. (2007) Preserving
the evolutionary potential of floras in biodiversity hotspots. Nature 445:757–760
Fraser R, Jamieson G (2003) Community environmental education: challenges within the
Biosphere Reserve Concept. Prospects 33:293–302
Garaventa A (1936) La deforestación en la zona de Chile central y su importancia. Rev Univ
21:131–135
Garaventa A (1964) Importancia e interés botánico que presenta la vegetación del cerro La
Campana. Bol Inf Soc Cient de Valparaíso 23:2
Gay C (1838) Sobre las causas de la disminución de los montes de la provincia de Coquimbo.
In: Squeo FA, Arancio G, Gutiérrez JR, (eds) Libro Rojo de la Flora Nativa y de los Sitios
Prioritarios para su Conservación: Región de Coquimbo (2001). Ediciones Universidad de La
Serena, La Serena, pp 281–286
Golte W (1973) Das südchilenische Seengebiet: Besiedlung und wirtschaftliche Erschliessung seit
dem 18. Jahrhundert. Bonn Geogr Abh 47:1–183. + Figs
Grez A, Simonetti J, Bustamante RO (2006) Biodiversidad en Ambientes Fragmentados de Chile:
Patrones y Procesos a Diferentes Escalas. Editorial Universitaria, Santiago
Guilhaumon F, Gimenez O, Gaston KJ, Mouillot D (2008) Taxonomic and regional uncertainty in
species-area relationships and the identification of richness hotspots. Proc Natl Acad Sci USA
105:15458–15463
Heads M (2009) Globally basal centres of endemism: the Tasman-Coral Sea Region (south-west
Hechenleitner P, Gardner MF, Thomas PI, Echeverría C, Escobar B, Brownless P, Martínez C

(2005) Plantas Amenazadas del Centro-Sur de Chile: Distribución, Conservación y
Propagación. Universidad Austral de Chile y Real Jardín Botánico de Edimburgo, Valdivia
Hoffmann AE, (ed) (1998) La Tragedia del Bosque Chileno. Defensores del Bosque Chileno, Ocho
Libros Editores, Santiago
Jax K, Rozzi R (2004) Ecological theory and values in the determination of conservation goals:
examples from temperate regions of Germany, United States of America, and Chile. Rev Chil
Hist Nat 77:349–366
Kareiva P, Levin SA, (eds) (2003) The importance of species: perspectives on expendability and
triage. Princeton University Press, Princeton, NJ
Lauenroth WK, Epstein HE, Paruelo JM, Burke IC, Aguiar MR, Sala OE (2004) Potential effects
of climate change on the temperate zones of North and South America. Rev Chil Hist Nat
77:439–453
Lauer W (1961) Wandlungen im Landschaftsbild des Südchilenischen Seengebietes seit Ende der
spanischen Kolonialzeit. Schr Geogr Inst Univ Kiel 20:227–276
Le Quesne C, Acuña C, Boninsegna JA, Rivera A, Barichivich J (2009) Long-term glacier varia-
tions in the Central Andes of Argentina and Chile, inferred from historical records and tree-ring
reconstructed precipitation. Paleogeogr Paleoclim Paleoecol. doi:10.1016/j.palaeo.200801.039
Lehmkuhl M (2008) Neither coincidence nor success: how to interpret the current debate about
climate change. GAIA 17:9–11
Lenoir J, Gégout JC, Marquet PA, de Ruffray P, Brisse H (2008) A significant upward shift in plant
species optimum elevation during the 20th century. Science 320:1768–1771
León-Lobos P, Way M, Pritchard H, Moreira-Muñoz A, León M, Casado F (2003) Conservación
ex situ de la Flora de Chile en banco de semillas. Chloris chilensis 6. http://www.chlorischile.cl
Lomolino MV, Riddle BR, Brown JH (2006) Biogeography. Sinauer Associates, Sunderland, MA
Luebert F, Becerra P (1998) Representatividad vegetacional del Sistema Nacional de Áreas
Silvestres Protegidas del Estado (SNASPE) en Chile. Ambiente y Desarrollo 14:62–69
Luebert F, Pliscoff P (2006) Sinopsis bioclimática y vegetacional de Chile. Editorial Universitaria,
Santiago
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press,
New York, NY
Mardones G, Henríquez C (1996) Áreas Silvestres Protegidas de Chile. Serie de Monografías, Doc.
de trabajo N◦ 1. CODEFF, Santiago
Margules CR, Pressey RL (2000) Systematic conservation planning. Nature 405:243–253
Margules CR, Sarkar S (2007) Systematic conservation planning. Cambridge University Press,
Cambridge
Marzluff JM (2005) Island biogeography for an urbanizing world: how extinction and colonization
may determine biological diversity in human-dominated landscapes. Urban Ecosyst 8:157–177
McDonald-Madden E, Baxter PWJ, Possingham HP (2008) Making robust decisions for conserva-
tion with restricted money and knowledge. J Appl Ecol 45:1630–1638
Morales V, Moreira-Muñoz A (2009) GIS and systematic conservation planning in Chile, abstracts,
international cartography conference, Santiago, Chile, Nov 15–21, 2009
Moreira-Muñoz A (2005) Conservation in a changing world: biodiversity hotspots and the
distracting paradigm. J Conserv Plann 1:1–6
Muñoz Pizarro C (1969) Áreas Naturales. En: Segunda Jornada Latinoamericana de Parques
Nacionales, Oct 1–4, 1969, Viña del Mar. Ministerio de Agricultura.
Muñoz Pizarro C (1973) Chile: plantas en extinción. Editorial Universitaria, Santiago
Muñoz-Schick M, Yánez J, Núñez H (1996) Libro Rojo de los Sitios Prioritarios para la
Conservación de la Biodiversidad en Chile. CONAF, Santiago
Myers AA, Giller PS, (eds) (1988) Analytical biogeography. Chapman and Hall, London
Naughton-Treves L, Buck Holland M, Brandon K (2005) The role of protected areas in conserving
biodiversity and sustaining local livelihoods. Annu Rev Env Resour 30:219–252
Olivares P, San Martín J, Santelices R (2005) Ruil (Nothofagus alessandrii): Estado del
conocimiento y desafíos para su conservación. Departamento de Protección de Recursos
Naturales, Comisión Nacional del Medio Ambiente Región del Maule, Talca
References 193
Oltremari JV, Jackson RG (2006) Conflicts, perceptions, and expectations of indigenous commu-
nities associated with natural areas in Chile. Nat Areas J 26:215–220
Parmesan C (2007) Influences of species, latitudes and methodologies on estimates of phenological
response to global warming. Glob Change Biol 13:1860–1872
Parmesan C, Yohe G (2003) A globally coherent fingerprint of climate change impacts across
natural systems. Nature 421:37–42
Pauchard A, Villarroel P (2002) Protected areas in Chile: history, current status and challenges.
Nat Areas J 22:318–330
Pauchard A, Aguayo M, Peña E, Urrutia R (2006) Multiple effects of urbanization on the biodiver-
sity of developing countries: the case of a fast-growing metropolitan area (Concepcion, Chile).
Biol Conserv 127:272–281
Pearson RG, Thuiller W, Araujo MB, Martínez-Meyer E, Brotons L, McClean C, Miles L,
Segurado P, Dawson TP, Lees DC (2006) Model-based uncertainty in species range prediction.
J Biogeogr 33:1704–1711
Peters RL, Lovejoy TE (1992) Global warming and biological diversity. Yale University Press,
London
Prendergast JR, Quinn RM, Lawton JH (1999) The gaps between theory and practice in selecting
nature reserves. Conserv Biol 13:484–492
Pisano E (1996) Implicaciones de la teoría de biogeografía de islas para el diseño de reservas
naturales. An Inst Pat (Ser Cienc Nat) 24:117–126
Posadas P, Miranda-Esquivel DR, Crisci JV (2001) Using phylogenetic diversity measures to set
priorities in conservation: an example from southern South America. Conserv Biol 15:1325–
1334
Quintanilla V (2008) Estado de recuperación del bosque nativo en una cuenca nordpatagónica de
Chile, perturbada por grandes fuegos acaecidos 50 años atrás (44º–45ºS). Rev Geogr Norte
Grande 39:71–91
Quintanilla V, Castro R (1998) Seguimiento de las cubiertas vegetales post-incendios forestales en
la zona mediterránea costera de Chile. Ser Geogr (Univ Alcalá) 7:147–154
Ramírez de Arellano PI (2007) Systematic conservation planning in Chile: sensitivity of reserve
selection procedures to target choices, cost surface, and spatial scale. PhD thesis, State
University of New York, College of Environmental Science and Forestry, Syracuse, New
York, NY
Regan HM, Ben-Haim Y, Langford B et al. (2005) Robust decision-making under severe
uncertainty for conservation management. Ecol Appl 15:1471–1477
Rivera A, Acuña C, Casassa G (2006) Glacier variations in central Chile (32◦ S–41◦ S). In: Knight
PG, (ed) Glacier science and environmental change. Blackwell, Oxford, pp 246–247
Rodrigues ASL, Gaston KJ (2002) Maximising phylogenetic diversity in the selection of networks
of conservation areas. Biol Conserv 105:103–111
Root TL, Liverman D, Newman C (2007) Managing biodiversity in the light of climate change:
current biological effects and future impacts. In: Macdonald D, Service K, (eds) Key topics in
conservation biology. Blackwell, Oxford, UK, pp 85–104
Rovira J, Ortega D, Alvarez A, Molt K (2008) Areas Protegidas en Chile. In: CONAMA (ed)
Biodiversidad de Chile: patrimonio y desafíos, 2nd edn. CONAMA, Santiago, pp 506–561
Rozzi R, Massardo F, Anderson C, (eds) (2004) Reserva de Biosfera Cabo de Hornos: una prop-
uesta de conservación y turismo para el desarrollo sustentable en el extremo austral de América.
Ediciones de la Universidad de Magallanes, Punta Arenas
Sarkar S, Pressey RL, Faith DP, Margules CR, Fuller T, Stoms DM et al. (2006) Biodiversity
conservation planning tools: present status and challenges for the future. Annu Rev Env Resour
31:123–159
Searle JP, Rovira J (2008) Cambio climático y efectos en la biodiversidad: el caso chileno. In:
CONAMA (ed) Biodiversidad de Chile: patrimonio y desafíos, 3rd ed. CONAMA, Santiago,
pp 502–503
Serey I, Ricci M, Smith-Ramírez C, (eds) (2007) Libro Rojo de la Región del Libertador Bernardo
O’Higgins. CONAF, Santiago
Shafer CL (1990) Nature reserves: island theory and conservation practices. Smithsonian
Institution Press, Washington, DC
Smith-Ramírez C (2004) The Chilean coastal range: a vanishing center of biodiversity and
endemism in South American temperate rainforests. Biodivers Conserv 13:373–393
Squeo FA, Arancio G, Gutiérrez JR, (eds) (2001) Libro Rojo de la Flora Nativa y de los Sitios
Prioritarios para su Conservación: Región de Coquimbo. Ediciones Universidad de La Serena,
La Serena
Squeo FA, Arancio G, Gutiérrez JR, (eds) (2008) Libro Rojo de la Flora Nativa y de los Sitios
Prioritarios para su conservación: Región de Atacama. Ediciones Universidad de La Serena, La
Serena, pp 137–163
Tognelli MF, Fernández M, Marquet PA (2009) Assessing the performance of the existing and
proposed network of marine protected areas to conserve marine biodiversity in Chile. Biol
Conserv 142:3147–3153
Torres-Mura JC, Castro S, Oliva D (2008) Conservación de la Biodiversidad. In: CONAMA (ed)
Biodiversidad de Chile: patrimonio y desafíos, 2nd ed. CONAMA, Santiago, pp 413–431
Walther GR, Post E, Convey P, Menzel A, Parmesan C, Beebee TJC, Fromentin JM, Hoegh-
Guldberg O, Bairlein F (2002) Ecological responses to recent climate change. Nature
416:389–395
Whittaker RJ, Araujo MB, Paul J, Ladle RJ, Watson JEM, Willis KJ (2005) Conservation
biogeography: assessment and prospect. Divers Distrib 11:3–23
Whittaker RJ, Triantis KA, Ladle RJ (2008) A general dynamic theory of oceanic island
biogeography. J Biogeogr 35:977–994
Wilson EO, Willis EO (1975) Applied biogeography. In: Cody ML, Diamond MM, (eds) Ecology
and evolution of communities. Belknap Press, Cambridge, pp 522–534
Wilson RJ, Gutierrez D, Gutierrez J, Monserrat VJ (2007) An elevational shift in butterfly
species richness and composition accompanying recent climate change. Glob Change Biol 13:
1873–1887
Part IV
Case Studies on Selected Families
Chapter 7
Cactaceae, a Weird Family and Postmodern
Evolution
Abstract One of the most charismatic American plant families, the Cactaceae, is
well represented in Chile. Biogeographic analysis of the Chilean representatives
is rarely undertaken, mainly due to the complex taxonomy of the group. Taking
account of the most recent taxonomic treatments and distributional knowledge,
an attempt is made here to develop a biogeographic regionalization by means of
parsimony analysis of endemism (PAE) and the program NDM/VNDM. The first
method yields three areas of endemism in northern Chile bordering the Atacama
Desert to the North and to the South, and the second method yields one area which
includes North-Central Chile. Advantages and restrictions of the methods are dis-
cussed. Finally, the evolution of the family is discussed in relation to the rise of the
Atacama Desert and the systematic interpretation of the group
“Cacti have a special fascination all of their own. Like the brilliantly colored hum-
mingbirds, the Cactaceae are creatures of the New World” (Barthlott 2001, p 9).
With representatives ranging from minute spiny dwarf cacti of a couple centimetres
in diameter to the majestic columns of the Arizona giant saguaro, the Cactaceae is
the second largest family of angiosperms almost exclusively restricted to the New
World. The richest American plant family is Bromeliaceae; both families have one
taxon occurring outside America: the Bromeliaceae has Pitcairnia as a native genus
in West Africa; the Cactaceae has Rhipsalis wider distributed in Africa, Madagascar,
Mauritius, the Seychelles and Ceylon. The family is present in America from cen-
tral Canada to Patagonia. The highest diversity is found in the arid environments
of North and South America, but contrary to the popular believe due to their spiny
morphology, the family is not strictly restricted to arid environments: it also occu-
pies temperate, sub-tropical, and tropical environments reaching mesic southern
Chile and Argentina and ranging from the coast to the high Andes (Kraus 1995;
Mauseth et al. 2002). The southern limit for the family is the Santa Cruz province
in Argentina (ca. 50◦ S, Anderson 2001), where Austrocactus, Gymnocalycium,
Maihuenia, Maihueniopsis, and Pterocactus occur.

DOI 10.1007/978-90-481-8748-5_7, ⃝
198 7 Cactaceae, a Weird Family and Postmodern Evolution
7.1 Cacti Classification

The morphological characteristics of cacti have long attracted botanists and ama-
teur plant lovers. First exemplars known in Europe supposedly were part of the gifts
that Columbus brought to Isabella when returning from his first voyage to the West
Indies (Barthlott 2001), and first reports on cultivation of cacti in Europe date back
to the middle of the sixteenth century (Mauseth et al. 2002). Cactaceae can be super-
ficially distinguished by the succulent stems (with exceptions) and characteristically
clustered spines (Mauseth et al. 2002) (Fig. 7.1).
The subdivision of the cacti into three subfamilies dates back to Schumann
(1898). He placed Pereskia and Maihuenia in the subfamily Pereskioideae, which
was already established by Engelmann in 1876 (Leuenberger 2008). In the middle
1990s, Maihuenia was transferred to its own subfamily, Maihuenioideae (Wallace
1995). Currently both subfamilies are recognized, together with the Opuntioideae
and the most evolved and species-rich Cactoideae (Griffith 2008).
– Pereskioideae is composed of leafy, spiny, often non-succulent trees and shrubs of
the genus Pereskia (16 species). Traditionally it is considered the most primitive
or relictual genus in the Cactaceae, supposedly resembling the “proto-Cactaceae”
ancestor. The subfamily is not represented in Chile, but is extensively distributed
from the Caribbean along the Andes to Argentina and SE Brazil, surrounding the
Amazonas basin (map in Leuenberger 2008).
– Opuntioideae have minute, barbed, deciduous spines (= glochids), and small,
often ephemeral leaves, and a bony seed aril. It comprises about 300 species,
occupying a geographic range from North America to Patagonia.
– Cactoideae are leafless, spiny stem succulents, considered the most “derived”
subfamily, occurring in America, Africa, Madagascar, Mauritius, Seychelles and
Ceylon due to the genus Rhipsalis.
– Maihuenioideae is composed of the two species of the genus Maihuenia,
distributed in southern Chile and Argentina (Leuenberger 1997, 2008). The
Maihuenioideae is a little subfamily that is the last to have been recognized, and
enjoys a huge interest with regard to the evolution of the whole family (Sect. 7.5).
7.2 Chilean Representatives and Their Distribution

The dynamic taxonomy of the group results in very different accounts of the rep-
resentatives in Chile: Lembcke and Weisser (1979) considered 23 genera, while
Marticorena (1990) gave a list of 21 genera and 154 native species for Chile (plus
almost 100 infraspecific taxa). More recently, collaborators of the Southern Cone
Checklist consider the Chilean Cactaceae as represented by 21genera and 112
species, plus quite a number of varieties and subspecies (Zuloaga et al. 2008). This
account, that also presents 88 Chilean endemic species, is considered here as the
most recent, valid checklist (Table 7.1). The inclusion of Miqueliopuntia as different
from Austrocylindropuntia is based on more recent results from Griffith and Porter
7.2 Chilean Representatives and Their Distribution 199
a b
c d
Fig. 7.1 Chilean cacti: a Eulychnia iquiquensis, Tocopilla; b Pyrrhocactus curvispinus, La

Campana; c Eriosyce aurata, Illapel; d Browningia candelaris, Tarapacá (photo credits: a–c A.
Moreira-Muñoz; d Carlo Sabaini);
(2009). All former Chilean Opuntia species have been transferred to other genera
like Maihueniopsis or Pterocactus. Tephrocactus nigrispinus (cfr Pinto and Kirberg
2009) is considered by the Southern Cone Checklist as a synonym of Maihueniopsis
nigrispina. Cylindropuntia tunicata is considered as an introduced species.
e f
g h i
Fig. 7.1 (continued) e Copiapoa cinerea, Tal Tal; f Cumulopuntia sphaerica, La Serena; g
Maihuenia poeppigii, Las Trancas; h Neoporteria subgibbosa, Fray Jorge; i Oreocereus leu-
cotrichus, Chusmiza (photo credits: e–i A. Moreira-Muñoz)
Clearly the Cactoideae contain most Chilean representatives of the family

(Table 7.1), since it is the richest subfamily of the Cactaceae. Most Chilean cac-
tus genera have their distribution in the highs of Peru, Bolivia, and northern Chile
(12 genera). 4 genera show more restricted distributions to Argentina and Chile.
And 5 genera are endemic to north central Chile (Copiapoa, Eriosyce, Eulychnia,
Neoporteria, and Thelocephala). These genera show an important role in the
building of a northern Chilean area of endemism (see Sect. 4.3).
7.2.1 Cacti Distribution in Chile

Lembcke and Weisser (1979) mapped the distribution of 23 genera encompassing
Chilean Cactaceae, but new collections and systematic treatments (Pinto 2002,
2003; Kiesling 2002; Zuloaga et al. 2008) imply a renewal of this early account.
Only the extreme northern regions have been intensively surveyed and mapped
(Pinto and Kirberg 2009). For this chapter, dot maps for Chilean genera have been
made by means of the collections in existence at the National Herbarium SGO
(Fig. 7.2a). A species richness map is presented as Fig. 7.2b. Distribution maps
Table 7.1 Classification and richness of Chilean cacti (according to Zuloaga et al. 2008)
Chilean Chilean
Subfamily Genus Nº spp. Distribution native spp. endemic spp.
Maihuenioideae Maihuenia 2 Arg, Chile 2 0

Opuntioideae Cumulopuntia 20 Bol, S Peru, N Chile 2 0
Maihueniopsis 16 Andes Peru, Bol, 13 9
Chile, Arg
Miqueliopuntia 1 Chile 1 1
Pterocactus 9 Patagonia Arg, Chile 2 0
Tunilla 9 Andes Peru, Bol, Chile 2 0
Cactoideae Austrocactus 4 Arg, Chile 3 1
Browningia 11 Peru, Bol, Chile, 1 0
Paraguay
Copiapoa 22 N Chile 22 22
Corryocactus 12 Bol, S Peru, N Chile 1 0
Eriosyce 5 Chile 5 5
Eulychnia 6 Chile 6 6
Haageocereus 20 Peru, Chile 2 1
Islaya 4 Peru, Chile 4 3
Lobivia 24 Andes Peru, Bol, 2 0
Chile, Arg
Neoporteria 6 Chile 6 6
Neowerdermannia 2 Peru, Bol, Chile, Arg 1 1
Oreocereus 7 Andes Peru, Bol, 4 1
Chile, Arg
Phyrrocactus 27 Arg, Chile 20 20
Thelocephala 6 Chile 6 6
Trichocereus 25 Chile, Arg, S Peru 7 6
112 88
for each genus within the three subfamilies present in Chile are shown in Figs. 7.3
and 7.4.
With the collection localities, a species richness map was made on the base of a
1 × 1 degree grid (Fig. 7.2b). Chilean Cactaceae are mainly distributed in central
and northern Chile from 18◦ to 38◦ S, concentrated at the coast of the Antofagasta
and Atacama regions, between 25◦ and 31◦ S (Fig. 7.2b). Genera showing a wider
distribution in Chile are Eulychnia, Phyrrocactus, and Trichocereus. In contrast,
other genera like Browningia, Islaya, Neowerdermannia, and Lobivia have a very
restricted distribution in Chile, inferior to one latitudinal degree. Most genera show
discontinuous distributions, but collection gaps cannot be ruled out. This disconti-
nuity is extreme in genera such as Austrocactus, with several collections in Central
Chile and in Patagonia (Fig. 7.3).
Skottsberg (1916) reported the species Pterocactus australis (=Opuntia aus-
tralis) and Pterocactus hickenii (=Opuntia skottsbergii) from Lago Buenos Aires
(Rio Fenix). A recent report (Kiesling 2002) for Pterocactus hickenii on the Chilean
side at 46◦ 30′ S (Chile Chico) confirms the early observations from Skottsberg
(1916). That means that the known southern limit for the family in Chile is at the
Lago General Carrera (46◦ 30′ S). At the limit with Argentina (ca 50◦ 23′ S) most
probably Austrocactus patagonicus can be found (Hoffmann and Walter 2004).
Fig. 7.2 Distribution of a b

Chilean Cactaceae: a
collection localities from
SGO; b species richness map 20!−
30!−
40!− 1–5
6 – 10
11 – 20
50!−
A dubious report of a southernmost occurrence of Maihuenia poeppigii at Torres del

Paine National Park, some 1,400 km south of its distribution, has been rejected by
Leuenberger (2008). That means that aside from the few localities in the Patagonian
steppe at the boundary with Argentina, the Chilean regions Araucanía, Los Lagos,
Aisén and Magallanes, encompassing the area from 38◦ to 56◦ S, have just a few
species according to the Southern Cone Checklist (Zuloaga et al. 2008), and no
collections at all in the National Herbarium SGO (Fig. 7.2a).
7.2.2 Areas of Endemism

The species richness core that appears at the coast between 25 and 31◦ S
(encompassing 11–20 species) can be superficially interpreted as an area of
endemism. This can be tested by means of specific methods: PAE and the
optimality method (see also Sect. 4.3).
33° −
18° −
20° −
23° −
43° − Browningia
18° −
53° − 35° −
23° −
Austrocastus Corryocactus Copiapoa
18° −
20° − 20° −
23° −
Haageocereus
18° −
35° − 35° −
23° −
Eriosyce Eulychnia Islaya
Fig. 7.3 Collection localities of the genera of the Cactoideae

18° −
20° −
23° −
Lobivia
18° − 18° −
35° −
23° − 23° −
Neowerdermannia Neoporteria Oreocereus
20° − 20° − 20° −
35° − 35° − 35° −
Pyrrhocactus Thelocephala Trichocereus
Fig. 7.3 (continued)

20! − 20! − 20! −
35! − 35! − 35! −
Cumulopuntia Maihueniopsis Miqueliopuntia
20! −
35! − 18! −
30! −
23! −
Tunilla
45! −
40! −
50! −
55! −
Pterocactus Maihuenia
Fig. 7.4 Collection localities of the genera of the Opuntioideae and Maihuenioideae (Maihuenia)
ROOT
40 74
A2 7
73 ”
40
A3 7 20 ”
39 40
A4 7
1 58 75
A16 5
49 75
A17 3
58
A17 5
49 58
A18 3
0
A33 3
42 A9 6
29
A10 6
49 54 58
45 A17 4
62 75
A18 4
12 13 59 69
48 71 A10 4
7 17 20 21 26 29 31 56 58 72
A10 5
26 32 55 60
49 58 67 75 A15 4
A16 4
A3 6
3 25 26 35 52 53
51
40 78 A1 6
A2 6
A5 8
40 43
37 53 74
41 A5 7
A6 8
Area a
58
A18 5
Area b
38 A19 4
A20 3 Area c
A20 4
65
A5 5
6 26
A6 5
26
33 36 A3 5
30
A4 5
4 24
A7 5
9 16 63 66
5 14 15 18 22 26 27 33 34 64
17 23 28 31 68 77 A8 5
8 9 19 20 63 70 72
A9 5
44 58
A15 5
26 47 75 75
27
A14 5
59 61
50 ”
A12 5
26 29 46 49 69 ”
A14 4
32 58 76
A13 4
10 26 31 47 26 47
A13 5
2 32 59 61 59 73
A12 4
27 50 58 71
11 16 26 34 76
a 39 41 46 A11 4
27 31 44 50 57 61 75 b
A11 5
Fig. 7.5 Results from PAE: a matrix obtained with Winclada; b map of terminals (quadrats)
representing areas of endemism
7.2.2.1 PAE
Parsimony Analysis of Endemicity (PAE) was applied by means of a matrix with
42 quadrats and the 79 species, in Winclada. 92 trees were obtained, with 137 steps
and CI = 57 (consistence index) and RI = 61 (retention index). The strict consensus
cladogram had 146 steps, CI = 54 and RI = 55. This cladogram (Fig. 7.5a) showed
an initial politomy and 56 geographic synapomorphies, but only 11 were used to
identify areas of endemism because they defined clades based on the presence of
two or more geographic synapomorphies. Three areas of endemism were found,
two of them with a nested pattern. The first area (Fig. 7.5b) includes three quadrats
between 18◦ –21◦ S and 69◦ –70◦ W, including four endemic species. The second
area (Fig. 7.5b) is nested in another wider area; it includes four endemic species,
comprises two quadrats at 24◦ –26◦ S and 70◦ –71◦ W. The third area (Fig. 7.5b)
includes the previous area plus another quadrat at 23◦ –26◦ S and 70◦ –71◦ W, and
added three endemic species (for the species list, see Table 7.2).
7.2.2.2 NDM/VNDM
The optimality method was performed with NDM/VNDM v. 2.6 (Goloboff 2005).
The georeferenced database of Chilean species of Cactaceae from the National
Herbarium SGO consists of 514 records of 79 species, including 473 unique local-
ities. Each locality was superposed to a grid of 1 × 1 degrees. The program was
run saving sets with two or more species, and with scores above 2.0; settings were
the same as for the analysis of endemic genera (Sect. 4.3). The search was per-
formed until the number of sets was shown stable in 25 repetitions with different
random seeds, using edge proportions. Eleven areas of endemism were obtained,
and 33 species were endemic to some area. A consensus was applied to the sets,
with 40% of similarity in species and the option against any of the other areas in the
consensus. The program yielded four consensus areas; some of them overlapping.
The endemic species of each consensus are shown in Table 7.2. Consensus area 0
is retrieved from just 1 set, and showed five endemic species and a score of 5.08. It
includes three quadrats at 18◦ –21◦ S latitude and 69◦ –70◦ W longitude. Consensus
area 1 was obtained from five sets, including 12 endemic species and a score of 5.23
(although two species have a score below 0.5); including 11 quadrats at 27◦ –33◦ S
and 70◦ –72◦ W. Consensus area 2 was built by four sets, had 14 endemic species and
a score of 6.83. It includes seven quadrats at 24◦ –29◦ S and 70◦ –72◦ W. Consensus
area 3 encompassed one set, including three endemic species and a score of 2.52. It
includes nine quadrats at 18◦ –24◦ S and 67◦ –70◦ W (Fig. 7.6).
Both programs usually yield different results (see discussion in Sect. 4.3). PAE
tends to retrieve less areas of endemism than NDM. This last program identified
11 areas of endemism, and therefore consensus had to be applied to the results.
Even the consensuses can retrieve overlapped areas, as was the case. The areas
of endemism identified by PAE are partially overlapped and mostly nested in the
ones from NDM. Area 0 from NDM is identical to Area a from PAE, supported by
4 species (e.g. Browningia candelaris) (Table 7.2). This Area 0 is nested in Area
Table 7.2 Areas of endemism and their supporting species (for NDM and PAE)
Consensus NDM Area PAE
Nº area Species a b c
73°
Browningia candelaris X
20° Corryocactus brevistylus X
0 Haageocereus fascicularis X
Oreocereus hempelianus X
Tunilla soehrensii
Thelocephala napina
Miqueliopuntia miquelii
Copiapoa coquimbana
Maihueniopsis ovata
Neoporteria clavata
1
Neoporteria nidus
Area 0
Area 1
Neoporteria villosa
Area 2 Pyrrhocactus eriosyzoides
Area 3 Pyrrhocactus heinrichianus
Trichocereus coquimbanus
Copiapoa serpentisulcata X X
Eriosyce rodentiophila X X
Thelocephala esmeraldana X X
50°
69° Trichocereus deserticola X X
Thelocephala napina
Copiapoa cinerea X
Pyrrhocactus paucicostatus X
Fig. 7.6 Result from NDM/VNDM 2
Pyrrhocactus taltalensis X
Copiapoa humilis
Copiapoa hypogaea
Copiapoa marginata
Eulychnia saint-pieana
Neoporteria sociabilis
Thelocephala odieri
Maihueniopsis boliviana
Oreocereus leucotrichus
3
Trichocereus atacamensis
3 that encompasses a wide area of the Chilean Altiplano. The gap that appears
in the core of the Atacama is obvious since it has the overall lowest presence of
plant taxa (Fig. 7.2b). At the coast south of the Atacama, both programs retrieve
areas of endemism: areas b and c from PAE are, excepting one quadrat, nested in
NDM’s Area 2, due to the presence of e.g. Eriosyce rodentiophila and Thelocephala
esmeraldana (Table 7.2 and Fig. 7.6). NDM’s Area 2 much more extends to the
South, and partially overlaps with Area 1 that reaches 33◦ S. Remarkably, Areas 2
and 1 retrieved by NDM, are almost perfectly coincident with Area 2 retrieved by
the analysis of the endemic genera (Sect. 4.3). Certainly in the regionalization by
7.3 Notes on Cacti Conservation 209
endemic genera (Fig. 4.9) the Cactaceae genera play an important role, and this is
corroborated here at the species level.
As discussed for the analysis of endemic genera (Sect. 4.3), these analyses are
still at the level of exercises, since they are much dependant on the strength of the
database, the grid size and the geographical origin of the grid, and the settings of
the programs. Also the observation that the areas of endemism do not show sharp
limits and tend to overlap (independent of the programs’ settings) is as valid for the
endemic genera as for the Cactaceae species.
7.3 Notes on Cacti Conservation

Areas of special interest for cactus conservation are highlighted by the richness
and endemism maps (Figs. 7.2, 7.5, and 7.6). The interest that these species gain
from private collectors from all over the world put many of the species at con-
servation risk. At least four species are classified in the red lists (Chap. 6) as
“highly endangered” in the regions Atacama and Coquimbo (26º–32ºS) (Eriosyce
rodentiophila, Maihueniopsis domeykoensis, Pyrrhocactus eriosyzoides, Eulychnia
breviflora), and 18 more species are classified as “vulnerable”. Another 29 species
are considered “not sufficiently known”, mostly due to the lack of precise knowl-
edge about their distribution. Among these are taxa of special interest for improving
collection efforts and population level studies (e.g. Pinto and Moscoso 2004), to
shorten the so-called Wallacean shortfall (Box 6.2). In the extreme north, cacti pop-
ulations are much more affected due to human activities, especially mining and
the extraction for urban gardening in the city of Iquique (Pinto and Kirberg 2009).
Coastal populations at this latitude (18º–21ºS) appear much damaged through long
drought periods (Fig. 7.7), but whether this is a cyclical situation related to El Niño
(Box 7.1) or the sign of a long term desiccation is still a matter of discussion (Schulz
2009; Pinto and Luebert 2009).
a b
Fig. 7.7 a Dead cactus exemplar of Eulychnia iquiquensis at the coast of Tocopilla (22ºS); b dry
exemplar of Puya boliviensis at the same locality (see Pinto and Kirberg (2009))
Box 7.1 Rise and Bloom of the Atacama Desert

Coastal deserts occur in western North- and South America at tropical and
extratropical latitudes, related to the atmospheric circulation (high pressure
belts), and the oceanic circulation (Richter 2001). Along the west coasts,
the oceanic upwelling generates a cool, stable coastal atmosphere, forming
coastal fog deserts like the Atacama, one of the driest ecosystems on earth
(Ezcurra 2006). Nevertheless, the exact time at which the climate in the
Atacama Desert became hyperarid is a topic of much debate with ages rang-
ing from 25 mya (Dunai et al. 2005), 14 mya (Alpers and Brimhall 1988) to 3
mya (Hartley and Chong 2002) (see Box 3.1).
Hartley et al. (2005) proposed that the sedimentary succession in the
Atacama Desert records deposition under an arid to semiarid climate from
the Late Jurassic (150 mya) to the present day. Prior to this, at the Middle
Jurassic, conifers, Bennettitales, ferns and sphenopsids dominated the land-
scape (Herbst and Troncoso 1996) (Sect. 1.2).
These results are in agreement with results presented by Evenstar et al.
(2005) on erosional and depositional surfaces analysed via remote sensing,
and are corroborated by Clarke (2006), who proposed that “the Atacama
Desert is almost certainly the oldest continuously arid region on Earth”.
This is in agreement with the geomorphologic/biogeographic analysis done
by Ochsenius (1999) who proposed the Atacama as being one of several
Triassic-Jurassic palaeodeserts, prior to the break-up of Gondwana. Ochsenius
(1999) could further identify Mesozoic relict floras of the intra-Andean val-
leys from Venezuela to Ecuador, which he described as an Interandean arid
track. Several tracks that connected the xeric natural areas of South America
have been also highlighted by Roig-Juñent et al. (2003, 2006).
Although the Atacama Desert has existed for at least 90 mya, the initial
onset of hyper-aridity was most likely to have developed progressively with
the uplift of the Andes as they reached elevations between 1,000 and 2,000 m
asl (see Box 1.5). The coupling with the intensification of the cold upwelling
current is dated by these authors at 15–10 mya (Middle Miocene). According
to Alpers and Brimhall (1988), the Middle Miocene climatic desiccation of the
Atacama is related to an increase in upwelling intensity as the Antarctic ice
cap became established at approximately 15–13 mya, generating a decrease
in temperature of coastal waters supplied by the ancestral Humboldt Current.
According to Hartley et al. (2005): “Rapid and extreme climatic fluctua-
tions during the Plio-Pleistocene were not sufficient to destabilize the climate
within the Atacama”.
Nevertheless, there is appealing evidence towards Atacamas’s increas-
ing aridity in the Late Pliocene, related to a phase of global cooling, the
rain-shadow effects of the growing Andean Cordillera (Box 1.5), and the
upwelling of the cold Humboldt Current (Hartley and Chong 2002). Hay
et al. (2002) further propose that the global climate system suffered a major
change with the advent and spread of C4 plants during the Late Miocene. The
authors called this a “climate change paradox”: these plants radically altered
the hydrologic cycle, changed atmospheric heat transport mechanisms, and
restructured weathering and erosion processes (Hay et al. 2002).
Climate oscillations occurred in the Pleistocene-Holocene of the Atacama
(Latorre et al. 2002, 2003; Maldonado et al. 2005) (Box 3.1). However, it
appears unlikely that they would have caused a major shift from arid condi-
tions, except locally through supply of both surface and groundwater (Clarke
2006).
Today, the scanty vegetation of the coastal Atacama is highly related to the
humidity available from fog, also known as “camanchaca” (Cereceda et al.
2008) (Fig. 7.8).
Fig. 7.8 Coastal fog supporting the sparse vegetation at Paposo (25ºS) (photo by
A. Moreira-Muñoz)
Humidity is highly restricted to the western slope of the coastal cordillera,

and rarely reaches the interior (Fig. 7.8). Nevertheless, in exceptional years,
the southern Atacama margin around 24◦ and 30◦ S suffers a drastic trans-
formation from the almost complete arid bareness to a full-color landscape,
known as desierto florido “blooming desert” (Muñoz-Schick 1991) (Fig. 7.9).
This highly attractive phenomenon is strictly related to short and intense rain-
falls that reach these latitudes from the south every 5–7 years. This is related
to the ENSO phenomenon (El Niño Southern Oscillation). El Niño effects
are recorded since 1525 (Ortlieb 1994), with a geological and archaeological
evidence of its existence of more than 5,000 years (Dillon 1998; Caviedes
2001). Its impact on the intertropical Pacific coast is related to the genera-
tion of exceptional rainfall on the coast of Peru and Chile. In this century
the most intense of these events were recorded in 1925–1926, 1982–1983,
and 1997–1998 (Dillon and Rundel 1990; Arntz and Fahrbach 1996; Dillon
1998).
Fig. 7.9 Flowering desert: a Cristaria cyanea fields, Carrizal Bajo; b Nolana paradoxa
fields, La Serena; c expecting the next flowering desert at Llanos de Challe (photo credits:
a, b, c A. Moreira-Muñoz)
Remarkably, El Niño-Southern Oscillation (ENSO) related rainfall anoma-

lies in north central Chile seem to be opposite to ENSO-related anomalies
in fog-frequency at 30◦ 30′ S (Garreaud et al. 2008). El Niño years are asso-
ciated with less foggy conditions at Fray Jorge forest (see Box 3.1), and
La Niña1 years produce temperature anomalies associated with higher spring-
time fog frequency. This seems to be very variable along the coast, since the
huge coastal blooming in the period 1997–1998 at 21◦ S (Iquique) was indeed
related to the increasing water content of the fog (Muñoz-Schick et al. 2001).
Recent results from tree ring analysis of the high Andes (Polylepis tarapacana
growing at 16◦ –23◦ S, between 4,000 and 5,200 m asl) show a high correlation
to ENSO, not necessarily to precipitation increase but to a rise in temper-
ature. This suggests nevertheless that the consequences of the phenomenon
reach much further than the coastal environments (Christie et al. 2009).
Relationships between ENSO and phenomena of smaller amplitude and the
low-frequency variability related to the Pacific Decadal Oscillation (PDO)
and the Antarctic Oscillation (AAO) are receiving much interest (Garreaud
et al. 2008). Moreover, the changes associated with ENSO events have shown
important implications for the research in geosciences, especially on agroe-
cosystems, ecosystem restoration, and biodiversity conservation (Holmgren
et al. 2006).
The increasing aridity of the Atacama Desert, associated with the Andes
uplift (Boxes 1.5 and 3.1), seems to have had important effects on the evo-
lution of the Chilean flora, promoting a rapid evolution in several groups
(Stebbins 1952; Solbrig 1976). This seems to be for example the case of
Heliotropium sect. Cochranea. Luebert and Wen (2008) found that isolation
episodes may be the reason for the morphological, ecological and geograph-
ical differentiation among this group. The uplift of the Andes may have
isolated section Cochranea on the western side from other lineages of the
South American Heliotropium on the eastern side of the Andes, promoting
vicariant speciation (Luebert and Wen 2008). According to these authors, the
gradual isolation of the individual species due to climatic fluctuations may
have been helped by geomorphologic events such as mass landslides. A simi-
lar situation could have been responsible for the diversification in Malesherbia
(Gengler-Nowak 2002) and in several Asteraceae genera like Polyachyrus and
Chuquiraga (see Chap. 8).
1 For the counter cycle of El Niño, a better denomination would be “El Viejo”, since “La Niña” is
a meaningless term in relationship with the origin of the concept associated to the occurrence of
warm waters towards Christmas (M. Dillon, pers. comm.)
7.4 Biogeographic Insights: Spatial and Temporal Origins

In spite of the conceptual difficulties in the search for centres of origin (Sect. 3.3),
the center of origin for the Cactaceae has been a constant question for researchers.
Backeberg (1942) proposed that the evolutionary origin of the cacti was in Central
America, with subsequent migrations north and south. Buxbaum (1969) cited both
the Caribbean and central South America as likely areas of origination, due to
the presence of Pereskia and of opuntioid and cactoid lineages that he considered
“ancestral”.
Wallace and Gibson (2002) hypothesized a central Andean origin for the fam-
ily. They assumed that Andean Pereskia, certain Opuntioideae, and the cactoid
Calymmanthium are basal cactus lineages, and all reside in Peru, Bolivia, and
northern Argentina.
Leuenberger (1986), in his monograph about Pereskia, concluded that northwest-
ern South America was a more likely center of origin, proposing a Late Cretaceous
origin of the group. Some recent studies (Hershkovitz and Zimmer 1997; Nyffeler
2002; Edwards et al. 2005), suggest that the cacti can’t be that old because the
sequence divergence between major clades is limited. Edwards et al. (2005) assert
that both Opuntioideae and Cactoideae originated in the central Andean region
of Peru, Bolivia, and northern Argentina. They refer to the Andean orogeny as
the cause of diversification for many plant lineages. Early uplift in the central
Andean region (25–20 mya) is hypothesized to have occurred under a fluctuat-
ing arid/semi-arid climate regime (Chap. 2), which presents a likely scenario for
early cactus diversification. Edwards et al. (2005) further suggest that the place-
ment of Cacteae (an exclusively North American lineage with its center of diversity
in Mexico) among the earliest diverging lineages of Cactoideae, implies relatively
early movements out of South America across the continent.
Centering the origin far away from Africa might explain the poor representation
of cacti in that continent. In fact, all major lineages of the cacti, i.e. Pereskioideae,
Maihuenioideae, Opuntioideae, and Cactoideae, occur mostly or exclusively in
South America. Furthermore, the closest relatives of the cacti from the “portu-
lacaceous cohort” (Applequist and Wallace 2001) have their highest diversity on
continents of the former Gondwana (Hershkovitz and Zimmer 1997). This is gener-
ally taken as evidence that the family Cactaceae originated in South America (e.g.
Buxbaum 1969). Hence, various groups of Pereskia, Opuntioideae, and Cactoideae
supposedly dispersed into Central and North America from their postulated north-
western South American center of origin (Leuenberger 1986). The presence of a
Rhipsalis species in tropical Africa, Madagascar, and Sri Lanka led some authors to
propose that this distribution indicates an old vicariance between South America and
Africa (e.g. Backeberg 1942) or even an origin of the cacti in the Old World (Croizat
1952). This would imply that the cacti originated before the split of the southern con-
tinents during the late Cretaceous and that all other cacti that might have naturally
occurred in Africa went extinct. More recently, however, this distribution pattern
of Rhipsalis baccifera, which is characterized by having very sticky seeds, has been
explained as the result of relatively recent long-distance dispersal by birds (Barthlott
7.5 Cactus Postmodern Evolution 215
1983; Gibson and Nobel 1986; Barthlott and Hunt 1993). This vision is not lacking
controversy: “I consider the bird-dispersal scenario to be as dead in the water as any
rain forest bird that tries to fly the Atlantic. This really leaves only the vicariance
explanation as a viable option. . . Another question must be asked: Why, of all the
cacti that have juicy fruits (and are therefore potentially attractive to birds) should it
be Rhipsalis that has this wide distribution?” (Maxwell 1998).
Since cacti are not well suited for the fossilization process, the fossil record
of the family is very sparse: only several reports related to microfossils exists for
the South American Cactaceae. Mautino (2002) reported two types of Miocene
pollen grains from Tucumán, northern Argentina. According to Barreda et al.
(2007), Cactaceae was one of the important groups that supports the early-
Middle Miocene Neotropical province in the actual Chaco Domain, together with
the Apocynaceae, Aquifoliaceae, Fabaceae, Sapotaceae, Moraceae, Anacardiaceae,
Ulmaceae, Arecaceae, Amaranthaceae (Chap. 1).
In spite the general consensus favouring a spatial origin of cacti in north-
ern South America (Nyffeler 2002), there is still disagreement about the time of
the cacti origin. A Late Cretaceous origin between 65 and 90 mya, relatedto the
breakup of western Gondwana, has been proposed by Gibson and Nobel (1986)
and Mauseth (1990). More recently, on the base of molecular data that show small
amount of sequence divergence, a much more recent origin of cacti in the Late
Palaeogene (30 mya) has been proposed (Hershkovitz and Zimmer 1997; Nyffeler
2002).
Edwards et al. (2005) still suggest a basal split in Cactaceae in a northern clade
comprising Pereskia species and a southern clade comprising also Pereskia species
and Maihuenia and other Cactoideae and Opuntioideae. Edwards et al. (2005) rec-
ognize that their sampling and resolution within Opuntioideae are insufficient to
make inferences about the geographic distribution of its basal members, and recog-
nize recent results that placed its earliest diverging lineages in Chile and Argentina
(Griffith 2004a).
7.5 Cactus Postmodern Evolution

For resolving the question about the geographical origin (if there is something like
that at all) and the time of origin of the family, we have to obtain a good picture about
the phylogeny of the family, including its closest relatives and the morphological
characteristics of the proto-cactus or the “first” cactus (i.e. “what did the first cactus
look like?”, Griffith 2004a).
Phylogenetic cladograms presented by Butterworth and Edwards (2008) still sup-
port the hypothesis of Pereskia representing the earliest members of the cacti. But
the authors recognize that accurate character reconstruction is still challenging.
Regarding the closest family, there has been a long standing view that the
Aizoaceae was the sister family to the cacti, but this was proved erroneous. Studies
of molecular data and vascular anatomy suggest that the closest relatives to the
cacti are the genera Portulaca, Talinum, and Anacampseros from the Portulacaceae
(Hershkovitz and Zimmer 1997; Applequist and Wallace 2001). Another possible
closely related family is the Didiereaceae. Extant Portulacaceae occur in South
Africa, Kerguelen Island and New Zealand. Didiereaceae occur on Madagascar. The
three families show therefore an old Gondwanic relationship.
As expressed by Gorelick 2008: “Good phylogenies matter. If Ceraria
and Portulacaria are most closely related to traditional Didiereaceae than to
Portulacaceae, as appears to be the case (Nyffeler et al. 2008), we may make
erroneous inferences about the evolution of succulence in cacti and related families.
This philosophical approach is not without problems, because it is often difficult to
discern which characters are basal and which are derived, much as it is difficult to
distinguish homology from analogy in evolution” (Gorelick 2008).
The difficulties in discerning the appropriate interpretation of characters are bet-
ter expressed by P. Griffith in two challenging papers. In Griffith’s vision (2004b),
the classical interpretation of cactus evolution is still influenced by the cultural con-
text of the 1500s, when the exotic members of the family were first seen in Europe.
“The striking, but culturally determined, exoticness of the Cactaceae still impacts
our concept of what is relictual and derived for the family” (Griffith 2004b). The
traditional view, codified in the early twentieth century by Britton and Rose, pro-
posed that leafy cacti are primitive or relictual; on the contrary, stem-succulent
cacti seem to be derived. Britton and Rose (1919) proposed the genus Pereskia,
due to its similarity to other woody flowering plants, the nearest cactus relative
to the other families. The most derived are the Cactoideae, exemplified by leaf-
less plants like Trichocereus. The Pereskia-primitive idea has been so influential
that “recent authors have sometimes misinterpreted new data to be consistent with
Britton and Rose, even when the data are more readily interpreted as contradictory”
(Griffith 2004a). That means that the evolutionary interpretations are bounded to
the contextual bias, i.e. the horticultural landscape from traditional botanical gar-
dens and colleges (see also the political aspects of Pereskia taxonomy discussed
by Leuenberger 2008, p 58). Griffith challenges current evolutionary interpretations
proposing that the nearest relatives of Cactaceae were not broad-leaved dicots super-
ficially similar to Pereskia; rather, Cactaceae likely evolved from diminutive, often
geophytic Portulacaceae; and the southern South American Maihuenia, with its
terete, succulent leaves and cushion habit form a deep, subfamilial lineage (Wallace
1995; Griffith 2004b).
As recently summarised by Griffith (2008), there are currently three crucial
lines of discussion: (a) the phylogenetic position of Pereskia; (b) the nature of
the outgroups; (c) the character evolution implied by each: “If Pereskia represent
transitional forms, what do they transition between?” (Griffith 2008, p 37).
The two deepest lineages in Opuntioideae include most species of Maihueniopsis
and all Pterocactus, and are characterized by diminutive, succulents, often of geo-
phytic habit, and leafless at maturity (Griffith and Porter 2009). The authors suggest
that Opuntioideae have originated in west-central South America.
As for other plant groups, the spatial split between several clades was most prob-
ably caused by the Andean uplift and the formation of intracontinental marine basins
during the Cenozoic (Ritz et al. 2007).
References 217
In spite of the efforts done by cacti experts and the advent of molecular data, and
due to the persisting lack of macrofossils and the difficulties in character interpre-
tation, early cactus evolution remains still an important unanswered question. Will
the future offer a fossil encounter of an early cactus outside America, as it was the
case for the first Old World hummingbird fossils recently discovered in European
Oligocene deposits? (Mayr 2004; Louchart et al. 2008).
References
Alpers C, Brimhall GH (1988) Middle Miocene climatic change in the Atacama Desert, northern
Chile: evidence from supergene mineralization at La Escondida. Geol Soc Am 100:1640–1656
Anderson EF (2001) The cactus family. Timber Press, Portland, OR
Applequist WL, Wallace RS (2001) Phylogeny of the portulacaceous cohort based on ndhF
sequence data. Syst Bot 26:406–419
Arntz WE, Fahrbach E (1996) El Niño. Experimento climático de la naturaleza. Causas físicas y
efectos biológicos. Traducción de C Wosnitza-Mendo y J Mendo. Fondo de Cultura Económica,
México
Backeberg C (1942) Cactaceae; Jahrbücher der DKG., Teil 2, Juni 1942: 55, Cactaceae Lindley;
Systematische Übersicht (Neubearbeitung) mit Beschreibungsschlüssel
Barreda VD, Anzótegui LA, Prieto AR, Aceñolaza PG et al. (2007) Diversificación y cambios de
las angiospermas durante el Neógeno en Argentina. In: Archangelsky S, Sánchez T, Tonni EP
(eds) Ameghiniana 50◦ aniversario, Publicación Especial 11: 173–191
Barthlott W (1983) Biogeography and evolution in neo- and paleotropical Rhipsalinae
(Cactaceae) – Sonderb. Naturwiss Verl Hamburg 7:241–248
Barthlott W (2001) Preface. In: Anderson EF, (ed) The cactus family. Timber Press, Portland, OR
Barthlott W, Hunt DR (1993) Cactaceae. In: Kubitzki K, Rohwer JG, Bittrich V, (eds) The families
and genera of vascular plants, Vol 2. Flowering plants: Dicotyledons: Magnoliid, Hamamelid
and Caryophyllid families. Springer, Berlin, Heidelberg, pp. 161–197
Britton NL, Rose JN (1919–1923) The Cactaceae, 4 vols. Carnegie Inst., New York, NY
Butterworth Ch, Edwards EJ (2008) Investigating Pereskia and the earliest divergences in
Cactaceae. Haseltonia 14:46–53
Buxbaum F (1969) Die entwicklungswege der Kakteen in Sudamerika. In: Fittkau EJ, Illies J,
Klinge H, et al. (eds) Biogeogaphy and ecology in South America. W. Junk, The Hague, pp
583–623
Caviedes C (2001) El Niño in history: storming through the ages. University Press of Florida,
Gainesville, FL
Cereceda P, Larrain H, Osses P, Farías M, Egaña I (2008) The spatial and temporal variability of
fog and its relation to fog oases in the Atacama Desert, Chile. Atmos Res 87:312–323
Christie DA, Lara A, Barichivich J, Villalba R, Morales MS, Cuq E (2009) El Niño-Southern
Oscillation signal in the world’s highest-elevation tree-ring chronologies from the Altiplano,
Central Andes. Palaeogeogr Palaeoclimatol Palaeoecol. doi:10.1016/j.palaeo.2007.11.013
Clarke JDA (2006) Antiquity of aridity in the Chilean Atacama Desert. Geomorphology 73:
101–114
Croizat L (1952) Manual of phytogeography, or an account of plant dispersal throughout the world.
Junk, The Hague
Dillon MO (1998) Chasing the 1997/1998 El Niño across Peru. Field Museums Memb Publ 69:2–4
Dillon MO, Rundel PW (1990) The botanical response of the Atacama and Peruvian desert
floras to the 1982–83 El Niño event. In: Glynn PW, (ed) Global ecological consequences
of the 1982–83 El Niño-Southern Oscillation. Elsevier Oceanography Series 52. Elsevier,
Amsterdam, pp 487–504
Dunai TJ, González López GA, Juez-Larré J (2005) Oligocene–Miocene age of aridity in
the Atacama Desert revealed by exposure dating of erosion-sensitive landforms. Geology
33:321–324
Edwards EJ, Nyffeler R, Donoghue MJ (2005) Basal cactus phylogeny: implications of Pereskia
(Cactaceae) paraphyly for the transition to the cactus life form. Am J Bot 92:1177–1188
Evenstar L, Hartley A, Rice C, Stuart F, Mather A, Chong G (2005) Miocene-Pliocene climate
change in the Peru-Chile Desert. 6th international symposium on Andean geodynamics (ISAG
2005, Barcelona), extended abstracts, pp 258–260
Ezcurra E, (ed) (2006) Global deserts outlook. United Nations Environment Programme (UNEP),
Nairobi
Garreaud RD, Barichivich J, Christie DA, Maldonado A (2008) Interannual variability of the
coastal fog at Fray Jorge relict forests in semiarid Chile. J Geophys Res 113:G04011. doi:
10.1029/2008JG000709
Gengler-Nowak KM (2002) Reconstruction of the biogeographical history of Malesherbiaceae.
Bot Rev 68:171–188
Gibson AC, Nobel PS (1986) The cactus primer. Harvard University Press, Cambridge, MA
Goloboff P (2005) NDM/VNDM ver. 2.6. Programs for identification of areas of endemism.
http://www.zmuc.dk/public/phylogeny/endemism
Gorelick R (2008) Early cactus evolution. Haseltonia 14:3–5
Griffith MP (2004a) What did the first cactus look like? An attempt to reconcile the morphological
and molecular evidence. Taxon 53:493–499
Griffith MP (2004b) Early cactus evolution: the postmodern view. Haseltonia 10:3–11
Griffith MP (2008) Pereskia, Portulacaceae, photosynthesis, and phylogenies: implications for
early Cactaceae. Haseltonia 14:37–45
Griffith MP, Porter JM (2009) Phylogeny of Opuntioideae (Cactaceae). Int J Plant Sci 170:107–116
Hartley AJ, Chong G (2002) A late Pliocene age for the Atacama Desert: implications for the
desertification of western South America. Geology 30:43–46
Hartley AJ, Chong G, Houston J, Mather AE (2005) 150 million years of climatic stability:
evidence from the Atacama Desert, northern Chile. J Geol Soc 162:421–424
Hay WW, Soeding E, DeConto RM, Wold CN (2002) The Late Cenozoic uplift – climate change
paradox. Int J Earth Sci 91:746–774
Herbst R, Troncoso A (1996) La tafoflora de Juan de Morales del Jurásico Medio (Formación
Chacarilla), Región de Tarapacá, Chile. Rev Geol Chile 23:3–15
Hershkovitz MA, Zimmer EA (1997) On the evolutionary origins of the cacti. Taxon 46:217–232
Hoffmann AE, Walter HE (2004) Cactáceas en la flora silvestre de Chile, 2nd edn. Ed. Fundación
Claudio Gay, Santiago
Holmgren M, Stapp P, Dickman CR, Gracia C, Graham S, Gutiérrez JR, Hice C, Jaksic F, Kelt DA
et al. (2006) A synthesis of ENSO effects on drylands in Australia, North America and South
America. Adv Geosci 6:69–72
Kiesling R (2002) Pterocactus (Cactaceae), new record for the Chilean flora. Gayana Bot 59:61–63
Kraus R (1995) The environmental conditions of Cactaceae in Chile. Haseltonia 3:110–129
Latorre C, Betancourt JL, Rylander K, Quade J (2002) Vegetation invasions into absolute desert; a
45 000 yr rodent midden record from the Calama–Salar de Atacama basins, northern Chile (lat
22◦ –24◦ S). Geol Soc Am Bull 114:349–366
Latorre C, Betancourt JL, Rylander K, Quade J, Matthei O (2003) A vegetational history from the
arid prepuna of northern Chile (22◦ –23◦ S) over the last 13,500 years. Paleogeogr Paleoclimatol
Paleoecol 194:223–236
Lembcke H, Weisser PJ (1979) The distribution of the genera of Chilean Cactaceae. Aloe 17:9–26
Leuenberger BE (1986) Pereskia (Cactaceae). Mem NY Bot Gard 41:1–141
Leuenberger BE (1997) Maihuenia—monograph of a Patagonian genus of Cactaceae. Bot Jahrb
119:1–92
Leuenberger BE (2008) Pereskia, Maihuenia, and Blossfeldia—taxonomic history, updates, and
notes. Haseltonia 14:54–93
References 219
Louchart A, Tourment N, Carrier J, Roux T, Mourer-Chauviré C (2008) Hummingbird with

modern feathering: an exceptionally well-preserved Oligocene fossil from southern France.
Naturwissenschaften 95:171–175
Luebert F, Wen J (2008) Phylogenetic analysis and evolutionary diversification of Heliotropium
sect. Cochranea (Heliotropiaceae) in the Atacama Desert. Syst Bot 33:390–402
Maldonado A, Betancourt JL, Latorre C, Villagrán C (2005) Pollen analyses from a 50 000-yr
rodent midden series in the southern Atacama Desert (25 degrees 30′ S). J Quat Sci 20:493–507
Marticorena C (1990) Contribución a la estadística de la flora vascular de Chile. Gayana Bot 47:
85–113
Mauseth JD (1990) Continental drift, climate and the evolution of cacti. Cact Succ J 62:302–308
Mauseth JD, Kiesling R, Ostolaza C (2002) A Cactus Odyssey: journal in the wilds of Bolivia,
Peru, and Argentina. Timber Press, Oregon
Mautino LR (2002) Cactaceae en la Formación San José (Mioceno Medio), Tucumán, Argentina.
8º Congreso Argentino de Paleontologia y Bioestratigrafía, Resúmenes: p. 26
Maxwell R (1998) The Rhipsalis Riddle – or the day the cacti came down from the
trees. http://rhipsalis.com/maxwell.htm (originally appeared in the New Zealand Cactus and
Succulent Journal)
Mayr G (2004) Old world fossil record of modern-type hummingbirds. Science 304:861–864
Muñoz-Schick M (1991) Flores del Norte Chico, 2nd edn. Dirección de Archivos, Bibliotecas y
Museos, Santiago
Muñoz-Schick M, Pinto R, Mesa A, Moreira-Muñoz A (2001) Oasis de neblina en los cerros
costeros del sur de Iquique, región de Tarapacá, Chile, durante el evento El Niño 1997–1998.
Nyffeler R (2002) Phylogenetic relationships in the cactus family (Cactaceae) based on evidence
from trnK/matK and trnL-trnF sequences. Am J Bot 89:312–326
Nyffeler R, Eggli U, Ogburn M, Edwards EJ (2008) Variations on a theme: repeated evolution of
succulent life forms in the Portulacineae (Caryophyllales). Haseltonia 14:26–36
Ochsenius C (1999) The neotropical aridity. CCOS PCSS Geowissenschaftliche Beiträge, Baden-
Württemberg
Ortlieb L (1994) Las mayores precipitaciones históricas en Chile Central y la cronología de eventos
ENOS en los siglos XVI–XIX. Rev Chil Hist Nat 67:463–485
Pinto R (2002) Lobivia ferox Britton et Rose (Cactaceae): new record for the Chilean flora. Gayana
Bot 59:65–72
Pinto R (2003) Maihueniopsis nigrispina (Cactaceae, Opuntioideae): nuevo registro para la flora
chilena. Chloris Chilensis 6. http://www.chlorischile.cl
Pinto R, Moscoso D (2004) Estudio poblacional de Echinopsis atacamensis (Cactaceae) en la
Región de Tarapacá (I), norte de Chile. Chloris Chilensis 2. http://www.chlorischile.cl
Pinto R, Kirberg A (2009) Cactus del Extremo Norte de Chile. AMF, Santiago
Pinto R, Luebert F (2009) Datos sobre la flora vascular del desierto costero de Arica y Tarapacá,
Chile, y sus relaciones fitogeográficas con el Sur de Perú. Gayana Bot 66:28–49
Richter M (2001) Vegetationszonen der Erde. Klett-Perthes, Ulmer, Gotha and Stuttgart
Ritz CM, Martins L, Mecklenburg R, Goremykin V, Hellwig FH (2007) The molecular phylogeny
of Rebutia (Cactaceae) and its allies demonstrates the influence of paleogeography on the
evolution of South American mountain cacti. Am J Bot 94:1321–1332
Roig-Juñent S, Flores GE, Mattoni C (2003) Consideraciones biogeográficas de la Precordillera
(Argentina), con base en artrópodos epígeos. In: Morrone JJ„ Llorente Bousquets J, (eds) Una
Perspectiva Latinoamericana de la Biogeografía. Las Prensas de Ciencias, Facultad de Ciencias,
UNAM, México, DF, pp 275–288
Roig-Juñent S, Domínguez MC, Flores GE, Mattoni C (2006) Biogeographic history of South
American arid lands: a view from its arthropods using TASS analysis. J Arid Environ
66:404–420
Schulz N (2009) Loma-Formationen der Küsten-Atacama/Nordchile unter besonderer
Berücksichtigung rezenter Vegetations- und Klimaveränderungen. PhD thesis, Geographical
Institute, Erlangen-Nürnberg University
Schumann K (1898) Gesamtbeschreibung der Kakteen. Neumann, Neudamm

Skottsberg C (1916) Botanische Ergebnisse der schwedischen Expedition nach Patagonien und
dem Feuerlande 1907–1909. V. Die Vegetationsverhältnisse längs der Cordillera de los
Andes südlich von 41 Grad südlicher Breite. K Sven Vetenskapsakademien Handl 56:1–366.
Almqvist & Wiksell, Stockholm
Solbrig O (1976) The origin and floristic affinities of the South American temperate, desert and
semidesert regions. In: Goodall W, (ed) Evolution of desert biota. University of Texas Press,
Austin, TX, pp 7–49
Stebbins GL (1952) Aridity as a stimulus to plant evolution. Am Nat 86:33–44
Wallace RS (1995) Molecular systematic study of the Cactaceae: using chloroplast DNA variation
to elucidate cactus phylogeny. Bradleya 13:1–12
Wallace RS, Gibson AC (2002) Evolution and systematics. In: Nobel PS, (ed) Cacti: biology and
uses. University of California Press, Berkeley, CA, pp 1–21
Sur: Argentina, Sur de Brasil, Chile, Paraguay y Uruguay. Monogr Syst Bot, 107: 1715–1830
Missouri Botanical Garden
Chapter 8
Asteraceae, Chile’s Richest Family
Abstract Being the richest family worldwide, the Asteraceae is also the richest
Chilean family at the genus and species level. According to the most up to date
knowledge, it encompasses 123 genera and 838 native species, that pertain to 20
different tribes. The 123 genera have been classified in 7 floristic elements and
9 generalized tracks and the geographical evolution of the family is discussed.
Analysis of endemism has been undertaken by means of the program NDM/VNDM,
resulting in 6 areas of endemism, some of them overlapping in Central Chile.
Finally, several aspects about the conservation of the Chilean Asteraceae are dis-
cussed, considering also the high degree of alien invasive taxa and the gaps in the
distributional knowledge. An exercise towards bridging these gaps is undertaken by
means of niche modeling of Mutisia species.
The Asteraceae or Compositae is the richest vascular plant family in the world, with
over 1,600 genera and 23,000 species (Anderberg et al. 2007). Only the Orchidaceae
(ca 880 genera and 22,000 species) and the Fabaceae (730 genera and 19,000
species) come close to these numbers (Stevens 2001 and onwards).
Members of the Asteraceae are found in almost every environment and on all
continents, except Antarctica, but the distribution at the genus and species level is
very heterogeneous and with high levels of endemism at different scales (Funk et al.
2005). Asteraceae representatives adopt almost every life-form: herbs, succulents,
lianas, epiphytes, trees, or shrubs (Figs. 8.1 and 8.2 ). They are mainly characterized
by the florets grouped in capitula, the fruit being a cypsela mostly with a pappus
(Fig. 8.3).
8.1 Classification of Chilean Asteraceae

Systematics of the Asteraceae is a very dynamic field. In the earlier stages of plant
taxonomy, Cassini (1819) recognized 19 tribes, while Bentham (1873) proposed a
system of 13 tribes. After many vicissitudes, currently under the predominance of
molecular systematics, the classification has been improved to include 10 subfam-
ilies and 39 tribes (Funk et al. 2009). The classification at the genus level is also

DOI 10.1007/978-90-481-8748-5_8, ⃝
222 8 Asteraceae, Chile’s Richest Family
C
b
d e
Fig. 8.1 Chilean Asteraceae: a Triptilion spinosum, Antuco; b Ophryosporus triangularis, Cuesta
Buenos Aires; c Chaetanthera pusilla, Cerro Provincia; d Mutisia cana, Catemu; e Haplopappus
rengifoanus, Cuesta Buenos Aires (photo credits: a–e A. Moreira-Muñoz)
very dynamic: every year several new genera are described and many more are res-
urrected or moved into synonymy (Bremer 1994). For the Chilean flora, one of the
most recently resurrected genera is Ocyroe (Bonifacino 2008).
The classification at the tribe level is far from having reached consensus, but an
attempt towards a synopsis has been proposed by Funk et al. (2005), (2009), as a
phylogenetic supertree based greatly on the last findings via molecular systematics
(Fig. 8.4).
The Chilean genera have been organized here under this scheme, following
Moreira-Muñoz and Muñoz-Schick (2007), updated by means of Anderberg et al.
(2007), the Southern Cone checklist (Zuloaga et al. 2008), and recent monographic
treatments (e.g. Chiliotrichum s.s., Bonifacino 2009). The Asteraceae is the richest
family in the Chilean flora (see Sect. 2.2). According to the current update, the fam-
ily is composed in Chile by 123 native genera and ca. 838 native species (Table 8.1).
8.2 Floristic Elements of Chilean Asteraceae 223
a b
Fig. 8.2 Chilean Barnadesieae (Asteraceae): a Dasyphyllum diacanthoides, Antuco; b

Chuquiraga oppositifolia, Juncal (photo credits: a, b A. Moreira-Muñoz)
A high number of these species (374 or 45%) are endemic to Chile (Table 8.1).
Regarding the tribe composition, 20 of 39 currently recognized tribes (according to
Funk et al. 2009) are recorded in Chile, i.e. half of the global diversity at the tribus
level is present in Chile. Best represented in Chile are the Astereae (20 genera),
Nassauvieae and Gnaphalieae (15) (Table 8.1).
At the species level the best represented tribe is by far the Senecioneae, due to
the presence of Senecio (ca 224 species, 103 of them endemic), the species-richest
Chilean plant genus (Sect. 2.2) (Teillier and Marticorena 2006). Other Compositae
species-rich tribes in Chile are the Astereae (173 species) and Nassauvieae (114);
species-rich genera are also Haplopappus (54 species), Leucheria (42), Baccharis
(40), and Chaetanthera (38).
The relative proportion of the genera in the respective tribes gives us another
perspective: the Chilean genera from the Onoserideae, Mutisieae, and Nassauvieae
represent 67, 57 and 60% of the generic richness respectively, while Chilean
Mutisieae and Nassauvieae represent one third of the species richness of these
tribes (Table 8.1). In contrast, tribes such as Cardueae, Anthemideae, Inuleae or
Eupatorieae, have a very sparse representativeness in Chile.
8.2 Floristic Elements of Chilean Asteraceae

Similar to the analysis done for the whole flora at the genus level (Chap 3) a specific
analysis of floristic elements and generalized tracks was undertaken for the genera
of the Asteraceae. As for the flora in general (Sect. 3.2), it seems that disjunction is
Fig. 8.3 Asteraceae fruits: a

female capitulum, Baccharis
linearis; b male capitulum, B.
linearis; c achene with
pappus, Pleocarphus
revolutus; d achene with
pappus, Taraxacum officinale;
e achene with pappus,
Hypochaeris incana
(illustrations from E. Sierra,
in Muñoz Pizarro 1966))
a b
d
c e
the norm under the Chilean Asteraceae. Floristic elements and generalized tracks of
Chilean Asteraceae genera are synthesized in Tables 8.2, 8.3 and Fig. 8.5.
Most genera within the Chilean Asteraceae (53 genera, 43%) can be classi-
fied into a Neotropical element, further composing four different tracks: the “wide
Neotropical”, “wide Andean”, “Central Andean”, and “South Amazonian” tracks.
The second most important element in the Chilean Asteraceae is the Antitropical
element, which reaches 21% with 26 genera. This Antitropical element can be
assigned to a “wide Antitropical track” that connects Chile with temperate North
America and Eurasia (9 genera, e.g. Adenocaulon, Hypochaeris) and a “Pacific-
Antitropical track” that includes most genera disjunct between central Chile and
western North America (13 genera, e.g. Bahia, Blennosperma, and Gutierrezia).
Four genera form a “circum-Pacific track”, showing a distribution in western North
America, South America, and Australasia (Flaveria, Gochnatia, Microseris, and
Soliva).
Goodeniaceae
Calyceraceae
Barnadesieae
Onoserideae
Nassauvieae
Mutisieae
Gochnatieae
Cardueae
Cichorieae
Senecioneae
Gnaphalieae
Astereae
Anthemideae
Inuleae
Helenieae
Coreopsideae
Tageteae
Bahieae
Heliantheae
Madieae
Perityleae
Eupatorieae
Fig. 8.4 Phylogenetic relationships of Chilean Asteraceae tribes. Goodeniaceae and Calyceraceae
appear as outgroups (adapted from Funk et al. (2005, 2009))
The South-temperate element (11 genera), encompasses genera distributed only

in central/southern Chile and Argentina (e.g. Belloa, Eriachaenium, and Triptilion).
Acrisione is a subendemic Chilean genus with a couple reported localities in
Argentina (Cabrera 1975).
The pantropical element is composed by 9 genera (e.g. Achyrocline, Centipeda,
Mikania) distributed along the world’s tropical belt.
The australasiatic element comprises also a generalized track that represents the
disjunct distribution between Chile and Australasia, as shown by the four genera
Abrotanella, Lagenophora, Leptinella, and Trichocline.
The Chilean endemic element is composed of 16 genera. Under those, 6
genera are endemic to the oceanic islands off Chile: Lycapsus (Perityleae)
and Thamnoseris (Cichorieae) are restricted to the Desventuradas Islands (San
Félix and San Ambrosio); Dendroseris (Cichorieae), Robinsonia (Senecioneae),
Centaurodendron, and Yunquea (both Cardueae) are restricted to the Juan Fernández
archipelago (see Chap 5). On the other hand, 10 genera are endemic to mainland
Chile, pertaining mostly to the Nassauvieae (Calopappus, Leunisia, Marticorenia,
226
Table 8.1 Summary of Chilean Asteraceae: tribes present in the Chilean flora, tribal distribution, global numbers of genera and species, Chilean numbers, and
relative presence in Chile (updated from Moreira-Muñoz and Muñoz-Schick 2007; Anderberg et al. 2007; Zuloaga et al. 2008; Funk et al. 2009). For detailed
genera and species numbers see Appendix
Gen. n◦ Gen. n◦ Spp. n◦ Spp. n◦ Endemic

Tribe Distribution global Chile % global Chile % Chilean spp.
Barnadesieae South America 9 3 33.3 91 9 9.9 3

Onoserideae Southern Andes 6 4 66.7 52 5 9.6 3
Nassauvieae Neotropics 25 15 60.0 315 114 36.2 39
Mutisieae Neotropics, few in Asia, Africa, Australia 14 8 57.1 200 71 35.5 29
Gochnatieae South America, few Mexico, USA 4 1 25.0 88 1 1.1 1
Cardueae Mediterranean, Asia, N hemisphere 72 3 4.2 2, 400 10 0.4 10
Cichorieae Temperate N hemisphere, few S Am, Islands 93 9 9.7 1, 400 55 3.9 31
Senecioneae Cosmopolitan 150 9 6.0 3, 500 258 7.4 118
Gnaphalieae Cosmopolitan, esp. Australia, S Africa, S Am 190 15 7.9 1, 240 74 6.0 27
Astereae Cosmopolitan 222 20 9.0 3, 100 174 5.6 88
Anthemideae Cosmopolitan, Mediterranean, Asia, S Africa 111 4 3.6 1, 800 6 0.3 0
8
Inuleae Subcosmopolitan, Eurasia, S Africa, S Am 66 2 3.0 687 2 0.3 0

Helenieae Anfitrops, esp. W USA 13 1 7.7 120 7 5.8 6
Coreopsideae North/South Am, few Australia, India 30 3 10.0 550 5 0.9 1
Tageteae Neotropics, esp. W USA 32 2 6.3 270 4 1.5 0
Bahieae Neotropics, esp. W USA, few Africa, Pacific 20 2 10.0 83 3 3.6 1
Heliantheae Pantropics, esp. Neotrops 114 10 8.8 1, 461 16 1.1 7
Madieae Anfitropics, esp. W USA 36 3 8.3 200 4 2.0 1
Perityleae SW USA, Chile/Peru, Desventuradas Is 7 3 42.9 84 4 4.8 2
Eupatorieae Subcosmop 182 6 3.3 2, 200 16 0.7 7
Total 123 838 374
Asteraceae, Chile’s Richest Family
Table 8.2 The generic composition of floristic elements and generalized tracks in the Chilean
Asteraceae (updated from Moreira-Muñoz and Muñoz-Schick (2007), by permission of John Wiley
and Sons)
N◦
Element Track Definition genera Genera
1. Pantropical Tropics 9 Achyrocline, Centipeda,

Conyza, Cotula, Mikania,
Pluchea, Sigesbeckia,
Spilanthes, Wedelia
2. Australasiatic 2.1 Australasiatic Southern 4 Abrotanella, Lagenophora,
hemisphere: Leptinella, Trichocline
America,
Australasia
3. Neotropical 3.1 Wide NW USA, 17 Ageratina, Ambrosia,
Neotropical México to Baccharis, Chaptalia,
Chile Coreopsis, Erechtites,
Gamochaeta, Grindelia,
Helenium, Heterosperma,
Schkuhria, Stevia,
Tagetes, Trixis,
Verbesina, Viguiera,
Villanova
3.2 Wide Andean Costa Rica, 9 Aristeguietia, Chuquiraga,
Colombia to Cuatrecasasiella,
Chile Diplostephium,
Luciliocline, Mutisia,
Perezia, Werneria,
Xenophyllum
3.3 Central Altiplano Perú, 18 Aphylloclados,
Andean Chile, Bolivia, Chaetanthera,
Argentina Chersodoma,
Haplopappus, Helogyne,
Laennecia, Leucheria,
Lophopappus, Mniodes,
Nardophyllum,
Nassauvia, Ocyroe,
Pachylaena,
Parastrephia, Plazia,
Polyachyrus, Proustia,
Urmenetea
3.4 South Andes and 9 Chevreulia, Dasyphyllum,
Amazonian southern Facelis, Lucilia,
Brazil Micropsis, Noticastrum,
Ophryosporus, Picrosia,
Tessaria
4. Antitropical 4.1 Wide Cool regions 9 Adenocaulon, Anaphalis,
antitropical both Antennaria, Artemisia,
hemispheres Erigeron, Hieracium,
Hypochaeris, Solidago,
Taraxacum
N◦
Element Track Definition genera Genera
4.2 Circum- Temperate 4 Flaveria, Gochnatia,

Pacific regions Microseris, Soliva
N Am, S Am
and Australasia
4.3 Pacific- Chile – W USA 13 Agoseris, Amblyopappus,
antitropical Bahia, Blennosperma,
Encelia, Flourensia,
Gutierrezia, Lasthenia,
Madia, Malacothrix,
Perityle, Psilocarphus,
Symphyotrichum
5. South- Temperate 11 Acrisione, Belloa,
temperate Chile/Argentina Brachyclados,
Chiliotrichum,
Doniophyton,
Eriachaenium,
Gamochaetopsis,
Lepidophyllum,
Macrachaenium,
Notopappus, Triptilion
6. Endemic Continental 10 Calopappus, Guynesomia,
Chile Gypothamnium,
Leptocarpha, Leunisia,
Marticorenia,
Moscharia, Oxyphyllum,
Pleocarphus, Podanthus
Chilean islands 6 Centaurodendron,
Dendroseris, Lycapsus,
Robinsonia,
Thamnoseris, Yunquea
7. Cosmopolitan Worldwide, 4 Bidens, Centaurea,
most continents Gnaphalium, Senecio
Table 8.3 Floristic elements

of Chilean Asteraceae genera Element N◦ Genera %
Pantropical 9 7.3
Australasiatic 4 3.3
Neotropical 53 43.1
Antitropical 26 21.1
South-temperate 11 8.9
Endemic 16 13.0
Cosmopolitan 4 3.3
123 100.0
8.3 Biogeographic regionalization of the Chilean Asteraceae 229
9
17
4
13
27 9
Australasiatic track Wide neotropical track

Circum-Pacific track Pacific antitropical track
Wide antitropical track Wide and central Andean tracks
Panbiogeographic nodes South-Amazonian track
Fig. 8.5 Generalized tracks of Chilean Asteraceae genera (updated from Moreira-Muñoz and
Muñoz-Schick (2007), by permission of John Wiley and Sons)
Moscharia, Oxyphyllum, Pleocarphus). Heliantheae are represented by Podanthus

and Leptocarpha; Onoserideae by Gypothamnium; and Astereae by Guynesomia.
Dot maps for the distribution of these endemic genera have been published in
Moreira-Muñoz and Muñoz-Schick (2007).
Finally, the cosmopolitan element includes only four widespread genera: Bidens
(Coreopsideae), Centaurea (Cardueae), Gnaphalium (Gnaphalieae), and Senecio
(Senecioneae).
The proportion of floristic elements in the Chilean Asteraceae is similar to that
of the whole vascular flora (compare with Table 3.2). Remarkably is the highest
presence of neotropical genera (43%) within the Asteraceae, resting from the cos-
mopolitan genera which in the whole flora reach a 17% while in the Asteraceae
corresponds only to a 3%.
8.3 Biogeographic regionalization of the Chilean Asteraceae

The latitudinal range of distribution of each genus of the Chilean Asteraceae has
been already illustrated in Moreira-Muñoz and Muñoz-Schick (2007), on the base
of the collections at the National Herbarium (SGO).
There is no obvious relationship between the geographic relationships and the
distribution in Chile along the latitudinal gradient: while the few australasiatic gen-
era tend to occupy more austral localities than the neotropical genera, there is one
exception: Trichocline, which reaches northernmost Chile. In contrast, not all the
neotropical genera just reach the subtropical latitudes of Central Chile: some of them
approach southernmost Chile, such as Gamochaeta, Nardophyllum, Nassauvia, or

the rich and widespread genus Baccharis. Apart from Baccharis, the most widely
distributed genera are Perezia, Leucheria, Conyza, Gamochaeta and Senecio, that
occupy the whole Chilean latitudinal extent from Parinacota (17◦ 35′ S) to Cabo de
Hornos (56◦ S). Of course we find genera with a more restricted distribution, e.g.
most genera pertaining to the Central Andean track reach the southern Atacama and
not disperse to the South. Chilean endemic genera are restricted to Central Chile,
such as Calopappus, Leunisia, Marticorenia, Moscharia. Only Leptocarpha reaches
more southern latitudes at 45◦ S.
The generic richness plotted on a grid of 1 × 1 degrees shows three centres
of richness (Fig. 8.6a): a reduced one at 18◦ –19◦ S in the Altiplano encompassing
many Neotropical genera shared with Peru and NW Argentina; a second and biggest
20°−
30°−
40° −
1 − 20 Area 0
21 − 40 Area 1
41 − 60 Area 2
Area 3
Area 4
Area 5
50°
Fig. 8.6 a Generic richness

of Chilean Asteraceae on a
1 × 1 degree grid; b areas of
endemism (based on
consensus) obtained by
NDM/VNDM
centre at 31◦ –35◦ S, encompassing many Chilean endemic genera; and a third centre
at 36◦ –38◦ S (coast). This last one seems to be an extension of the Central Chilean
centre, separated most probably due to collection gaps. The number of genera shows
an obvious decline in the Atacama Desert and South of 38◦ S, entering the temperate
zone that suffers a general reduction in the plant generic richness. This figure has
to be taken only as referential, since collection gaps cannot be ruled out, and much
effort needs to be done on the improvement of the georeferencing of the Herbaria
collections and the acquirement of new field data.
Reiche (1905) made, with the knowledge of his time, a biogeographic analysis
of the Chilean composites, further proposing the first biogeographic regionaliza-
tion for the Chilean flora (see also Sect. 4.3). He proposed five floristic regions and
five subregions based on the endemic or restricted genera and species of the family
(Table 8.4 and Fig. 8.7). Reiche further drew a second map in which he proposed
the dispersal routes for the Chilean Compositae (Fig. 8.7).
Reiche considered both the genera present in certain zones, as well as the genera
that were absent, i.e. the turnover in geographic distribution. This is the essence of
modern methods of endemism analysis like PAE or NDM/VNDM (Goloboff 2005)
(see Sects. 4.3 and 7.2). Remarkably, Reiche made a similar analysis intuitively and
with a much more limited distributional knowledge than we manage today. To repli-
cate the analysis, a database with 19,900 records of 116 genera of the Asteraceae
native to continental Chile was built. Collections pertain to the Herbaria SGO and
CONC (C. Marticorena pers. comm.). Records were checked regarding georefer-
ence, corresponding to 15,809 unique localities that were overlapped by a grid of
quadrats of 1◦ × 1◦ degree covering Chile.
The optimality method was carried out with NDM/VNDM v. 2.6 with the same
options as for the endemic genera and Cactaceae species (Sects. 4.3 and 7.2). The
Table 8.4 Floristic regions of the Chilean Compositae (Reiche 1905)
Regions and subregions Extention Characteristic taxa
I North Coastal 22◦ –29◦ S coast Gypothamniun, Oxyphyllum,

Amblyopappus
II North Andes 18◦ –31◦ S Andes Diplostephium, Schkuhria, Plazia,
Werneria
III Central and southern Andes 31◦ –52◦ S Andes Nassauvia, Leucheria, Perezia, Mutisia
(A) Coquimbo to Chillán 31◦ –36◦ S Andes Leucheria, Perezia
(B) Chillán to Magallanes 36◦ –52◦ S Andes Adenocaulon, Leucheria, Baccharis
magellanica
IV Central coast 30◦ –36◦ S Chaptalia, Moscharia, Triptilion
(A) Coquimbo to Maule 30◦ –36◦ S Absence of Polyachyrus
(B) Maule to Concepción 36◦ –37◦ S Dasyphyllum diacanthoides
(C) Concepción to the south 37◦ –56◦ S Lagenophora, Leucheria
V Magallanes and Tierra del 52◦ –56◦ s Chiliophyllum fuegianum, Artemisia
Fuego magellanica
a b
Fig. 8.7 Biogeographic maps of Chilean Asteraceae, from Reiche (1905): a floristic regions;
b migration routes
search was performed until the number of sets was shown stable with different
random seeds, using edge proportions.
16 areas of endemism were obtained, and 41 genera were endemic to some area.
A consensus was further applied to the obtained sets, with 10% of similarity in
species (genera) and the option against any of the other areas in the consensus
(Fig. 8.6b).
This resulted in 6 consensus areas, some of them overlapping. The consensus
area with the lower score has 2.27, and the higher has 5.88 (this score is relative,
considering that 1 is the highest score a genus can get, meaning it is found in all
quadrats of the area of endemism: i.e. a consensus built by 3 species (genera) can
reach maximally a score of 3).
Consensus Area 0 was obtained from two sets, showing four endemic genera and
a score of 2.3. It includes 10 quadrats between 28◦ and 33◦ S. Genera supporting this
area are Brachyclados, Flourensia, Guynesomia and Pleocarphus.
Consensus Area 1 was obtained from seven sets, built by the distribution of 13
genera and having a score of 3.81 (that is relatively low for a total of 13 genera). It
includes 25 quadrats between 29◦ and 39◦ S. Its supporting genera are Ageratina,
Anaphalis, Aristeguietia, Blennosperma, Calopappus, Chaptalia, Gochnatia,
Marticorenia, Micropsis, Mikania, Moscharia, Podanthus and Psilocarphus.
Consensus Area 2 is built by one set, made up of 3 genera and getting a score of
2.5 (that is relative high for just 3 genera). It encompasses the whole country’s latitu-
dinal extent, including 121 quadrats between 17◦ and 56◦ S. The widespread genera
supporting this area are Baccharis, Perezia and Senecio. This Area is included in the
map as a blank base (Fig. 8.6b).
Consensus Area 4 was obtained from 4 sets, had 13 genera and a score of
5.63 including 13 quadrats in the northernmost Chilean Altiplano between at 17◦
and 24◦ S. This area has a rather high score. Its genera are Chersodoma, Cotula,
Coreopsis, Diplostephium, Helogyne, Heterosperma, Lophopappus, Luciliocline,
Mniodes, Parastrephia, Pluchea, Trixis and Xenophyllum.
Consensus Area 5, obtained from one set, had 3 genera. It shows a score of
2.83 and includes six quadrats between 23◦ and 28◦ S. Its genera are Erechtites,
Gypothamnium and Oxyphyllum.
Consensus Area 3 was set aside in the final result, because the genera got very
low scores (< 0.5). The area included 22 quadrats between 32◦ and 42◦ S, and the
genera were Belloa, Chevreulia, Dasyphyllum, Lucilia and Noticastrum.
Taking account of the accumulation of knowledge in 100 years, the areas of
endemism presented here logically differentiate from the proposal by Reiche (1905),
but there are a couple of remarkable similarities: consensus area 5 is highly coher-
ent with the North coastal floristic region (Table 8.4), sustained by the genera
Oxyphyllum and Gypothamnium. Also Area 4 was recognized by Reiche as a “North
Andean” region, although not so strictly limited as in the present results (Fig. 8.6b).
Reiche’s “Central and Southern Andes” and “Central Coastal” regions are partly
consistent with Areas 0 and 1. Reiche proposed a sharp limit between coastal and
Andean units, which are not retrieved by NDM. This can be an effect of the scale
and origin of the grid and that can only be overruled with better distributional data.
The low richness of Magallanes does not allow recognition of a specific floristic
region, as proposed by Reiche.
Future analyses will have to include not only genera but species, and especial
attention has to be paid to the adequate taxonomy of the taxa and further increase in
distributional knowledge.
8.4 Asteraceae Evolutionary Biogeography

After analyzing the distribution of the genera and visualizing the migration routes
for the Compositae (Fig. 8.7b), Reiche reached the conclusion that only an old cir-
cumpolar continent connecting South America with New Zealand could be the cause
of the close floristic relationships between the genera that composed his “antarctic
element” (Reiche 1905). Impossible to be known to Reiche, only during the 1990s
several authors proposed that the family seems to have an ancient origin in the
South Hemisphere related to the Gondwana break-up (Bremer 1993; Bremer and
Gustafsson 1997). DeVore and Stuessy (1995) proposed a Southern Hemisphere
origin for the three families Goodeniaceae, Calyceraceae, and Asteraceae, with
Goodeniaceae separating from Calyceraceae–Asteraceae when America got isolated
from Australia/Antarctica (in the Early Eocene, 43–53 Ma) (Boxes 1.3 and 8.1).
Box 8.1 Goodeniaceae and Calyceraceae in the Familiar

Relationships of the Asteraceae
Molecular analyses suggest that the closest families to the Asteraceae are
the Goodeniaceae and Calyceraceae. The Goodeniaceae show a predominant
distribution in Australia, New Zealand and the Pacific. The distribution of
Selliera, on the Australian Continent, Pacific islands, and Chile, is particu-
larly interesting. The Calyceraceae are completely restricted to southern South
America. From the 6 genera and 60 species, 5 genera (Boopis, Calycera,
Gamocarpha, Moschopsis, and Nastanthus) and 30 species are to be found in
Chile (Fig. 8.8). Fossil pollen from the Calyceraceae related to Gamocarpha
type has been recently reported by Palazzesi et al. (2010) from Miocene
deposits (Chubut, Patagonia).
The fossil record of the family is sparse and consists mostly of dispersed pollen
grains. Turner (1977) suggested a Cretaceous origin for the Asteraceae, while the
revision made by Graham (1996) appears to indicate an Eocene origin for the
Asteraceae in South America. Eocene fossil Asteraceae pollen has been found more
recently also in Palaeocene/Eocene (55 mya) deposits in South Africa (Zavada
8.4 Asteraceae Evolutionary Biogeography 235
a b
c d
Fig. 8.8 Global distribution of the sister families to the Asteraceae: a Goodeniaceae;
b Calyceraceae; c illustration of Boopis alpina, from Poeppig and Endlicher (1835–1845) (Chap 2);
d Nastanthus spathulatus (photo by A. Moreira-Muñoz)
and de Villiers 2000; Scott et al. 2006). Anderberg et al. (2007) emphasize sev-
eral uncertainties of identification and/or dating of all pre-Oligocene records. Early
to Middle Oligocene records are found in western USA, Chile, central Europe
(Anderberg et al. 2007), and in the Late Oligocene of North America and Patagonia
(Katinas et al. 2007). Recently Palazzesi et al. (2009) reported the presence of fos-
sil pollen grains in the Miocene of Patagonia, pertaining to the basal Barnadesieae;
Stuessy et al. (1996) have suggested an origin in the early Oligocene (38 mya) for
Barnadesieae (Box 8.2).
Box 8.2 The Barnadesieae/Barnadesioideae

The tribe (currently considered as a subfamily), comprises 9 genera and 91
species, and is located at the base of the cladogram as a sister group to
the rest of the tribes (Fig. 8.4). The genera of the Barnadesieae are mainly
characterized by the absence of a chloroplast DNA inversion characteris-
tic of the rest of the family (Jansen and Palmer 1987; Bremer 1994). The
tribe has an exclusively South American distribution, with genera distributed

from Venezuela to Patagonia, including Eastern Brazil, but surrounding
the Amazon basin (Fig. 8.9). In Chile three genera occur: Chuquiraga,
Dasyphyllum and Doniophyton (Fig. 8.9). The disjunct distribution of the
arborescent Dasyphyllum species (= subgenus Archidasyphyllum) of south-
ern Chile/Argentina is (Fig. 8.2), in comparison with the distribution of the
rest of the genus, very challenging. Cabrera (1959) proposed the subgenus
Archidasyphyllum as the most primitive within the tribe, but newer phylo-
genetic analyses by Gruenstaeudl et al. (2009) support a basal evolutionary
split within the subfamily into a clade comprising Chuquiraga, Doniophyton,
and Duseniella and a clade comprising all other genera. Archidasyphyllum
species (i.e. Chilean Dasyphyllum) appear most closely related to Fulcaldea
and Arnaldoa from the western northern Andes. This relationship, together
with the east-west disjunction across the Andes between both subgenera, sug-
gests the future elevation of these taxa to the generic rank (Gruenstaeudl et al.
2009).
Fulcaldea
Arnaldoa
Chuquiraga
Dasyphyllum
Barnadesia
Schlechtendahlia
Doniophyton
Huarpea
Duseniella
Fig. 8.9 Distribution of Barnadesieae (from Moreira-Muñoz (2006), by permission of

Jardín Botánico Chagual)
8.4 Asteraceae Evolutionary Biogeography 237
Most tribes appear in the fossil record only in the middle Miocene (14
mya) (Graham 1996). The fossil record encompasses both primitive clades (e.g.
Mutisieae) as well as derived ones (e.g. Heliantheae). This supports the idea of an
older age than the Eocene for the family (Graham 1996). The fossil record gets
richer into the Miocene (5 mya), where representatives of most of the present tribes
have been found (Graham 1996).
The geographical spread of the Asteraceae during the Miocene (10–20 mya) was
probably correlated with the development of extensive dry to semi-arid ecosystems
with seasonal precipitation (Anderberg et al. 2007). The family members show
several adaptations to exploit conditions of limited water availability under cli-
mates with seasonal rainfall (Hendry 1996). This, coupled with the development
of effective dispersal mechanisms and effective chemical defenses against herbi-
vores, enabled the Asteraceae to spread in the grassland vegetation types which
became widespread during the Late Miocene, and pre-adapted them to the cooler,
xeric conditions arising during the Pliocene (Anderberg et al. 2007).
Luebert et al. (2009) emphasize the role of the Andean uplift and the Atacama
aridity (see also Boxes 1.5 and 7.1) in the evolution of the endemic Asteraceae gen-
era Oxyphyllum and Gypothamnium. Interestingly, the sympatric distribution range
of these two genera arose under different scenarios: Oxyphyllum seems to be one
of the northernmost representatives of a group with its highest presence in central
Chile, like the closely related Leucheria, Moscharia, and Polyachyrus. Contrary to
this pattern, the ancestral group related to Gypothamnium, encompassing genera like
Onoseris and Lycoseris, is distributed in the tropical areas of north-western South
America and Mesoamerica (Luebert et al. 2009). Therefore, the current superposed
distribution is to be traced along two different evolutionary lines, both related to
the uplift of the Andes and the continuous drought of the Atacama. According to
Luebert et al. (2009) the Andes played its role in the origin and diversification of the
Atacama Desert flora in three different ways: (1) acting as a corridor that allowed
the north–south geographical range expansion of taxa, causing the pattern of some
related groups to establish a distribution range along the Andes; (2) the Andes have
acted as a vicariant barrier separating closely related groups on the eastern and
western sides; (3) they served as a new niche for speciation in high-elevation envi-
ronments, which may explain the fact that there are phylogenetically related taxa
in the basal areas of both sides of the Andes and also at high elevations (see also
Hershkovitz et al. 2006, for the evolution of Chaetanthera on Andean elevations).
Muellner et al. (2005) further reported a complex system of recolonization routes
after Pleistocene glaciations in the temperate Andes by Hypochaeris palustris.
In spite of the fact that it seems so recent, several authors propose the
Pliocene/Pleistocene, in relation to the final uplift of the Andes, as the time
of origin of several endemic genera of the Mutisieae in Central Chile, such as
Moscharia and Polyachyrus that apparently descended from a Pliocene/Pleistocene
ancestor (Katinas and Crisci 2000). Simpson (1973) also interpreted the split-
ting of the Prenanthoides group in Perezia, as a recent Pleistocene event.
Also Abrotanella (Senecioneae) (Swenson and Bremer 1997) and Chuquiraga
(Barnadesieae) (Ezcurra 2002) have a supposedly Pleistocene origin.
8.4.1 Origin and Dispersal Routes

The many trans-Pacific relationships led Bremer (1993) to suggest the Pacific basin
as related to the early evolution of the family. This Pacific relationship is to be found
among the Chilean Asteraceae in the circum-Pacific, australasiatic, and Pacific
antitropical tracks, encompassing 21 genera belonging to different tribes (Table 8.2).
The early evolution of the family seems to be related to the fragmentation of West
Gondwana during the Palaeocene, but there is still the question of how the rep-
resentatives of the family reached the rest of the world. Same as for the whole
flora (Sect. 3.3), currently many researchers are appealing to long-distance dispersal
scenarios to explain the global spread of the family.
The genera of the Cichorieae shared between South America and North America,
such as Microseris, supposedly have undergone a greater radiation in North America
with later dispersion towards South America and Australia/New Zealand (Chambers
1963; Vijverberg et al. 1999; Lee et al. 2003). Recently discovered Peruvian
Microseris populations are supposedly derived from Chilean progenitors instead of
being remnants of stepping-stone dispersal from North America through the Andes
to Chile (Lohwasser et al. 2004). Analogous molecular analyses of Hypochaeris
suggest that the closest relatives to the South American species are in Northwest
Africa. This has been interpreted as a direct dispersion Africa-South America,
without a previous stop in Europe-North America (Tremetsberger et al. 2005).
The remarkable arborescent Cardueae, Cichorieae and Senecioneae from Juan
Fernández have been also interpreted as the result of long-distance dispersal from
the mainland, but the closest relatives are unknown, and the oceanic nature of the
islands has been long part of the debate (Chap. 5). This is valid also for the endemic
genera from the Desventuradas Islands, Lycapsus and Thamnoseris (Chap. 5). The
oldest taxa of the Senecioneae were apparently herbs and the arboreal habit and
the palmate venation of the leaves arose later; the arboreal genus Robinsonia is an
example. The genus suffered an explosive radiation in the Juan Fernández Islands,
reaching eight species endemic to this archipelago (Sanders et al. 1987), but the
probable origin and extension of this genus is unknown (Chap. 5). Small (1919)
proposed the Senecioneae as the basal group for the phylogeny of the family, in
agreement with the “Age and area” theory of Willis (1922). The theory supposes
that the more diversified groups had more time to evolve and therefore they are
the oldest. Present evidence suggests that at least for the Asteraceae this is not the
case, and that Senecioneae is nested well within the family (Fig. 8.4). Also from
the Senecioneae, current Pacific disjunction in Abrotanella has been also proposed
as the result of long-distance dispersal (Swenson and Bremer 1997; Wagstaff et al.
2006), in spite of lacking a pappus suited for this.
8.4.2 Dispersal Capacities
There is a common believe that Asteraceae taxa have a good dispersal capacity,
due to their wind-dispersed achenes; therefore the taxa should have reached far
8.5 Conservation v/s Invasions 239
away territories by means of long-distance dispersal. But specific experiments have

shown that the dispersal capacity is sometimes very restricted: even wind-dispersed
achenes can travel only several meters under specific environmental conditions
(Sheldon and Burrows 1973; McEvoy and Cox 1987).
Taking account of the fruit characteristics, taxa having a pappus not suited for
wind dispersal, such as Abrotanella and its Pacific disjunct distribution, have been
interpreted as the result of vicariant form-making prior to the Palaeogene, in relation
to processes of subduction, accretion and orogenesis associated with the tectonic
evolution of the Pacific region (Heads 1999). To the same deduction came Grehan
(2007) while analyzing the disjunct distribution of Microseris. Another eight gen-
era with a Pacific distribution do not have a pappus suited for wind transport:
Blennosperma, Centipeda, Flaveria, Lagenophora, Lasthenia, Leptinella, Soliva,
and Psilocarphus.
The high level of endemism at the genus and species level also challenges the
supposedly good dispersal capacity, since theoretically floristic exchange would
obscure endemism and disjunct patterns. Also the low presence of cosmopolitan
genera (only four) in the Chilean Asteraceae flora contrasts with the assumption
that the Asteraceae have facilities of dispersion by wind or animals. Emphasizing
the notable patterns of endemism existent in all taxonomic levels, and challenging
any proposed centre of origin, Heads (2009) maintains that the Asteraceae might
descend from a globally widespread ancestor. This ancestor or few ancestors “might
have undergone a phase of active mobilism in the Mesozoic in which they occu-
pied much of Earth’s land surface, before settling down into a Cenozoic phase of
immobilism and speciation” (Heads 2009, p 765).
There seems to be still a lot to do in Asteraceae biogeography: “Perhaps the
size of the family, its global distribution, the lack of macrofossils and paucity of
discriminating characters in fossil pollen, and the lack of an agreed upon phylogeny
have restricted attempts to understand its history” (Funk et al. 2009, p 755).
8.5 Conservation v/s Invasions

Almost a third of the Chilean Asteraceae genera (35) show very restricted to
medium-small distribution ranges between just one locality to 5 latitudinal degrees
(Moreira-Muñoz and Muñoz-Schick 2007). This is not irrelevant, since one of the
most important criteria for classifying endangered species is the geographic range
extent (IUCN 2001). Furthermore, half of the genera (53) have their distribution in
central Mediterranean Chile from 27◦ to 37◦ S, in an intensively modified human
landscape (see Sect. 6.1). As shown by the map of generic richness (Fig. 8.6), cen-
tres of richness located between 31◦ –35◦ S and 36◦ –38S◦ , coincide with the most
populated regions (Santiago, Valparaíso, Concepción). This means a huge challenge
for the conservation of these taxa, since the intensive land occupation creates huge
gaps in protected areas (see Figs. 6.2 and 6.3).
An early recognized target of conservation is the species Dasyphyllum excelsum
(Box 8.2), which is considered as Endangered in the Species Red List (Sect. 6.1).
Dasyphyllum excelsum, a characteristic tree of the sclerophyllous forest of Central

Chile, pertains to the basal Barnadesieae and is therefore a critical species for the
understanding of the early evolution of the family. Regional Red Books (see Chap
6) also include many Asteraceae species. Almost 80 Asteraceae species have been
catalogued as vulnerable or endangered in the Atacama, Coquimbo and O’Higgins
regions (see Sect. 6.1). Considering the serious lack of adequate distributional data
to allow a good assessment, these types of assessments must be applied with caution:
sometimes a species that is considered highly vulnerable or even extinct can be
rediscovered with adequate field work, as in the case of Plazia cheiranthifolia, that
was considered as regionally extinct and was newly rediscovered in the Coquimbo
region (Faúndez and Saldivia 2008).
Nevertheless, this seems to be sadly not the case for the Fernandezian endemic
composites since the islands’ flora is the most endangered of the country and one of
the most threatened island territories worldwide (see Chap 5). This directly affects
several Asteraceae species: two species, Robinsonia berteroi and R. macrocephala
are already extinct (Danton and Perrier 2005), while ca. 30 species (i.e. all of the
islands’s Asteraceae!) are classified ranging from critically endangered (CR) to
vulnerable (VU) (Danton and Perrier 2006).
On the other hand, the Chilean Asteraceae flora, on the islands as well as on the
continent, is constantly “enriched” by alien species that are becoming rapidly nat-
uralized. Castro and Muñoz-Schick (2006) reported the presence of 58 Compositae
genera and 99 species naturalized in continental Chile.
Only 15 (26%) of these genera were present in the flora as native genera. Most
of the genera (43; 74%) are new for the Chilean flora. The newly incorporated gen-
era tend to be less diverse than the others, since these are represented by a mean
of 1.4 species, while genera already present in Chile contain a mean of 2.4 intro-
duced species. This fact suggests that the phylogenetic relationships could affect
the effectiveness of introduced species: genera already represented in Chile by
means of native species have major possibilities of being enriched by introduced
species. The mechanisms underlying these observations are unknown, but Castro
and Muñoz-Schick (2006) speculate a sort of pre-adaptation of several species to
occupy a specific niche in a new territory. Of course this is a very interesting field
of research due to the richness of “invaders” in the family. The competition that one
would expect in closely related species is a totally undeveloped field of research. The
Asteraceae are certainly one of the best study objects in the research of invasions
(sensu Pauchard et al. 2009).
Most of the introduced Asteraceae genera are represented by one species.
Several genera represented by two to four species are: Acmella, Anthemis, Carduus,
Carthamus, Chrysanthemum, Cirsium, Conyza, Hypochaeris, Lactuca, Tanacetum,
Ambrosia, Calendula, Chamomilla, Leontodon, Xanthium, Cotula, Crepis and
Sonchus (see examples in Fig. 8.10). The most diverse genera are represented by
five introduced species, from the genera Bidens, Centaurea, Hieracium and Senecio.
These genera are originally present in Chile, although the relation is not propor-
tional: Senecio has more than 220 native species (Teillier and Marticorena 2006)
Fig. 8.10 Chrysanthemum

coronarium and
Chrysanthemoides
monilifera, two highly
invasive Asteraceae species in
coastal central Chile (original
illustration by Sergio
Elórtegui Francioli)
while Bidens, Centaurea and Hieracium have between six and seven native species,
i.e. introduced species can soon overpass native species in these genera.
Regarding the geographic origin, most of the naturalized Compositae come from
Mediterranean Europe and northern Africa (72% of the species). Only 10% comes
from other South American countries. Interestingly, the naturalized Compositae
flora also shows a “natural” spatial segregation: the northern assemblage is different
from the one from Central Chile and from the southern one, the central assemblage
being the species-richest. This fact may reflect the different human colonization
history of the territory, or/and that the environmental differences indeed affect the
establishment of the introduced flora (Castro and Muñoz-Schick 2006).
8.5.1 Invading Biogeography

Invasive species, not just Asteraceae, are putting biogeography in trouble; Schram
(2008) asks: “. . .does biogeography have a future in a globalized world. . .?” Or
in the words of Nijman and Vonk (2008): “. . .in a globalised world, with species
reintroductions, invasions of alien species, and large-scale extinctions, unravel-
ling the true biogeographic relationships between areas and species may become
increasingly difficult” (Nijman and Vonk 2008). But the analysis of invasions shows
also good research opportunities: several authors consider invasions as “natural

experiments” for the study of spatially explicit phenomena like dispersal, coloniza-
tion, range expansion, and population dynamics. In this sense, the study of invasions
contributes to our understanding of the spatial dimensions of ecological processes,
and to biogeography! (Pauchard 2007; Crawford and Hoagland 2009; Pauchard
et al. 2009). One of the most promising research topics is the range expansion of
naturalized Asteraceae species. Several of them can be traced back to the moment
of their arrival and it seems attractive to evaluate possible scenarios of maximal
future expansion, via GIS niche modeling, and phylogeography (Ortiz et al. 2008).
Distributional modeling has been indeed used for the analysis of Chilean native
Senecio species (Rovito et al. 2004).
It is worth to emphasize the need of strengthening the herbaria collections, for
the native as well as for the naturalized species, thus minimizing the “Wallacean
shortfall” (Box 6.2). In spite of new research programs like niche modeling on GIS
platforms (Sect. 6.2), which can support conservation initiatives, good botanical col-
lections are essential to a better performance of the models. An exercise done with
the native Mutisia species is presented in Box. 8.3 (Fig. 8.11 ). A better knowledge
of the geographic distribution and ecological situation of the Chilean Asteraceae will
enable us to propose new lines of inquiry that will allow us a better understanding
of the family’s evolution in time and space.
Box 8.3 Niche Modeling of Mutisia Species

Niche modelling techniques are getting widely used, together with GIS tech-
niques and remote sensing, to identify and understand current distributions
as well as to predict range shifts in the future (Guisan and Thuiller 2005;
Kosak et al. 2008; Gillespie et al. 2008). In Chile the application of niche
modelling is a very promising field of research (e.g. Zizka et al. 2009, for
Chilean Bromeliaceae). Moreira-Muñoz et al. (2009) showed an example of
its application with the distribution modelling of Chilean Mutisia species.
Twenty one species were modelled, based on the collections from the SGO
Herbarium and the monograph (Cabrera 1965), encompassing 700 georefer-
enced localities in Chile and adjacent localities in Argentina, Perú and Bolivia.
Twenty percentage of the data was set apart for testing the fit of the models in
the program MaxEnt. Nineteen environmental variables were obtained from
Worldclim (www.worldclim.org), together with the digital elevation model
(DEM). Results yielded a bad fit for 7 species that were those with less than
13 collection localities. Five environmental variables better explained the dis-
tribution of the analyzed species: altitude, Min T◦ colder month, Mean T◦ of
the humid quarter, Mean T◦ warmer trimester, precipitation warmer trimester.
The program was run again and models of probability were obtained and
transferred to binary maps with the cut-value obtained for each species. These
results are still preliminary; the models seem to be very sensitive to the num-
ber of presences. Testing of the climatic surfaces seems to be also necessary, as
well as adequate georeferencing, which is critical for the work with Herbarium
collections (see the richness map of Chilean Mutisia species in Fig. 8.11).
Fig. 8.11 Distribution niche modelling of Chilean Mutisia species: a M. decurrens; b M.

latifolia; c M. oligodon; d M. rosea; e richness map based on niche modelling
References
Anderberg AA, Baldwin BG, Bayer RG, Breitwieser J, Jeffrey C, Dillon MO, Eldenäs P, Funk V
et al. (2007) Compositae. In: Kubitzki K, (ed) Asterales. Kadereit JW, Jeffrey C (vol eds), The
families and genera of vascular plants, vol 8. Springer, New York, NY, pp 61–621
Bentham G (1873) Notes on the classification, history and geographic distribution of Compositae.
J Linn Soc Bot 13:355–577
Bonifacino JM (2008) Reinstatement of Ocyroe (Compositae: Astereae). Brittonia 60:205–212
Bonifacino JM (2009) Taxonomic revision of the Chiliotrichum group sensu stricto (Compositae:
Astereae). Smithson Contrib Bot 92:1–115
Bremer K (1993) Intercontinental relationships of African and South American Asteraceae: a
cladistic biogeographic analysis. In: Goldblatt P, (ed) Biological relationships between Africa
and South America. Yale University Press, New Haven, CT and London, pp 105–135
Bremer K (1994) Asteraceae: cladistics and classification. Timber Press, Portland, OR
Bremer K, Gustafsson MHG (1997) East Gondwana ancestry of the sunflower alliance of families.
Proc Nat Acad Sci USA 94:9188–9190
Cabrera AL (1959) Revisión del género Dasyphyllum (Compositae). Rev Mus Plata (Ser. 2) 9:
21–100
Cabrera AL (1965) Revisión del género Mutisia (Compositae). Opera Lilloana 13:1–227
Cabrera AL (1975) Nota sobre cinco Compositae de la República Argentina. Bol Soc Arg Bot
16:255–260
Cassini H (1819) Suit du sixiéme mémoire sur la familie des Synanthérées, contenat les caractères
des tribus. J Phys Chem, Hist Nat Arts 88:189–204
Castro S, Muñoz-Schick M (2006) Compuestas naturalizadas en Chile: Importancia de la flora
exótica como agente del cambio biotic. Rev Chagual (Jard Bot Santiago) 4:29–38
Chambers KL (1963) Amphitropical species pairs in Microseris and Agoseris (Compositae:
Cichorieae). Q Rev Biol 38:124–140
Crawford PHC, Hoagland BW (2009) Can herbarium records be used to map alien species invasion
and native species expansion over the past 100 years? J Biogeogr 36:651–661
Danton P, Perrier C (2005) Notes sur la disparition d’une espèce emblématique: Robinsonia bert-
eroi (DC.) Sanders, Stuessy and Martic. (Asteraceae), dans l’île Robinson Crusoe, archipel
Juan Fernández (Chili). J Bot Soc Bot France 31:3–8
Danton P, Perrier C (2006) Nouveau catalogue de la flore vasculaire de l’archipel Juan Fernández
(Chili). Acta Bot Gall 153:399–587
DeVore ML, Stuessy TF (1995) The place and time of origin of the Asteraceae, with additional
comments on the Calyceraceae and Goodeniaceae. In: Hind DJN, Jeffrey C, Pope GV, (eds)
Advances in Compositae systematics, vol. 1. Royal Botanic Gardens, Kew, pp 23–40
Ezcurra C (2002) Phylogeny, morphology, and biogeography of Chuquiraga, an Andean-
Patagonian genus of Asteraceae-Barnadesioideae. Bot Rev 68:153–170
Faúndez L, Saldivia P (2008) Plazia cheiranthifolia, especie ‘extinta’, redescubierta en la
precordillera de Ovalle, region de Coquimbo, Chile. Noticiario Mens Mus Nac Hist Nat
360:18–21
Funk VA, Bayer RJ, Keeley S, Chan R, Watson L, Gemeinholzer B, Schilling E, Panero JL,
Baldwin BG, Garcia-Jacas N, Susanna A, Jansen RK (2005) Everywhere but Antarctica:
Using a supertree to understand the diversity and distribution of the Compositae. Biol Skr 55:
343–374
Funk VA, Susanna A, Stuessy TF, Bayer RJ, (eds) (2009) Systematics, evolution, and biogeography
of Compositae. International Association for Plant Taxonomy, Vienna
Gillespie TW, Foody GM, Rocchini D, Giorgi AP, Saatchi S (2008) Measuring and modelling
biodiversity from space. Prog Phys Geogr 32:203–221
Goloboff P (2005) NDM/VNDM ver. 2.6. Programs for identification of areas of endemism.
http://www.zmuc.dk/public/phylogeny/endemism
References 245
Graham A (1996) A contribution to the geologic history of the Compositae. In: Hind DJN, Beentje
HJ (eds) Compositae: Systematics. Proceedings of the international Compositae conference,
Kew, 1994, vol 1: 123–140
Grehan JR (2007). A brief look at Pacific biogeography: the trans-oceanic travels of Microseris
(Angiosperms: Asteraceae). In: Ebach MC, Tangney RS, (eds) Biogeography in a changing
world. Systematics association special volumes. CRC Press, Boca Raton, FL, pp 83–94
Gruenstaeudl M, Urtubey E, Jansen RK, Samuel R, Barfuss MHJ, Stuessy TF (2009) Phylogeny
of Barnadesioideae (Asteraceae) inferred from DNA sequence data and morphology. Mol Phyl
Evol 51:572–587
Guisan A, Thuiller W (2005) Predicting species distributions: offering more than simple habitat
models. Ecol Lett 8:993–1009
the genus Abrotanella (Compositae). Biol J Linn Soc 67:391–432
Heads M (2009) Inferring biogeographic history from molecular phylogenies. Biol J Linn Soc
98:757–774
Hendry GAF (1996) Fructan and the ecology and evolution of the Compositae. In: Caligari
PDS, Hind DJN (eds) Compositae: biology and utilization. Proceedings of the international
Compositae conference, Kew Royal Botanic Gardens, 1994, vol 2, pp 121–128
Hershkovitz MA, Arroyo MTK, Bell C, Hinojosa LF (2006) Phylogeny of Chaetanthera
(Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages.
Mol Phyl Evol 41:594–605
IUCN (2001) The IUCN List of Threatened Species. Categories and Criteria (version 3.1).
http://www.iucn.org/themes/ssc/redlists/RLcats2001booklet.html
Jansen RK, Palmer JD (1987) A chloroplast DNA inversion marks an ancient evolutionary split in
the sunflower family, Asteraceae. Proc Nat Acad Sci USA 84:5818–5822
Katinas L, Crisci JV (2000) Cladistic and biogeographic analyses of the genera Moscharia and
Polyachyrus (Asteraceae, Mutisieae). Syst Bot 25:33–46
Katinas L, Crisci JV, Tellería MC, Barreda V, Palazzesi L (2007) Early history of Asteraceae in
Patagonia: evidence from fossil pollen grains. NZ J Bot 45:605–610
Kosak KH, Graham CH, Wiens JJ (2008) Integrating GIS-based environmental data into evolu-
tionary biology. Trends Ecol Evol 23:141–148
Lee J, Baldwin BG, Gottlieb LD (2003) Phylogenetic relationships among the primarily North
American genera of Cichorieae (Compositae) based on analysis of 18S–26S nuclear rDNA ITS
and ETS sequences. Syst Bot 28:616–626
Lohwasser U, Granda A, Blattner FR (2004) Phylogenetic analysis of Microseris (Asteraceae)
including a newly discovered Andean population from Peru. Syst Bot 29:774–780
Luebert F, Wen J, Dillon MO (2009) Systematic placement and biogeographical relationships of
the monotypic genera Gypothamnium and Oxyphyllum (Asteraceae: Mutisioideae) from the
Atacama Desert. Bot J Linn Soc 159:32–51
McEvoy PB, Cox CS (1987) Wind dispersal distances in dimorphic achenes of ragwort, Senecio
jacobaea. Ecology 68:2006–2015
Moreira-Muñoz A (2006) Posición filogenética y distribución de los generous de Compuestas
chilenas, con algunas notas biogeográficas. Rev Chagual (Jard Bot Santiago) 4:12–28
Moreira-Muñoz A, Muñoz-Schick M (2007) Classification, diversity, and distribution of
Chilean Asteraceae: implications for biogeography and conservation. Divers Distrib 13:
818–828
Moreira-Muñoz A, Morales V, Pliscoff P (2009) Aplicación del programa ‘Maxent’ para mode-
lar la distribución geográfica de las especies del género Mutisia (Asteraceae). XXX Congreso
Nacional de Geografía Talca, Chile, 14 de octubre de 2009
Muellner AN, Tremetsberger K, Stuessy TF, Baeza CM (2005) Pleistocene refugia and recoloniza-
tion routes in the southern Andes: insights from Hypochaeris palustris (Asteraceae, Lactuceae).
Mol Ecol 14:203–212
Muñoz Pizarro C (1966) Sinopsis de la Flora Chilena, 2nd edn. Ediciones Universidad de Chile,
Santiago
Nijman V, Vonk R (2008) Blurring the picture: introductions, invasions, extinctions – biogeography
in a global world. Contrib Zool 77:67–70
Ortiz MA, Tremetsberger K, Terrab A, Stuessy TF, Garcia-Castano JL, Urtubey E, Baeza CM,
Ruas CF, Gibbs PE, Talavera S (2008) Phylogeography of the invasive weed Hypochaeris rad-
icata (Asteraceae): from Moroccan origin to worldwide introduced populations. Mol Ecol 17:
3654–3667
Palazzesi L, Barreda V, Tellería MC (2009) Fossil pollen grains of Asteraceae from the Miocene
of Patagonia: Barnadesioideae affinity. Rev Palaeobot Palynol 155:83–88
Palazzesi L, Barreda V, Tellería MC (2010) First fossil record of Calyceraceae (Asterales): Pollen
evidence from southern South America. Rev Palaeobot Palynol 158:236–239
Pauchard A (2007) Invasions as spatially explicit processes: contributions to ecology (Letter to the
editor). Front Ecol Environ 5:123–124
Pauchard A, Kueffer C, Dietz H, Daehler CC, Alexander J, Edwards PJ, Arévalo JR, Cavieres L
et al. (2009) Ain’t no mountain high enough: plant invasions reaching high elevations. Front
Ecol Environ 7:479–486
Reiche C (1905) La distribución geográfica de las Compuestas de la flora de Chile. An Mus Nac
Chile Secc Bot 17:1–45
Rovito SM, Arroyo MTK, Pliscoff P (2004) Distributional modelling and parsimony analysis of
endemicity of Senecio in the Mediterranean-type climate area of Central Chile. J Biogeogr
31:1623–1636
Sanders RW, Stuessy TF, Marticorena C, Silva M (1987) Phytogeography and evolution of
Dendroseris and Robinsonia, tree-Compositae of the Juan Fernández Islands. Opera Bot
92:195–215
Schram FR (2008) Does biogeography have a future in a globalized world with globalized faunas?
Contrib Zool 77:127–133
Scott L, Cadman A, Mcmillan I (2006) Early history of Cainozoic Asteraceae along the Southern
African west coast. Rev Palaeobot Palynol 142:47–52
Sheldon JC, Burrows FM (1973) The dispersal effectiveness of the achene-pappus units of selected
Compositae in steady winds with convection. New Phytol 72:665–675
Simpson BB (1973) Contrasting modes of evolution in two groups of Perezia (Mutisieae;
Compositae) of southern South America. Taxon 22:525–536
Small J (1919) The origin and development of the Compositae. New Phytol 18:201–234
Stevens PF (2001 onwards) Angiosperm Phylogeny Website. http://www.mobot.org/MOBOT/
research/APweb/
Stuessy TF, Sang T, DeVore ML (1996) Phylogeny and biogeography of the subfamily
Barnadesioideae with implications for early evolution of the Compositae. In: Hind DJN,
Beentje HJ (eds) Compositae: Systematics. Proceedings of the international Compositae
conference, Royal Botanic Gardens, Kew, 1994, vol. 1, pp 463–490
Swenson U, Bremer K (1997) Pacific biogeography of the Asteraceae genus Abrotanella
(Senecioneae, Blennospermatinae). Syst Bot 22:493–508
Teillier S, Marticorena A (2006) El género Senecio (Asteraceae): el más diverso de Chile. Rev
Chagual (Jard Bot Santiago) 4:39–48
Tremetsberger K, Weiss-Schneeweiss H, Stuessy TF, Samuel R, Kadlec G, Ortiz MA,
Talavera S (2005) Nuclear ribosomal DNA and karyotypes indicate a NW African
origin of South American Hypochaeris (Asteraceae, Cichorieae). Mol Phylogenet Evol
35:102–116
Turner BL (1977) Fossil history and geography. In: Heywood VH, Harborne JB, Turner BL, (eds)
The biology and chemistry of the Compositae, vol. 1. London, Academic Press, pp 21–39
Vijverberg K, Mes THM, Bachmann K (1999) Chloroplast DNA evidence for the evolution
of Microseris (Asteraceae) in Australia and New Zealand after long-distance dispersal from
western North America. Am J Bot 86:1448–1463
References 247
Wagstaff SJ, Breitwieser I, Swenson U (2006) Origin and relationships of the austral genus
Abrotanella (Asteraceae) inferred from DNA sequences. Taxon 55:95–106
Willis JC (1922) Age and area: a study in geographical distribution and origin of species.
Cambridge University Press, Cambridge
Zavada M, de Villiers S (2000) Pollen of the Asteraceae from the Paleocene-Eocene of South
Africa. Grana 39:39–45
Zizka G, Schmidt M, Schulte K, Novoa P, Pinto R, König K (2009) Chilean Bromeliaceae:
diversity, distribution and evaluation of conservation status. Biodivers Conserv 18:2449–2471
Sur: Argentina, Sur de Brasil, Chile, Paraguay y Uruguay. Monographs in systematic botany,
vol 107. Missouri Botanical Garden, St. Paul, MO, 1715–1830
Chapter 9
Nothofagus, Key Genus in Plant Geography
Abstract The genus Nothofagus is the only representative of the Nothofagaceae. It

comprises 36 species distributed in disjunct territories of the southern hemisphere:
southern South America, New Zealand, East Australia, New Caledonia and New
Guinea. This southern distribution pattern, together with the rich fossil record that
is also found in Antarctica, sustains the status of the genus as a “key genus” in plant
geography. The evolution of the genus is discussed, confronting different visions,
and integrating the phylogeny of related genera, such as the fungi species of the
genus Cyttaria.
Nothofagus has been considered as a “key genus” in plant geography (Darlington

1965; Van Steenis 1971) for at least three reasons: (1) because of the characteristic
disjunct pattern of its species between South America and Australasia, together with
the dominant character of these species in their territories of occurrence (Donoso
1993; Veblen et al. 1996) (Fig. 9.1); (2) because of its abundant fossil record,
expressed in a rich array of macro and microfossils found in the southern territories
including Antarctica (Dettmann et al. 1990); and (3) because the use of the genus
as a model for testing different biogeographic methods and the various contesting
visions that different studies have reached (Humphries 1981; Heads 1985, Linder
and Crisp 1995, Cook and Crisp 2005; Knapp et al. 2005).
9.1 Taxonomy and Phylogeny

The genus Nothofagus was described by Blume in 1850, as a member of the family
Fagaceae, associated with Fagus in the subfamily Fagoideae (hence the name: “false
Fagus”). The family Nothofagaceae was proposed in 1962 by Kuprianova mainly on
the basis of pollen characteristics. Since then, morphological and molecular studies
highly support recognition of the family (Nixon 1982; Hill and Jordan 1993).
Li et al. (2004) have placed the Nothofagaceae as a sister clade of the rest of the
families of the Fagales. However, the phylogenetic relationship of Nothofagus with
members of the family Fagaceae remains unclear. Several authors have proposed
a close relationship between Nothofagus and Fagus (Van Steenis 1971; Hill and

DOI 10.1007/978-90-481-8748-5_9, ⃝
250 9 Nothofagus, Key Genus in Plant Geography
b c
Fig. 9.1 Nothofagus species: a N. macrocarpa, Cerro El Roble; b N. glauca, flowers, Altos de
Lircay (photos A. Moreira-Muñoz); c N. antarctica (plate 123 in Hooker JD (1847) The Botany of
the Antarctic Voyage . . . London (Chapter 2))
9.1 Taxonomy and Phylogeny 251
d e
f g
Fig. 9.1 (continued) d N. alpina; e N. alessandrii, Los Ruiles; f N. dombeyi and N. glauca,
Altos de Lircay; g Cyttaria sp., Altos de Lircay (photo/illustration credits: e, f A. Moreira-Muñoz;
g S. Elórtegui Francioli; d plate 197 in Poeppig and Endlicher (1835–1845) Nova genera et species
plantarum . . . Leipzig (Chapter 2))
Jordan 1993), on the base of morphological similarities like the cupule. However,
Nixon (1989) argues that Nothofagus is not closely related to Fagus, but it shares
some important apomorphic features with the family Betulaceae. In accordance with
Nixon’s point of view, temperate Nothofagus species share some leaf characters
with representatives of the Betulaceae. These characters, such as a leaf margin with
compound teeth, and analogous venation types, are distinct from those of Fagus
(Li and Zhou 2007). These authors suggest that extant Nothofagus species could
be classified into two groups: tropical species usually having an entire leaf margin,
and temperate species having teethed or crenate margins. Since the venation types
of these two groups are also different, Li and Zhou (2007) further suggest that the
genus may be split in several genera.
Van Steenis (1971) indeed divided the genus Nothofagus into two sections:
Nothofagus and Calusparassus: the first section encompassing the deciduous
species from South America and the deciduous N. gunnii from Tasmania. Van
Steenis (1971) interpreted the characteristics of the cupule as very relevant for the
phylogeny, suggesting a “progression rule” in the ornamentation of the cupule and
a reduction in seed number (Fig. 9.2). In the section Calusparassus he included
the evergreen species of Australasia and the three South American evergreen
species (N. betuloides, N. dombeyi, and N. nitida). Most recent taxonomic studies
tend to dismiss the deciduous/evergreen character as important in the classification,
favoring instead the morphological characteristics of the pollen. The deciduous/
evergreen character would have more ecological than phylogenetic significance
(Ramírez et al. 1997). The possibility of hybridization between deciduous species
(Donoso et al. 1990) and between deciduous and evergreen species (Stecconi et al.
2004) supports this view. According to the differences in pollen, several authors rec-
ognize four monophyletic groups classified as subgenera: Brassospora, Nothofagus,
Progression
towards
simplification of
the cupule
Progression Progression 21- 22 reduction of the
towards towards
ornamentation cupule segments
ornamentation
of the cupule of the cupule
23-25
pinnatipartite
lamellae
with glands
5
26, 27
divided cupula recurved
lamellae lamellae reduction to 1nut
with glands with glands
1, 2 6 10, 14, 15 28 - 30, 35
reduction
divided to 2-segmented
lamellae cupule
4, 16 7, 8, 9 13 17, 20
31, 34
3 11, 12
3 ridge nut 2 ridge nut
Section Nothofagus Section Calusparassus
Fig. 9.2 Illustration of the cupules of Nothofagus species (adapted from Van Steenis 1971, also
reproduced in Gottlieb 2001). Numbers refer to the species in Table 9.1
9.1 Taxonomy and Phylogeny 253
Table 9.1 Species composition and geographical distribution of the Nothofagus subgenera.
Source: several authors in Veblen et al. (1996) and Rodríguez and Quezada (2003) for South
American species. (Numbers after the species refer to numbers in Fig. 9.2). Nothofagus leonii
is considered as a hybrid between N. glauca and N. macrocarpa
Subgenus Pollen type Distribution N◦ sp. Species
Nothofagus fusca type b South America (SA) 5 N. antarctica (4), N. betuloides

(7), N. dombeyi (9), N. pumilio
(6), N. nítida (8)
Fuscospora fusca type a South America (SA) 1 N. alessandrii (3)
Tasmania (TAS) 1 N. gunnii (16)
New Zealand (NZ) 3 N. truncata (12), N. fusca (11),
N. solandri (13)
Lophozonia menziesii South America (SA) 4 N. alpina (5), N. glauca (2),
N. macrocarpa, N. obliqua (1)
New Zealand (NZ) 1 N. menziesii (10)
Australia (AUS) – 1 N. cunninghamii (15)
Tasmania (TAS)
Australia (AUS) 1 N. moorei (14)
Brassospora brassi New Caledonia (NC)a 5 N. balansae (33), N. baumanniae
(34), N. codonandra (32),
N. discoidea (35),
N. aequilateralis (31)
New Guinea (NG) 14 N. brassii (20), N. carrii (23),
N. crenata (28), N. flaviramea
(24), N. grandis (29), N. nuda
(18), N. perryi (17),
N. pseudoresinosa (27),
N. pullei (22), N. resinosa
(21), N. rubra (30),
N. starkenborghii (19),
N. stylosa (25), N.
womersleyi (26)
a New Caledonian species still need a careful taxonomic revision, according to Munzinger (2010).
Fuscospora and Lophozonia (Hill and Read 1991; Hill and Jordan 1993; Manos
1997) (Fig. 9.3).
Hill and Jordan (1993) proposed the subgenus Fuscospora as basal in the phy-
logeny, including the species N. alessandrii,1 also proposed by van Steenis (1971)
as the most ancient species of the genus because of their domes of 7 flowers
(Fig. 9.2). However, Manos (1997), based on morphological and molecular anal-
ysis, placed the subgenus Lophozonia at the basis of the phylogeny, as shown
in Fig. 9.3. Nevertheless, Manos (1997) recognized that Fuscospora shows many
ancestral characters, so it does not appear as monophyletic in all the trees. The
subgenera Nothofagus and Brassospora appear as the most derived groups.
1 Apart from its phylogenetic relevance, this is one of the most threatened plant species in Chile
(Torres-Díaz et al. 2007, see also Sect. 6.2).
Fig. 9.3 Phylogenetic N. grandis

hypothesis for Nothofagus
Brassospora
N. perryi
(adapted from Manos (1997),
updated for subgenus N. brassii
Nothofagus from Acosta and N. resinosa
Premoli (2010))
N. balansae
N. aequilateralis
N. antarctica
Nothofagus
N. pumilio
N. nitida
N. betuloides
N. dombeyi
N. truncata
Fuscospora
N. fusca
N. solandri
N. gunnii
N. alessandrii
N. alpina
N. glauca
Lophozonia
N. obliqua
N. cunninghamii
N. moorei
N. menziesii
The interespecific relationships within of the subgenus Nothofagus has been

recently studied by Acosta and Premoli (2010). They analyzed morphology and
sequences of chloroplast and nuclear DNA and found good support for the
monophyly of the subgenus, and according to the results shown by nuclear DNA,
the evergreen species N. dombeyi and N. betuloides differentiated more recently.
These species together with N. nitida are sisters to the deciduous N. antarctica.
Nothofagus pumilio (deciduous) seems to have a different and earlier evolutionary
history (Fig. 9.3). Chloroplast DNA did not show a clear specific differentiation, but
varied instead with geographic location.
9.2 Diversity and Distribution

The extant 36 species of the genus Nothofagus are distributed in the territories of the
South Pacific, from 33◦ to 55◦ S in South America (Chile and Argentina), and from
0◦ to 47◦ S in Australasia: New Guinea (including the Islands d’Entrecasteaux and
9.2 Diversity and Distribution 255
Nothofagus - Brassospora
NG
NC
SA
Fuscospora
NZ
TAS
SA
SA
TAS/ Lophozonia
AUS
NZ
Fig. 9.4 Area cladogram (areagram) of Nothofagus subgenera; tracks represent the disjunct dis-
tribution of the subgenera: abbreviations of the territories according to Table 9.1 (adapted from
Moreira-Muñoz 2004)
New Britain), New Caledonia, Australia, Tasmania and New Zealand (van Steenis
1971) (Fig. 9.4 and Table 9.1).
The geographic relationships have been represented in Fig. 9.4 as an area clado-
gram, following the phylogenetic hypothesis of Manos (1997). Figure 9.4 also show
the tracks that join the species that compose the subgenera, showing three dis-
junctions: between species in Lophozonia, between species in Fuscospora, and the
disjunct pattern between the subgenera Nothofagus and Brassospora.
The distribution of the genus in Chile encompasses half of the country’s latitu-
dinal extent from 33◦ S to the South (Fig. 9.5). Two cores of species richness can
be identify: a little one at 36◦ –37◦ S and a second spreader one between 39◦ and
42◦ S. Several species occupy both cores, i.e. N. alpina, N.antarctica, N. dombeyi,
N. obliqua, and N. pumilio, while N. glauca is restricted to the northern centre. The
southern centre lacks N. glauca but incorporates N. betuloides and N. nitida from
40◦ to the South. N. alpina reaches only 41◦ S (Fig. 9.5).
33°
43°
53°
N. alessandrii N. alpina N. antarctica N. betuloides
33°
20°
1
2–3
43° 4–5
6
30°
53°
N. dombeyi N. glauca N. macrocarpa
33° 40°
43°
50°
53°
N. nitida N. obliqua N. pumilio Richness
Fig. 9.5 Distribution and species richness of Nothofagus in Chile (collections SGO)
9.3 Speciation v/s Extinction

Hill (2001a), among others, suggests that Nothofagus was much more diverse in the
past and the number of species remaining today is relatively small compared to its
peak diversity, possibly during the Late Oligocene-Early Miocene. He also argues
9.3 Speciation v/s Extinction 257
that this was also the time of maximum spatial distribution of the genus (although in
Chile the northernmost expansion was during the Maastrichtian; see Chap. 1). Hill
(2001a) emphasizes that it is difficult to study the evolutionary history of Nothofagus
based exclusively on evidence from extant species.
Nothofagus fossils are found in almost all regions of occurrence of the extant
species. Fossils have also been discovered in Antarctica, particularly in the Antarctic
Peninsula and its nearby islands (Torres 1984, 2003) (Box 9.1). Among macrofos-
sils, leaves are most common, followed by fossil wood and cupules; these latter
are extremely rare, reported only from Tasmania (Hill 2001b). Fossil wood resem-
bling extant Nothofagaceae has been traditionally classified under the organ genus
Nothofagoxylon (Poole 2002).
While the description and interpretation of both macro-and microfossils of
Nothofagus is not without difficulties (Hill 2001a), the fossil record of the genus is
one of the most informative in the palaeobotany of the southern hemisphere. Even its
absence is informative: the lack of fossils of the genus in India and Africa suggests
an origin and radiation after the separation of these territories at the beginning of the
fragmentation of West Gondwana. Both stratigraphically and geographically, pollen
fossils are the most common. Fossil pollen is especially useful because it is pro-
duced in large quantities, is morphologically distinctive and very well preserved in
sediments (Dettmann et al. 1990; Hill and Dettmann 1996; Hill 2001a). The oldest
known occurrence involves the presence of the four types of pollen (brassii, fusca a,
fusca b, menziesii) in deposits from the upper Cretaceous of Antarctica (Campanian
= ca 83 mya) and South America (Maastrichtian = ca 70 mya) (Table 9.2). The
four types have been also found in more modern deposits from Australia, Tasmania
and New Zealand. This suggests an ancestral continuous distribution of the four
subgenera, from South America to Australasia via Antarctica, until the removal of
Nothofagus forests from Antarctica during the Pliocene (Hill and Dettmann 1996)
(Box 9.1).
Table 9.2 Microfossil Nothofagus (pollen) for different epochs and territories during the Late
Cretaceous and the Cenozoic. Simplified from Swenson et al. (2001)
Lophozonia Fuscospora Nothofagus Brassospora
Miocene (23–5 – – – New Guinea

mya)
Eocene (53–34 New Zealand, – – New Zealand
mya) Tasmania
Palaeocene Australia Australia, Australia, Australia,
(65–53 mya) Tasmania, Tasmania, Tasmania
New Zealand New Zealand
Campanian/ Antarctica, Antarctica, Antarctica, Antarctica,
Maastrichtian South South South South
(83–65 mya) America America America America
Box 9.1 The Green Dress of Antarctica

Antarctica was covered by forests from the Permian onwards (Chap. 1,
Fig. 1.10) (Torres 2003; Francis et al. 2008). The evolution of the Antarctic
climate from a greenhouse into the Neogene icehouse is captured within a
rich record of micro- (pollen) and macrofossils (leaves, wood, and flowers)
from the Antarctic Peninsula and the Transantarctic Mountains (Francis
et al. 2008). In the Early Cretaceous, the Antarctic forest ecosystem was
dominated by a conifer-fern community similar to that in the warm temperate
araucarian–podocarp rainforests of present-day New Zealand’s North Island
(Falcon-Lang et al. 2001). In the Late Cretaceous flowering plants radiated
throughout Gondwana changing the Antarctic physiognomy to one more
similar to the angiosperm-dominated cool temperate Valdivian rainforests
of present-day Chile. Characteristic taxa were Nothofagaceae, Myrtaceae,
Eucryphiaceae (=Cunoniaceae), Lauraceae, Monimiaceae, Araucariaceae,
Cupressaceae, Podocarpaceae and Winteraceae (Eklund 2003; Poole and
Gottwald 2001; Poole and Cantrill 2006). The similarities to the Valdivian
rainforests strengthened into the Cenozoic and become dominant up until
the Eocene (Poole and Cantrill 2006). By the Eocene (48 mya), Antarctic
rainforests were also inhabited by a rich vertebrate assemblage (Reguero
Fig. 9.6 Illustration of the biotic assemblage from the Oligocene of the Antarctic
Peninsula. Steppes were inhabited by ratites, palaeolamas, penguins’ ancestors. Forests of
Araucaria and Nothofagus were inhabited by little marsupials, among others. The con-
tinent’s inland showed already the beginning of the gradual massive glaciation (original
drawing by Sergio Elórtegui Francioli)
9.3 Speciation v/s Extinction 259
et al. 2002). Forests were increasingly reduced in area and diversity, due to
the global climate cooling and the initial formation of the Antarctic ice sheets
at the Eocene/Oligocene boundary (34–35 mya) (Fig. 1.10 and 9.6) (Zachos
et al. 2001; Francis and Poole 2002; Siegert et al. 2008). The development
of the Antarctic Circumpolar Current (ACC), related to the openings of the
Drake Passage and the Tasman Gateway (~45–30 mya), may have gener-
ated Antarctica’s own glaciation and processes of continental cooling (Denton
et al. 1991; Convey et al. 2008). There is no agreement on exactly when
between 41 and 24 mya the ACC was initiated (Lawver and Gahagan 1998,
2003; Scher and Martin 2006; Livermore et al. 2007), but it most certainly
allowed the ice sheets persistence on Antarctica during the Oligocene (till 27
mya). From this stage until the middle Miocene (~15 mya), the global ice vol-
ume remained low, due to a warming trend that reduced the extent of Antarctic
ice. This warm phase peaked in the Mid-Miocene climatic optimum (17–15
mya), and was followed by a gradual cooling and reestablishment of a major
ice-sheet on Antarctica (Zachos et al. 2001, see Sect. 1.2).
Antarctic vegetation became steadily more depauperate towards the end of the
Miocene (Fig. 1.18). The Miocene–Pliocene was probably more similar to the cur-
rent Magellanic tundra of southern Chile/Argentina (see Sect. 1.2), including local
stands of angiosperms like Nothofagus and a few conifer taxa (Francis et al. 2008).
It is likely that the vegetation went through a krummholz phase (e.g. Nothofagus
decreasing in stature) prior to becoming dominant by low prostrate shrubs in the
Pliocene tundra (Poole and Cantrill 2006). The timing of the Antarctic extinctions
caused by the climate cooling are still under debate due to the paucity of Neogene
fossil sites, possibly following the mid-Miocene warm interval at c. 17 mya or a
mid-Pliocene warm interval at about 3 mya (Ashworth and Cantrill 2004). Pollen
evidence from the Ross Sea basin indicates that tundra vegetation existed in East
Antarctica from the Oligocene into the Early Miocene (Ashworth et al. 2007). As
Antarctica’s climate cooled further into the Plio-Pleistocene deep-freeze, and the
continent became deprived of summer heat and liquid water, it lost almost all its vas-
cular plants (Francis et al. 2008), with the only exception of Deschampsia antartica
and Colobanthus quitensis.
As the four major lineages (subgenera) already existed in the Late Cretaceous
(80 mya), before the total breakup of Gondwana, extant taxa could have under-
gone a process of sympatric speciation in a continuous geographic range (Swenson
et al. 2001). The possibilities of hybridization between extant species support this
hypothesis. Subsequent events of fragmentation of the biota would merely exacer-
bate the process of speciation. However, today’s scenario might be mainly the result
of extinctions (Manos 1997; Swenson et al. 2001). Fossil leaves and cupules sim-
ilar to N. betuloides and N. dombeyi (subgenus Nothofagus) found in Oligocene
deposits on Tasmania show the presence and subsequent extinction of this group
in Australasia (Hill 2001b). Also the micro-fossil record shows the extinction of
Brassospora species in South America, Antarctica and New Zealand, and species of
the subgenus Nothofagus in Antarctica, Tasmania and New Zealand (Manos 1997).
The greatest diversity of species of Nothofagus in south-eastern Australia hap-
pened between the Oligocene and Miocene, including the four subgenera and 10
species (Hill 2001a, b). Currently there are only two species in Australia (Table 9.1).
The extinction in Australia includes the subgenera Brassospora and Nothofagus
(as well as 10 genera of the Podocarpaceae and two genera of the Araucariaceae,
according to Crisp et al. 2004).
9.4 Vicariance v/s Dispersal and Centres of Origin

The current disjunct distribution of the subgenera and the lack of any obvious means
of trans-oceanic dispersal have led many authors to infer a vicariance explanation
for the distribution, related to the fragmentation of Gondwana (Van Steenis 1971;
Humphries et al. 1986; Gottlieb 2001). However, the species of the four extant
subgenera do not group neatly into four geographic regions; instead there is consid-
erable overlap (Heads 2006) (Fig. 9.4). There is also no clear congruence between
species cladograms and the breakup sequence of Gondwana, and so a simple vicari-
ance model has been rejected (Knapp et al. 2005; Cook and Crisp 2005). Thus
discussion on the biogeography of the genus has developed into the usual vicariance
vs. dispersal debate (Sect. 3.3), and shows little sign of resolution (Heads 2006).
The impression of JD Hooker upon his early visit to the southern territories
was a scenario of an ancient continuous biota (including Nothofagus and other
disjunct groups), then fragmented by geological and climatic events. Later biogeog-
raphers invented complicated narratives including centers of origin and dispersal.
Darlington (1965) proposed, for example, a center of origin of Nothofagus in
the northern hemisphere (Asia), with subsequent migration events to the South.
Cranwell (1963), on the contrary, proposed a southern origin and subsequent
radiation. Van Steenis (1971) raised the idea of contemporary developments of
Fagus-Nothofagus from a parental “Fagacean” matrix in Southeast Asia, where
there are now representatives from nearly every genus of the family Fagaceae. Van
Steenis supported his evolutionary scenario by means of “land bridges”, whose
emergence at different times would have connected the territories of the extant
disjunct species (see Sect. 10.4).
This hypothesis has been replaced by modern models of plate tectonics, giving
support for the vicariance hypothesis, and several authors found that the phe-
nomena of speciation are quite consistent with the events of fragmentation of
Gondwana from the Cretaceous onwards (Linder and Crisp 1995). Swenson et al.
(2001) specifically proposed vicariance events affecting the three subgenera: an
event in the subgenus Lophozonia that occurred in the Eocene (35 mya), which
corresponds to the separation of Antarctica/South America/Australia (Chap. 1);
an event affecting Fuscospora when New Zealand/New Caledonia separated from
Gondwana (Cretaceous, 80 mya), and which corresponds to the separation between
New Zealand and New Caledonia and the rest of Gondwana; and possible also in the
9.4 Vicariance v/s Dispersal and Centres of Origin 261
Cretaceous an event that would have separated the species of Brassospora between
New Caledonia and New Guinea. Disjunct taxa that are not explained by vicariance
are explained as the result of long-distance dispersal, as in N. gunnii, N. alessandrii
and N. menziesii.
Recently, Knapp et al. (2005) calibrated a relaxed molecular clock for 12 species
(not including the subgenus Brassospora) using fossil ages, i.e. minimum ages.
This analysis suggests that the relationships of the Australian and New Zealand
Lophozonia and Fuscospora species are too recent to have roots in Gondwana,
suggesting long-distance oceanic dispersal. The evolutionary relationship between
the Australasian and South American Fuscospora lineages, however, seems to be
consistent with vicariance.
Knapp et al. (2005) provided the first “unequivocal molecular clock evidence”
that, whilst some Nothofagus transoceanic distributions are consistent with vicari-
ance, trans-Tasman Sea distributions can only be explained by long-distance
dispersal (see discussion in Box 3.3). Knapp et al. (2005) interpreted the absence
of Lophozonia and Fuscospora pollen types in the New Zealand Cretaceous as evi-
dence for Cenozoic dispersals of Nothofagus to New Zealand. The origins of New
Zealand’s biota have been the subject of much debate due to many authors proposing
a complete drowning of these islands in the Oligocene and a relative recent recolo-
nization (see Box 10.1). But others are much more cautious in the interpretation of
the geological evidence.
Knapp et al. (2005) suggested that the biogeographic history of Nothofagus is
more complex than envisaged under simplistic dispersal or vicariance scenarios.
Also Cook and Crisp (2005) emphasized that both vicariance and dispersal played
a role in the distributional patterns of extant Nothofagus. But Heads (2006) pro-
posed that “the vicariant main massings of the four subgenera are compatible with
largely allopatric differentiation and no substantial dispersal since at least the Upper
Cretaceous (Upper Campanian), by which time the fossil record shows that the four
subgenera had evolved”. He further suggested that the complex geological origin
of New Caledonia-New Guinea in which several allochtonous terranes played their
role, has to do with the early differentiation of species in the subgenus Brassospora.
He finally proposed that the breakup of Gondwana, was possibly too late to have
been involved in the evolution of the extant subgenera. All four extant subgenera
have a fossil record in the Upper Cretaceous, which gives a minimum age for all of
them (Heads 2006).
Knapp et al. (2005) and Heads (2006) coincided in calling for the need for
caution in evaluating fossil evidence. The fossil record doesn’t necessarily cap-
ture when a species first appeared. Knapp et al. finally proposed that “the strength
of our molecular analyses highlights the importance of future research into poten-
tial mechanisms of long-distance dispersal, and in particular reinvestigation of the
transoceanic dispersal properties of Nothofagus seeds”. There seems not too much
to do in this respect, however, since the limited dispersal capacity of the fruits was
already experimentally tested by Preest (1963), who showed that they cannot float
and are not transported by birds. Burrows and Lord (1993) further calculated a limit
of 10–15 km for wind transport.
9.5 Nothofagus and Associated Taxa

Several attempts have been made to integrate the biogeographic analysis of
Nothofagus with its parasites, (e.g. Humphries et al. 1986). A very interesting
relationship is that composed by several Nothofagus species and their associated
Cyttaria species. Cyttaria is a fungi genus exclusively related to Nothofagus
species, encompassing 7 species in Chile/Argentina and 4 in Australasia (Barrera
2004; McKenzie et al. 2000) (Fig. 9.1).
The phylogenetic hypothesis of this group (Crisci et al. 1988) shows an interest-
ing relationship with the phylogeny of Nothofagus (Fig. 9.7): the most basal species
of Cyttaria parasitize on species of the South American subgenus Nothofagus, while
the most derived Cyttaria species parasitize on Lophozonia species, both South
American and Australasian.
An independent phylogenetic analysis supports the monophyly of these two sub-
genera of Nothofagus. However, the phylogeny of Cyttaria challenges the relations
between the subgenera, suggesting a closer relationship between Lophozonia and
Nothofagus. Specifically regarding the relations of parasitism, “Fahrenholz’s rule”
predicts that in groups of permanent parasites the classification of the parasites usu-
ally corresponds directly to the natural relationships of the hosts (Mitter and Brooks
1983; Humphries et al. 1986).
These facts aggregate another point of complexity to the already questioned
dispersal capacity of Nothofagus. Nothofagus menziesii, usually treated as a long-
distance dispersal taxon (Knapp et al. 2005), is parasited by three species of Cyttaria
of which C. pallida and C. nigra are exclusive parasites. That a species could
disperse with its own parasites is as improbable as an orchid dispersing with its
mycorrhiza (Sect. 3.3).
C. septentrionalis
C. nigra Parasites subgenus

Lophozonia in Australia,
C. pallida New Zealand, Tasmania.
C. gunnii
C. berteroi Parasites subgenus

C. espinosae Lophozonia in South America
C. exigua
C. darwinii
Parasites subgenus
C. hariotii
Nothofagus (South America)
C. johowii
C. hookeri
Fig. 9.7 Cyttaria species and their hosting subgenera (from the phylogeny by Crisci et al. 1988)
References 263
9.6 Synthesis and Outlook

In spite of the fact that Nothofagus has long been a traditional study object in bio-
geography, few facts really show “unequivocal evidence” (sensu Knapp et al. 2005).
Results from the most recent studies on extant and fossil leaves (Li and Zhou 2007),
on molecular phylogeny (Acosta and Premoli 2010), on molecular dating (Knapp
et al. 2005), and species main massings (Heads 2006) suggest several challenges for
the future research.
(1) Nothofagus most likely originated in high latitude regions of the Southern
Hemisphere, encompassing Antarctica, southern South America, New Zealand
and Australia, during the Early Campanian (Late Cretaceous). The genus
quickly diversified there, and then dispersed gradually to lower latitudes during
the Latest Cretaceous and Palaeogene; whether this process of dispersal was
over terrestrial surfaces or across oceanic barriers is still a matter of discussion;
(2) A more profound analysis of closely related taxa of host-specific parasites, like
Cyttaria, could throw more light on this still controversial issue. Other relations
of parasitism like the one with the temperate South American Misodendrum
could be also explored (see Muñoz-Schick 2004)
(3) Nothofagus fossil pollen is usually considered as more reliable than macro-
fossils like leaf impressions (Hill 2001a), but Li and Zhou (2007) argue that
research on fossil leaf impressions has emerged into a new era, greatly con-
tributing to the investigations on biogeography, evolution and systematics.
Specific morphological features still need to be analyzed under contrasting
evolutionary scenarios/hypotheses.
(4) Molecular phylogenies need to be based on both nuclear and chloroplast
DNA, together with comprehensive sampling methods (Acosta and Premoli
2010). Recent results from phylogeographic analysis of Nothofagus are adding
interesting results to longstanding controversies like the “refugial debate”
(Azpilicueta et al. 2009) (Box 1.7). In molecular dating, the interpretation of
minimal v/s maximal ages in the fossil record is especially critical (Heads
2006).
(5) All this means that Nothofagus, extant and fossil, will most probably con-
tinue to be a “key genus in plant geography”. The gained status as endangered
species of several members of the family, as is the case with Nothofagus
alessandrii (see Sect. 6.2), confirm the urgent necessity of improve conservation
actions on order to assure the possibilities of future research in this challenging
biogeographic group.
References
Acosta MC, Premoli AC (2010) Evidence of chloroplast capture in South American Nothofagus
(subgenus Nothofagus, Nothofagaceae). Mol Phylogenet Evol 54:235–242
Ashworth AC, Cantrill DJ (2004) Neogene vegetation of the Meyer Desert Formation (Sirius
Group) Transantarctic Mountains, Antarctica. Palaeogeogr Palaeoclimatol Palaeoecol 213:
65–82
Ashworth AC, Lewis AR, Marchant DR (2007) Mid-Miocene extinction in Antarctica. 2007 GSA
Denver annual meeting (28–31 Oct 2007)
Azpilicueta MM, Marchelli P, Gallo LA (2009) The effects of Quaternary glaciations in Patagonia
as evidenced by chloroplast DNA phylogeography of Southern beech Nothofagus obliqua. Tree
Genet Genomes 5:561–571
Barrera E (2004) Especies chilenas de Cyttaria Berkeley (Cyttariaceae). Rev Chagual (Jard Bot
Santiago) 2:62–65
Burrows CJ, Lord JM (1993) Recent colonisation by Nothofagus fusca at Cass, Canterbury. NZ J
Bot 31:139–146
Convey P, Gibson JAE, Hillenbrand C-D, Hodgson DA, Pugh PJA, Smellie JL, Stevens MI (2008)
Antarctic terrestrial life – challenging the history of the frozen continent? Biol Rev 83:103–117
Cook LG, Crisp MD (2005) Not so ancient: the extant crown group of Nothofagus represents a
post-Gondwanan radiation. Proc R Soc B Biol Sci 272:2535–2544
Cranwell LM (1963) Nothofagus: living and fossil. In: Gressitt JL, (ed) Pacific basin biogeography.
Bishop Museum Press, Honolulu, HI, pp 387–400
Crisci JV, Gamundí I, Cabello MN (1988) A cladistic analysis of the genus Cyttaria (Fungi-
Ascomycotina). Cladistics 4:279–290
Crisp M, Cook L, Stean D (2004) Radiation of the Australian flora: what can comparisons of
molecular phylogenies across multiple taxa tell us about the evolution of diversity in present-
day communities? Philos Trans R Soc B Biol Sci 359:1551–1571
Darlington PJ Jr (1965) Biogeography of the southern end of the world. Harvard University Press,
Cambridge, MA
Denton GH, Prentice ML, Burckle LH (1991) Cainozoic history of the Antarctic ice sheet. In:
Tingey RJ, (ed) The geology of Antarctica. Clarendon Press, Oxford, pp 365–433
Dettmann ME, Pocknall DT, Romero EJ, Zamaloa MdelC (1990) Nothofagidites Erdman ex
Potonie 1960: a catalogue of species with notes on the paleogeographic distribution of
Nothofagus BL (Southern Beech). N Z Geol Surv Palaeontol Bull 60:1–79
Donoso C (1993) Bosques templados de Chile y Argentina: variación, estructura y dinámica.
Editorial Universitaria, Santiago
Donoso C, Morales J, Romero M (1990) Hibridación natural entre roble (Nothofagus obliqua
(Mirb.) Oerst.) y raulí (Nothofagus alpina (Poepp. & Endl.) Oerst.) en bosques del sur de Chile.
Eklund H (2003) First Cretaceous flowers from Antarctica. Rev Palaeobot Palynol 127:187–217
Falcon-Lang HJ, Cantrill DJ, Nichols GJ (2001) Biodiversity and terrestrial ecology of a mid-
Cretaceous, high-latitude floodplain, Alexander Island, Antarctica. J Geol Soc 158:709–724
Francis JE, Poole I (2002) Cretaceous and Tertiary climates of Antarctica: evidence from fossil
wood. Palaeogeogr Palaeoclimatol Palaeoecol 182:47–64
Francis JE, Ashworth A, Cantrill DJ, Crame JA, Howe J et al. (2008) 100 million years of Antarctic
climate evolution: evidence from fossil plants. In: Cooper AK, Barrett PJ, Stagg H, et al. (eds)
Antarctica: a keystone in a changing world. Proc 10th international symposium on Antarctic
earth sciences. National Academies Press, Washington, DC, pp 19–27
Gottlieb M (2001) Zur Phylogenie un Phyto-/Paläophytogeographie der Südbuchen. Gattung
Nothofagus. Mitt Dtsch Dendrol Ges 86:85–106
Heads M (1985) Biogeographic analysis of Nothofagus (Fagaceae). Taxon 34:474–480
Heads M (2006) Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species
massings. J Biogeogr 33:1066–1075
Hill RS (2001a) Biogeography, evolution and palaeoecology of Nothofagus (Nothofagaceae): the
contribution of the fossil record. Aust J Bot 49:321–332
Hill RS (2001b) Nothofagus cupules from Late Oligocene-Early Miocene sediments at Balfour,
northwest Tasmania, Australia. Int J Plant Sci 162:683–690
Hill RS, Read J (1991) A revised infrageneric classification of Nothofagus (Fagaceae). Bot J Linn
Soc 105:37–72
Hill RS, Jordan GJ (1993) The evolutionary history of Nothofagus (Nothofagaceae). Aust Syst Bot
6:111–126
References 265
Hill RS, Dettmann ME (1996) Origin and diversification of the genus Nothofagus. In: Veblen TT,
Hill RS, Read J, (eds) The ecology and biogeography of Nothofagus forests. Yale University
Press, New Haven, CT, pp 11–24
Humphries CJ (1981) Biogeographic methods and the southern beeches. In: Forey PL, (ed) The
evolving biosphere. Cambridge University Press, Cambridge, pp 283–297
Humphries CJ, Cox JM, Nielsen ES (1986) Nothofagus and its parasites: a cladistic approach to
coevolution. In: Stone AR, Hawksworth DL, (eds) Coevolution and systematics. Clarendon
Press, Oxford, pp 55–76
Knapp M, Stöckler K, Havell D, Delsuc F, Sebastiani F, Lockhart PJ (2005) Relaxed molecular
clock provides evidence for long-distance dispersal of Nothofagus (southern beech). PLoS Biol
3:38–43
Lawver LA, Gahagan LM (1998) Opening of Drake Passage and its impact on Cenozoic ocean
circulation. In: Crowley TJ, Burke KC, (eds) Tectonic boundary conditions for climate recon-
structions. Oxford Monographs on Geology and Geophysics 39. Oxford University Press,
Oxford, pp 212–223
Lawver LA, Gahagan LM (2003) Evolution of Cenozoic seaways in the circum-Antarctic region.
Palaeogeogr Palaeoclimatol Palaeoecol 198:11–37
Li RQ, Chen ZD, Lu AM, Soltis DE, Soltis PS, Manos PS (2004) Phylogenetic relationships in
Fagales based on DNA sequences from three genomes. Int J Plant Sci 165:311–324
Li HM, Zhou ZK (2007) Fossil nothofagaceous leaves from the Eocene of western Antarctica and
their bearing on the origin, dispersal and systematics of Nothofagus. Sci China Ser D Earth Sci
50:1525–1535
Linder HP, Crisp MD (1995) Nothofagus and Pacific biogeography. Cladistics 11:5–32
Livermore R, Hillenbrand C-D, Meredith M, Eagles G (2007) Drake Passage and Cenozoic
climate: an open and shut case? Geochem Geophy Geosyst 8:Q01005. doi: 10.1029/2005
GC001224
Manos PS (1997) Systematics of Nothofagus (Nothofagaceae) based on rDNA spacer sequences
(ITS): taxonomic congruence with morphology and plastid sequences. Am J Bot 84:1137–1155
McKenzie EHC, Buchanan PK, Johnston PR (2000) Checklist of fungi on Nothofagus species in
New Zealand. N Z J Bot 38:635–720
Mitter C, Brooks DR (1983) Phylogenetic aspects of coevolution. In: Futuyama DJ, Slatkin M,
(eds) Co-evolution. Sinauer Associates, Sunderland, MA, pp 65–98
Moreira-Muñoz A (2004) Nothofagus Bl., pieza clave en la reconstrucción biogeográfica del
hemisferio austral. Rev Chagual (Jard Bot Santiago) 2:48–56
Munzinger J (2010) Origin of the Flora of New Caledonia, past and present knowledge. Abstract
southern Connection congress, Feb 15–19, 2010, Bariloche, Argentina
Muñoz-Schick M (2004) El género Misodendrum DC. en Chile. Rev Chagual (Jard Bot Santiago)
2:57–61
Nixon KC (1982) In support of recognition of the family Nothofagaceae Kuprianova. Bot Soc Am
Publ 162:102
Nixon KC (1989) Origins of Fagaceae. In: Crane PR, Blackmore S, (eds) Evolution, systematics,
and fossil history of the Hamamelidae, vol 1. Clarendon Press, Oxford, 23–43
Poole I (2002) Systematics of Cretaceous and Tertiary Nothofagoxylon: implications for southern
Hemisphere biogeography and evolution of the Nothofagaceae. Aust Syst Bot 15:247–276
Poole I, Gottwald H (2001) Monimiaceae sensu lato, an element of Gondwanan polar forests:
evidence from the Late Cretaceous-early Tertiary wood flora of Antarctica. Aust Syst Bot
14:207–230
Poole I, Cantrill DJ (2006) Cretaceous and Cenozoic vegetation of Antarctica integrating the fossil
wood record, vol 258. Geological Society, London, Special Publications, 63–81
Preest DS (1963) A note on the dispersal characteristics of the seed of New Zealand podocarps and
beeches and their biogeographical significance. In: Gressitt JL, (ed) Pacific Basin biogeography.
Bishop Museum Press, Honolulu, Hawaii, pp 415–424
Ramírez C, San Martín C, Oyarzún A, Figueroa H (1997) Morpho-ecological study on the South
American species of the genus Nothofagus. Plant Ecol 130:101–109
Reguero MA, Marenssi SA, Santillana SN (2002) Antarctic Peninsula and South America
(Patagonia) Paleogene terrestrial faunas and environments: biogeographic relationships.
Paleogeogr Paleoclimatol Paleoecol 179:189–210
Rodríguez R, Quezada M (2003) Fagaceae. In: Marticorena C, Rodríguez R, (eds) Flora de Chile,
vol 2. Ed. Universidad de Concepción, Concepción, pp 64–75
Scher HD, Martin EE (2006) Timing and climatic consequences of the opening of the Drake
Passage. Science 312:428–430
Siegert MJ, Barrett P, Decont R, Dunbar R, Cofaigh CO, Passchier S, Naish T (2008) Recent
advances in understanding Antarctic climate evolution. Antarctic Sci 20:313–325
Stecconi M, Marchelli P, Puntieri J, Picca P, Gallo L (2004) Natural hybridization between a decid-
uous (Nothofagus antarctica, Nothofagaceae) and an evergreen (N. dombeyi) forest tree species:
evidence from morphological and isoenzymatic traits. Ann Bot 94:775–786
Swenson U, Hill RS, McLoughlin S (2001) Biogeography of Nothofagus supports the sequence of
Gondwana break-up. Taxon 50:1025–1041
Torres T (1984) Nothofagoxylon antarcticus n. sp., Madera fósil del Terciario de la isla Rey Jorge,
isles Shetland del Sur, Antártica. Ser Cient INACHI 31:39–52
Torres T (2003) Antártica: un mundo oculto bajo el hielo. Instituto Antártico Chileno, Punta Arenas
Torres-Díaz C, Ruiz E, González F, Fuentes G, Cavieres LA (2007) Genetic diversity in Nothofagus
alessandrii (Fagaceae), an endangered endemic tree species of the coastal maulino forest of
Central Chile. Ann Bot 100:75–82
Van Steenis CGGJ (1971) Nothofagus, key genus of plant geography, in time and space, living and
fossil, ecology and phylogeny. Blumea 19:5–98
Veblen TT, Hill RS, Read J (eds) (1996) The ecology and biogeography of Nothofagus forests.
Yale University Press, London
Zachos J, Pagani M, Sloan L, Thomas E, Billups K (2001) Trends, rhythms, and aberrations in
global climate 65 Ma to present. Science 292:686–693
Part V
Where to from Here? Projections of
Chilean Plant Geography
Chapter 10
All the Possible Worlds of Biogeography
For history is like a nymph glimpsed bathing between leaves:

the more you shift perspective, the more is revealed. If you want
to see her whole you have to dodge and slip between many
different viewpoints
(Fernández-Armesto 1997, p 228).
Abstract Today anyone intending to integrate the different views that shape
modern biogeography must confront the differences inherent to the diverse
approaches involved in the discipline. In spite of the attempts to integrate different
approaches into one coherent program of synthetical biogeography, the biogeo-
graphic arena is getting more and more fragmented due to a plethora of methods,
and the ultimate synthesis is getting more and more elusive. The so-called “crisis of
biogeography” seems to be related to a more general crisis of reductionistic mod-
ern science in its failure to account for the real world problems, as challenged by
scientists inspired by postmodern theory. To what extent biogeography assumes and
reflects the conflicts, presumptions and challenges inherent to (post)modern science
must be kept in mind while analysing the Chilean plant geography.
10.1 The Fragmented Map of Modern Biogeography

As noted by Riddle (2005), among others, modern biogeography seems to suf-
fer a protracted identity crisis, since there is an evident lack of fully integrative
approaches. The plethora of methods that are in common use (see a revision by
Crisci et al. 2003; Morrone and Crisci 1995) leaves no opportunity for this integra-
tion. Nelson and Ladiges (2001) characterized current biogeography as a “mess of
methods” (Fig. 10.1).
No few attempts have been undertaken to propose a more integrative research
program in biogeography (e.g. Croizat 1958; Myers and Giller 1988; Salomon 2001;
Crisci et al. 2006). Any proposal will be challenging and conflicting: e.g. Donoghue
and Moore’s (2003) “integrative biogeography” program has been criticized because
it has been used to reinforce equivocal divisions, such as that between phylogeneti-
cally and ecologically focused methods (Parenti and Ebach 2009). Indeed, together

DOI 10.1007/978-90-481-8748-5_10, ⃝C Springer Science+Business Media B.V. 2011
270 10 All the Possible Worlds of Biogeography
Fig. 10.1 Biogeographers’ diverse conceptual and methodological “suitcases”, according to

Udvardy (1981), by permission of Columbia University Press
with the traditional conflict between dispersalism versus vicariance (Chap. 3), one
of the most striking divides has been the one between ecological and historical
biogeography. This “probably reflects the predominance of narrative rather than
analytical methods in biogeography because narratives allow authors to cast their
explanations in terms of rival beliefs rather than rigorous inferences. On the other
hand, when analytical methods are used in biogeography, the patterns obtained
from them may be neither wholly historical nor wholly ecological, and consider-
able refinement, testing and reasoning is needed if the effects of these processes
are to be distinguished” (Rosen 1988, p 33). Several authors have maintained that
this is rather an artificial conflict (e.g. Parenti and Ebach 2009) and there are sev-
eral researchers working towards a more appropriate framework to overcome these
binary opposites to generate a “free play of the opposites” (Crisci and Katinas 2009,
p 1031).
10.2 Postmodern Biogeography: Deconstructing the Map

We are living in a time in which postmodernism is constantly confronting the hegemonic
speech of science to denounce its excesses and to expose its limits (Morrone 2003, p. 87)
After the period of discovery that marked the origin of modern nature sciences (with
A. von Humboldt) “historical and ecological biogeography and their many subdivi-
sions evolved, diverged, and eventually flourished (or languished) as increasingly
more distinct disciplines” (Lomolino and Heaney 2004, p 1). This diversifica-
tion and growth of distinctive scientific disciplines established a presumed need
10.2 Postmodern Biogeography: Deconstructing the Map 271
to specialize that resulted in more and more splintering. As a consequence of sci-

entific specialization “. . .the grand view, the ultimate synthesis across space and
time, became murky and more elusive” (Lomolino and Heaney 2004, p 1). But
there seems to be still hope for this great synthesis, as it might “encourage creative
development and applications of the comparative approach, deconstructing and
reassembling more comprehensive explanations for the diversity and distribution of
biotas” (Lomolino and Heaney 2004, p 2).
The deconstruction suggested by Lomolino and Heaney (2004) is in tune with
the revision that is today occurring in many disciplines. After decades of reduction-
ism, science has failed in helping to construct a more harmonic and inclusive world
for humans and non humans, coming to the current global crisis of ethic values and
nature destruction (Chap. 6). The reaction towards actual scientific splintering has
canalized critics and alternatives in the so called postmodern science. This new pre-
tended scientific paradigm has rapidly gained advocates and opponents (between
the last e.g. Dawkins 1998; Attwell and Cotterill 2000). Advocates maintain that
only postmodern science would “nurture the human hunger for quality: for beauty,
balance, creative advance” (Ferré 1997, see also Dear and Wassmansdorf 1993). In
his context, used and abused dualistic concepts like “objectivity” and “subjectivity”
have been questioned: “objects are to be understood through their relations rather
than as ‘pure objects’. . . our knowledge of objects is neither objective nor subjec-
tive, but a complex outcome of knowing and action in an environment” (Jones 2008,
p 1606).
The critical view of postmodernism left nothing untouched and our Weltbild is
changing rapidly; the potential consequences are immense, and may provide the
only possibility to heal our “fragmented culture” (Goodwin 2007) (see further Sect.
10.5). The way in which we do science is inseparable from the culture and the
historical moment. As already discussed in Chap. 7: “. . .the striking, but culturally
determined, exoticness of the Cactaceae still impacts our concept of what is relictual
and derived for the family” (Griffith 2004).
Perhaps the first explicit proposal for a deconstruction of biogeography is
a guest editorial that appeared in the Journal of Biogeography as: “Mesozoic
tectonics and the deconstruction of biogeography” (Heads 1990). There, Heads
proposed an alternative view for the evolution of the Australasian biota, arguing
that “approaches to biogeography have been based all too often on consideration
of particular lineages, emphasizing purely theoretical ancestor-descendant rela-
tionships and have maintained a blind spot towards the general effect of phases
of modernization on a landscape and its biota”. Since then M. Heads has been
championing the developments in biogeographic theory integrating the findings of
Croizat and philosophers like Derrida (Heads 2005a). Nowadays, deconstruction
is being promoted not as an alternative to traditional academic research, but as a
systematic approach in science: “deconstruction should be consciously performed
as a methodological strategy” (Marquet et al. 2004, p 192; see also Ebach et al.
2003; Terribile et al. 2009). This method favoured recently a better interpretation
of the fossil evidence of Ericaceae in New Zealand (Jordan et al. 2010) (see also
Box 10.1).
Box 10.1 Circular Logic in the Drowning of New Zealand

The best example of the contest between geological versus biogeographical
hypotheses affects interpretations on the origin of the New Zealand biota,
which has been perhaps the most important of any regions in the world for
biogeographic analysis (Nelson 1975; Gibbs 2006). According to a recent
revision by Goldberg et al. (2008) “molecular studies of terrestrial animals
and plants in New Zealand indicate that many taxa arrived since isolation of
the land, and that diversification in most groups is relatively recent”. This is
in accordance with the dispersalist view that most of New Zealand’s biota dis-
appeared during the Oligocene (Pole 1994; Winkworth et al. 2002; McGlone
2005; McDowall 2008), and the area was recolonized over the last 22 mil-
lion years across a huge oceanic gap via long-distance dispersal events, rather
than simply persisting there for 80 million years. Waters and Craw (2006)
criticized the apparent logical circularity in proposing the persistence of sev-
eral groups like Agathis (Fig. 10.2), ratites, wrens, mammals, for 80 million
years in “splendid isolation”. They invoke lack of geological evidence neglect-
ing biogeographical evidence, but finally concluded that “available geological
data thus neither confirm nor reject Oligocene drowning of New Zealand”
(Waters and Craw 2006, p 352).
a b
Fig. 10.2 Southern forests: a broad-leaved forest with Agathis and ferns at Coromandel,
New Zealand; b Valdivian forest at Oncol, Chile. Photo credits: a A. Moreira-Muñoz;
b S. Elórtegui Francioli
10.3 Sloppy Biogeography v/s Harsh Geology? 273
In support of the long splendid isolation is the continuous presence of

Agathis in New Zealand (Stöckler et al. 2002; Lee et al. 2007), together with
other plant taxa. Wanntorp and Wanntorp (2003) propose a continuous pres-
ence of Gunnera since the Cretaceous and others appoint for drastic drowning,
but not all submersion of land (to ca 20% of the present land area, sensu
Cooper and Cooper 1995). If we have a look at two independent recent global
palaeoreconstructions, New Zealand does not appear submerged in the last
120 million years, neither at 25 mya in the Oligocene (Hall 2002; Sdrolias
et al. 2003).
According to Michaux and Leschen (2005) “we have no way of knowing
how much of the microcontinent [the Campbell Plateau] was above water by
25 mya: some must have done so to account for the persistence of palaeo-
endemics”. They concluded that “the New Zealand Subantarctic Islands,
Fiordland, Southland, Otago, and southern South America preserve a unique
collection of west Gondwanan flora and fauna“. These findings are coher-
ent with results by Knapp et al. (2007): “ratites, wrens, mammals, and New
Zealand Araucariaceae fossils do not support the hypothesis that New Zealand
was completely submerged during the Oligocene”. Physiognomic and floristic
similarities between southern Chile and New Zealand last till today (Ezcurra
et al. 2008).
Newest evidence from faunistic groups seems to support the continen-
tal nature and the persistence of New Zealand’s biota: i.e. in the case of
centipedes and meat harvestmen (Edgecombe and Giribet 2008; Boyer and
Giribet 2009). Recent evidence also suggests that the subterranean freshwater
fauna survived the presumed Oligocene inundation of New Zealand (Wilson
2008), as well as the fossil ancestors of extant tuatara recently found and dated
at 19–16 mya (early Miocene) (Jones et al. 2009).
The dispersalist view was already contested by JD Hooker (see Sect. 3.3):
“If the number of plants common to Australia and New Zealand is great, and
quite unaccountable for by transport, the absence of certain very extensive
groups of the former country is still more incompatible with the theory of
extensive migration by oceanic or aerial currents. This absence is most con-
spicuous in the case of Eucalypti, and almost every genus of Myrtaceae, of the
whole immense genus of Acacia, and its numerous Australian congeners . . .”
(Hooker 1853, as quoted by Turrill 1953, p 152).
10.3 Sloppy Biogeography v/s Harsh Geology?

Evolution is not a constant battle between the eaters and those to be eaten, but rather changes
in organisms that are living and surviving on a dynamic Earth (Ebach and Humphries 2003).
As explained by McLoughlin (2001), detailed biogeographic understand-

ing of the Southern Hemisphere floras must ultimately depend on a thorough
understanding of the timing of geological and biological events that led to mod-
ern floristic distributions. In this sense, the fragmentation of Gondwana should have
a big impact on biogeography and evolution of the southern floras. For most parts
of Gondwana, the amalgamation and breakup phases and palaeoclimatic regimes
are well documented via palaeomagnetic studies, radiometric dating, tectonostrati-
graphic analyses, palaeontological investigations and comparison of sedimentary
facies (McLoughlin 2001) (see Table 1.1). Biogeographic hypotheses are being con-
fronted with palaeogeographic scenarios, in an attempt to discover the processes
underlying patterns. Biogeographic explanations are thus getting complicated: dif-
ferent models for the Gondwana fragmentation are entering the discussion and none
of them fits well for every biotic group (Krause et al. 2006; Upchurch 2008).
Upchurch et al. (2002), expressed it in this way: “Clearly, there is not a perfect fit
between the biogeographical patterns and palaeogeographical history, but there are
several reasons why it would be premature to reject the biological signal: (i) palaeo-
geographical reconstructions are themselves hypotheses that potentially contain
errors; (ii) congruence may increase as time-slicing and area selection are refined;
(iii) the degree of congruence partly depends on a priori expectations regarding the
effect of barriers on dispersal (e.g. phylogenetic divergence may commence before
a barrier is fully developed); and (iv) the repeated area relationships are statistically
supported signals that stand by themselves as patterns that require explanation”
(Upchurch et al. 2002, pp 618–619). Croizat (1958) was the first to call for this
independent view of biogeography as a discipline with own methods and tools.
He proposed that biogeographers should not base all their study on geologically
“well established” hypotheses. Biogeography, as a mature and independent disci-
pline, should be able to develop its own hypotheses and theories to compare these
with the geological theories. That is the way Alfred Wegener (1915) could develop
his worldwide accepted theory (continental drift). At his time he was emphatically
criticized by most geologists, but in the end the evidence imposed itself.
In the words of Heads (2005b, p 64): “assuming a priori that any particular geo-
logical event, such as the break-up of Gondwana, is relevant to biogeography is
a fatal flaw of much biogeography, both dispersalist and vicariance. . . In fact, a
great deal of evidence suggests biogeographic patterns involving New Zealand, New
Guinea, New Caledonia, etc. were determined by earth history events both prior and
subsequent to the break-up of Gondwana”.
In concordance with this, recently some authors are calling for avoiding the
circular logic in confronting biogeographical hypothesis with geological evidence
(e.g. Renner 2005; Waters and Craw 2006) (Box 10.1). “Constraining nodes in a
phylogenetic tree by geological events risks circularity in biogeographical analy-
ses because it already assumes that those events caused the divergence, rather than
testing temporal coincidence” (Renner 2005, p 552).
“Unfortunately, at the time we wrote this paper [on Proteaceae], we were mis-
led by conservative geologists who had not got around to accepting continental
drift, and our phytogeographic understanding was much distorted by this” (Johnston
1998, as quoted by McCarthy 2005b). McCarthy continues: “some still tend to
elevate geological speculation over basic distributional realities. Implicit in papers
10.4 Just Some Possible Worlds 275
that indulge in extravagant dispersalism and a plethora of just-right fossil absences

is the notion that the basic principles of biogeography are wispy and yielding while
geophysical theories are made of sterner stuff. Such papers appear to extend the
legend that planetary scientists work in a field devoid of speculation, the belief that
when a biogeographer and a geologist confront each other on a narrow path, the
biogeographer must step aside” (McCarthy 2005b).
Under a vicariance paradigm the classical pre-drift reconstruction of Pangea cannot ade-
quately explain trans-Pacific tracks. Therefore, alternative paleogeographic models may
be invoked as explanatory hypotheses: the lost continent Pacifica, island integration, a
new reconstruction of eastern Gondwanaland, an expanding earth. None of these alter-
native models is fully compatible with all geological and biogeographic data available at
present. It is stressed that biogeographic data and theories should not be made subservient
to geological theories (Sluys 1994, p 42).
Also Linder and Crisp (1995) found that the biogeographic pattern found in
Nothofagus (Chap. 9) was not congruent with geological hypotheses and wrote:
“This is not congruent with the current geological theories, nor with the pat-
terns evident from insect biogeography. We suggest that concordant dispersal is an
unlikely explanation for this pattern, and propose that the solution might be found
in alternative geological hypotheses” (Linder and Crisp 1995, p 5).
Here lies a big paradox in modern biogeography: palaeoreconstructions have
been traditionally done integrating geological and paleontological information, i.e.
biogeographical evidence, stratigraphic evidence, isotopic signature and palaeomag-
netic data (Rapalini 2005). The integration of these different lines of evidence is
full of conflict depending on the point of view and the data analysed. But usually
botanists take the most accepted reconstruction, and try to fit the extant disjunct
distributions with the major events in e.g. the splitting of Gondwana (e.g. split of
Africa-America, split of Antarctica-South America, see Table 1.1). But the regional
tectonic reconstructions suggest that the tectonic history is much more complicated
and everywhere in the southern hemisphere there are geological and biotic compos-
ite areas: in Australasia (Morley 2001), New Zealand (Craw 1988), the Subantarctic
Islands (Michaux and Leschen 2005), Tasmania (Heads 1999), New Guinea (Heads
2002) and certainly in southern South America (Crisci et al. 1991; Katinas et al.
1999).
10.4 Just Some Possible Worlds

As revised in the preceding section, alternative biogeographic scenarios are often
in conflict with tectonic reconstructions, and gain therefore much less accep-
tation in the scientific community. Alternative palaeoreconstructions rest often
on alternative methods that are not fully developed, like Dobson’s (1992) spa-
tial logics, which allows building alternative hypothesis of plates’ fragmentation.
Palaeoreconstructions with specific biogeographic implications for the Chilean flora
will be briefly revised hereafter.
10.4.1 Connections Over Land Bridges

The idea of former land bridges on the sites of today’s oceans, contrary to the
current paradigm of plate tectonics, is not an old one, and like a circular idea,
is constantly re-presented and re-rejected. Probably the first one who proposed
and illustrated former lands between today’s continents was Hermann von Ihering
(1907), while studying the biogeographical connections of the Neotropical biota.
He proposed ancient land connections between America and Africa and between
Antarctica and Australasia that he called Archinotis and Archhelenis, respectively
(Fig. 10.3a). Also Croizat (1952) proposed vast emerged lands between the shapes
of current continents (Fig. 10.3b). The theory of land bridges was extendedly
defended by CG van Steenis (1962). The idea of the land bridges has been recently
called for again by Morley, explaining the biogeographical relationships between
the American and African tropics (Morley 2003), the pantropics (Zhou et al. 2006)
and between Cenozoic disjunct relict floras from Eurasia – North America (Milne
2006). Davis et al. (2002) favoured Palaeocene-Oligocene migrations between
North America and South America via the Caribbean Basin for the Malpighiaceae,
avoiding the classical dichotomy vicariance/long-distance dispersal.
10.4.2 And What About a Closer Pacific Basin?

If we remove the Pacifica hypothesis we are left to choose another causal explanation for
the repeating pattern of area (floristic) relationships, and other parts of the scenario remain
intact. Even if we remove some taxa, we do not change anything significant about the whole
picture. But if we remove the initial transpacific area pattern, everything collapses: there is
no repeating pattern from which to extrapolate and for which to build complex scenarios
(Grande 1994, p 76).
As discussed in Chap. 5 for the Juan Fernández islands, the floristic relationships
between the islands and the austral territories of South America call for a better
explanation than a simple relationship to the current closer territory. The proposal
of a Pacifica continent West of South America fits perfectly within this scheme.
This old idea has been revitalized under the umbrella of vicariance biogeography
(Nur and Ben-Avraham 1977, 1981; Kamp 1980; Dickins et al. 1992) (Fig. 5.10b).
The hypothesis has been firmly rejected on the base of geological arguments, not
based on biogeographical patterns (Cox 1990). Ocean surface maps certainly do not
leave us recognizing anything like a sunken continent west of South America, but
we could still question if the old land was perhaps sunken into the trench?
The formation of the Pacific plate is crucial in this sense but has still many
unresolved questions. It seems that it began to form in the Mid-Jurassic at the
Tongareva triple junction (Larson et al. 2002). From here, the Pacific plate expanded
at the expense of the Phoenix, Farallon and Izanagi plates that surrounded it (Smith
2007). Terrane accretion on the Gondwana–Pacific margin seems to have accom-
panied the plate formation (Vaughan and Livermore 2005). These authors proposed
a Late Triassic–Early Jurassic deformation associated with the Pangaea breakup,
Fig. 10.3 a Land bridges proposed by von Ihering (1907), called Archinotis and Archhelenis; b
land bridges proposed by Croizat (1952); numbers represent evolutionary “gateways”
and a mid-Cretaceous deformation. Possible effects of the last event are evident for
the austral biogeography but the earlier one was probably just as important (Heads
2009).
In close relationship to the Pacifica continent hypothesis, lays the expanding
Earth theory. First proposed by Lindemann (1927) and Hilgenberg (1933), it the-
orizes that all continents were united to form a single crust that encompassed a
much smaller pre-Jurassic globe. The theory lost force during the twentieth cen-
tury, in spite of being still defended by some researchers (e.g. Carey 1988; Vogel
1990). The theory was reanalysed specifically for biogeography by Shields (1998)
and more recently by McCarthy (2003, 2005a,b, 2007).
McCarthy (2003) noted that the juxtaposition of New Zealand and southern Chile
on expanding Earth palaeomaps provides a simple explanation (i.e. most parsimo-
nious) for the distributions without need for additional hypotheses. According to
this view, the oceanic crust did not begin forming between New Zealand and South
Chile until 83.5 mya and even then the amount of Pacific Ocean crust between the
two locations did not become significant for many tens of millions of years. “If the
existence of Panthalassa is not assumed, then New Zealand and Chile would have
remained in proximity until the Late Cretaceous and would have been separated by
a narrow sealike barrier during the Eocene” (McCarthy 2003, p 1542). The close
floristic relationship between the austral territories, i.e. the austral floristic realm
(Chap. 4), and the persistence of numerous ancient groups in New Zealand seem
to support this hypothesis. As recognized by Stevens (1997), closure of the Pacific
Ocean considerably simplifies the links that occurred between New Zealand, West
Antarctica and South America (Fig. 10.4).
These arguments are far from convincing traditional biogeographers, which para-
doxically are rejecting the theory on the base of geologic arguments, not on the base
a b
North
America
Africa
Asia
South
America
Australia
India
New Antarctic peninsula

Zealand
Antarctica
Fig. 10.4 Contrasting palaeoreconstructions: a the traditional model for the Late Jurassic based
on the Time Trek program; b position of Australia and New Zealand close to South America
considering a close Pacific basin (redrawn from Stevens 1997)
of biogeographical signal (Cox 1990; Briggs 2004; Ali 2006). But the theory is far
from been amply rejected, as expressed by the collection of papers by the Italian
geophysist G Scalera and collaborators (Scalera 2006, 2008; Scalera and Jacob
2003). Scalera (2007) proposes several methodological and theoretical aspects for
gaining advance in this research field, like the development of changing-radius
cartography and the reinterpretation of palaeomagnetism according to a different
globe’s radius at the epoch of primary magnetization. As a sign of the many chal-
lenges affecting modern biogeography, an expanding globe illustrates the cover of
the book Biogeography in a Changing World (Ebach and Tangney 2007).
10.4.3 Three Models of Gondwana Fragmentation +

One Dispersal
Not only radical views have to be considered as alternative explanations. When

the model of Gondwana fragmentation has gained general acceptance, emerging
views are challenging the timing of separation of the different landmasses and its
implications for biogeography. As an example, most plate tectonic reconstructions
assume that major continental blocks (Eurasia, Greenland, North America, South
America, Africa, India, Australia and Antarctica) have separated from one another
by the end of the Early Cretaceous. Hay et al. (1999) suggests this simplistic view
may be incorrect, proposing instead three large continental blocks (North America-
Eurasia; South America-Antarctica-India-Madagascar-Australia; and Africa) with
large contiguous land areas surrounded by shallow epicontinental seas (Hay et al.
1999). Krause et al. (2006), explicitly compare two different separation scenarios
for explaining the known distribution of Late Cretaceous terrestrial vertebrates from
Madagascar (Fig. 10.5).
Upchurch (2008) newly proposed a practical approach for testing different
models of Gondwana fragmentation with biogeographic data. Upchurch (2008)
grouped the various models of Gondwanan biogeography in three alternative break-
up models and one dispersal model: (a) Samafrica model (South America and
Africa): break-up of Gondwana beginning in the Jurassic (175–140 mya), along
a line from Somalia in the North to the Weddell Sea in the South; (b) Africa-first
model: according to this vision Africa became isolated 140–120 mya whereas
South America remained in contact with East Gondwana until ~80 mya; (c) Pan-
Gondwana model: Gondwanan areas remained connected until ~80 mya and then
separated almost simultaneously. The Pan-Gondwana model does not predict vicari-
ance, because the simultaneous, rather than sequential, formation of barriers should
not impose a distribution pattern on taxa and phylogenies (Upchurch 2008). These
models are not necessarily exclusive.
At a more detailed scale, uncertainties persist regarding the separation of spe-
cific territories: “The time of onset of the ACC (Antarctic Circumpolar Current) is
uncertain, and abundant speculation exists about its onset and effects. As knowledge
of modern ocean circulation and the uses of the geological record have developed,
Pan-Gondwana Hypothesis Africa-first Hypothesis
C C
C
C
C
C
120 mya
C C
C C
100 mya
? C
C
80 mya
C
60 mya
Fig. 10.5 Two different fragmentation scenarios for explaining the distribution of Late Cretaceous
terrestrial vertebrates from Madagascar: C = connection; arrow = separation (adapted from Krause
et al. 2006)
and as the palaeoceanographic data base has grown, so speculation has been refined,
but the uncertainties in the ACC onset and subsequent variation remain significant”
(Barkera and Thomas 2004) (see also Boxes 1.3 and 9.1).
Upchurch (2008) finished his discussion of the different models of Gondwanan
fragmentation describing trans-oceanic long distance dispersal as a fourth biogeo-
graphical model, and he concluded that the current focus on vicariance versus
dispersal is too narrow because it ignores geodispersal (i.e. expansion of species

into areas when geographical barriers disappear), extinction, sampling errors, and
palaeogeographical uncertainty (Upchurch 2008, pp 234–235).
He further emphasizes five critical points for the advancement in the field:
(1) conclusions arisen from current studies that seek to evaluate the importance of
vicariance versus long-distance dispersal are dubious if they do not assess the
effects of reticulation and distributional noise;
(2) palaeontological studies usually develop synthetic scenarios lacking analyti-
cal and statistically explicit methods that distinguish “genuine biogeographical
patterns from distributional noise”;
(3) too often researchers tend to oversimplify biogeographical reticulation assum-
ing that there is a single vicariance pattern caused by, and therefore congruent
with, Gondwanan break-up;
(4) molecular biologists usually fail to explore (or even mention) uncertainties in
palaeogeographical and palaeoclimatic reconstructions;
(5) a false impression that long-distance dispersal is the correct explanation for the
distributions of organisms can appear as the consequence of sampling errors
and multiple signals (Upchurch 2008, p 235).
He suggests that specialized methods (like pattern-spotting) “must be applied if an

unbiased understanding of southern hemisphere biogeography is to be achieved”
(Upchurch 2008, p 229). This seems to be in accordance with the more inte-
grated and complementary programs in biogeography, like systematic biogeogra-
phy (Williams and Ebach 2008), and evolutionary biogeography (Morrone 2007,
2009) (Box 10.2).
Box 10.2 Evolutionary Biogeography: Conciliating

Distributions, Palaeogeographies, and Evolution
An evolutionary biogeographical analysis may involve five steps (Morrone

2007, 2009):
(1) recognition of biotic components (sets of spatio-temporally integrated

taxa due to common history), through panbiogeography and methods
used to identify areas of endemism;
(2) contrasting of the biotic components and identification of the vicari-
ant events that fragmented them, through cladistic biogeography and
comparative phylogeography;
(3) establishment of a hierarchic arrangement of the components in a bio-
geographic system of realms, regions, dominions, provinces and districts
(i.e. biogeographic regionalization);
(4) identification of cenocrons (sets of taxa with similar origins and ages),
dated using intraspecific phylogeography, molecular clocks and fos-
sils; and
(5) formulation of a geobiotic scenario, that explains the evolution of the
biotic components and cenocrons, integrating geological and tectonical
information.
This evolutionary biogeography is a great step towards a more integrated

biogeography and explicitly considers the systematic classification of areas,
i.e. a systematic biogeography (cfr Williams and Ebach 2008). In the search
for an even more integrative program, Parenti and Ebach (2009) propose a
comparative biogeography that integrates both systematic and evolutionary
biogeography.
These are good signals towards more integrated and analytical approaches
in biogeography that will gain a more accurate picture from complex and
mostly elusive patterns and processes in an increasingly changing and threat-
ened global biota.
10.5 The “New Biogeography”

. . . most of us are held captive by the notion of the world “out there”, separate from us
“in here.” The moment we wake up to the fact that we are part of the world and engage
in a conversation with her to get to know her (and ourselves) better, the captivity of a
dualistic world view ends. We are freed to engage as participants in the world (Holdrege
2005).
The consideration of different scenarios for hypothesis testing is one practical

solution to avoid the ad-hoc explanations that relegate biogeography as a mere
descriptive discipline, usually “relegated to the interesting pay-off” in systematic
studies (Upchurch 2006, p 6). “The greatest strides we can make in unlocking the
mysteries and complexities of nature in this fundamentally interdisciplinary science
are those from new synthesis and bold collaborations among scientists across the
many descendant disciplines, long divergent but now reticulating within a strong
spatial context – the new biogeography” (Lomolino and Heaney 2004, p 1). “We
believe that the best means for advancing the frontiers of our science is to fos-
ter reintegration and reticulations among complementary research programs. The
new series of synthesis –more complex, scale-variant, and multi-factorial views of
how the natural world develops and diversifies – may be less appealing to some
researchers, but it is likely to result in a much more realistic and more illuminat-
ing view of the complexity of nature” (Lomolino and Heaney 2004, p 2). Alas, this
10.5 The “New Biogeography” 283
new biogeography will have to deal with complex science, with uncertainties, with
deconstructions, all the challenges that shape the current scientific endeavour.
The new biogeographic synthesis seems to be in tune with Goodwin’s new biol-
ogy: “The new biology is biology in the form of an exact science of complex systems
concerned with dynamics and emergent order. Then everything in biology changes.
Instead of the metaphors of conflict, competition, selfish genes, climbing peaks in
fitness landscapes, what you get is evolution as a dance. It has no goal. As Stephen
Jay Gould says, it has no purpose, no progress, no sense of direction. It’s a dance
through morphospace, the space of the forms of organisms” (Goodwin in Brockman
1995, p 97). In the same line is the critique of reductionist ideological biologies
and the search for alternative models that respects the complexity and freedom of
organisms (Rose 1997). This has certainly to do with the still insufficiently known
principles of the auto-organisation of life on the base of the proposals of Varela
et al. (1974), and the recent increasing developments in evolutionary biology (evo-
devo) (García Azkonobieta 2005). This development is already affecting systematic
problems in plant biology (e.g. Frohlich 2006), and traditionally “hard” fields like
palaeobotany are beginning a concerted effort with developmental biologists to
reevaluate fossils (Sanders et al. 2007; Miller 2009 onwards). Specific (and old)
questions like the origin of the flower (see Box 2.4), are being approached integrat-
ing molecular phylogenetic and palaeobotanical evidence (Doyle 2008), together
with early environmental constraints (Feild and Arens 2005).
According to Goodwin (2007), newer developments show signs of a return to
the idealistic integral science proposed by Alexander von Humboldt (Ette 2007;
Greppi 2008), inspired by a Goethean approach to science. Goethean science is
only recently getting recognition (Bortoft 1996; Ebach 2005; Goodwin 2007). In
contrast to the dominating mechanistic doctrine, Goethe maintained that the scientist
is not a passive observer of an external universe. Rather, the scientist is involved in
a reciprocal, participatory relationship with nature. Goethean science is a science
of relationship, of quality and of wholeness. “And we can use this kind of science
to ask questions about all forms and functions of life” (Myers 1997), as a more
qualitative approach which intends to encompass the complexity of biogeographic
systems and geosystems (Spedding 2003).
Lomolino and Heaney (2004) continue with optimism: “The revitalization will
continue in earnest, largely through the efforts of broad-thinking scientists who no
longer shy away from but embrace the complexity of nature, and who foster col-
laborations and conceptual reticulations in modern biogeography” (Lomolino and
Heaney 2004, p 2). “We are equally confident that the current and future genera-
tions of biogeographers will continue to advance its frontiers, developing a more
comprehensive understanding of, and more successful strategies for conserving, the
geography of nature” (Lomolino and Heaney 2004, p 3).
The practical applications of these novel methodological and theoretical develop-
ments in comparative and evolutionary biogeography are a powerful and promising
framework for a potentially rapid and coherent development of Chilean plant
geography in the near future.
10.6 Coda: The Geographical Nature of Biogeography
Between geographers and historical biogeographers there has been relatively little com-
munication and, like Stoddart [1985], I consider this a problem. By far the majority of
those publishing work with biogeographical themes are biologists. Most biology-trained
biogeographers appear to have little or no familiarity with the theoretical, philosophical,
and methodological literature of geography; this, at least, seems to be the only conclusion
that can be drawn from the almost total absence of referral to such in their papers (Smith
1989).
Modern geographic science has intrinsically a comparative nature, since the

early theoretical and empirical advances by Bernhardus Varenius, Ferdinand von
Richthofen and especially Alfred Hettner (Holt-Jensen 1999). Two approaches per-
meate the essence of geography since its very beginning as a modern science:
“chorography” (detailed description of a place), and “chorology” (interrelations
among places). “The goal of the chorological point of view is to know the char-
acter of regions and places through comprehension of the existence together and
interrelations among the different realms of reality” (Hettner 1927, as quoted by
Hartshorne 1960, p 13) (see Chap. 4). “It was reserved for our time, to see the com-
parative geography cultivated in masterly fashion, in its widest compass, indeed
in its reflex on human history, on the relations of the form of the earth to the direc-
tion of the characteristics of peoples and the progress of civilization” (Humboldt in
the first volume of the Kosmos [1845], as quoted by Hartshorne 1939, p 230 [54]).
Humboldt was referring to Carl Ritter’s work Allgemeine Vergleichende Geographie
which “has shown that the comparative geography attains thoroughness only when
the whole mass of facts that have been gathered from various zones, is compre-
hended in one view, is placed at the disposal of the integrating (combinierenden)
intelligence” (as quoted by Hartshorne 1939, p 230 [54]).
Current comparative biogeography (cfr Parenti and Ebach 2009) is to some
extent inherited from these early developments in geography, in spite of chorology
following its own line of development (see Williams 2007). “We aim to establish
a comparative biogeography, a method or approach that incorporates systematic
biogeography (biotic relationships and their classification and distribution) and
evolutionary biogeography (proposal of possible mechanisms responsible for distri-
butions). We focus first on description and application of a systematic biogeography,
to demonstrate biotic area homology, and then explore mechanisms or processes that
may have given rise to general patterns” (Parenti and Ebach 2009, p 47).
It seems to be that, as expected by Lomolino and Heaney (2004), the “great
synthesis” will come from the “encouraging creative development and applications
of the comparative approach. . .” (Lomolino and Heaney 2004, p 2).
In this sense, challenges to biogeography are the same as for geography in
its intent to apprehend the complexity of geosystems. In spite of its supposed
holistic nature, geography is not exempted of a tendency to specialization and
reductionism, and the field is on a search towards a modern applied systems the-
ory (Egner 2006). “General system science’s concern with historic processes and
the dynamic self-determined relationships between systems structure, functioning
and selfcreation may be more appropriate to geographical research” (Haigh 1985).
10.6 Coda: The Geographical Nature of Biogeography 285
The new discourse of science policy called “mode 2” (Gibbons et al.1994), “insists
that transdisciplinarity and engagement with end users and contexts of application
are necessary both to meeting major societal challenges, such as sustainability and
wealth creation, and to ensuring the public relevance (and funding) of science”
(Demeritt 2009).
Turning to the beginning of this last chapter, the (post)modern map of
(bio)geography shows enormous challenges but at the same time tremendous oppor-
tunities. The capacity and inspiration of researchers in geography and biology
will hopefully open new horizons and help reconstruct the “fragmented map” of
biogeography (Box 10.3).
Box 10.3 Willi Hennig’s “Criterion of Veracity”

Willi Hennig, the great theorist behind the current cladistics paradigm,
explained the “criterion of veracity” of his phylogenetic approach, with the
reconstruction of a fragmented map:
Suppose a geographer has obtained fragments of a topographic map of an unknown
land. He will make every effort to reconstruct the map from the fragments. How can
he succeed if the original map is unknown to him? He was not present when the
map was torn up. The geographer must try to assign each fragment to its original
place in the total of all the recovered fragments. He will proceed by trying to find,
for a portion of a river present on one fragment of the map, the adjoining piece of
the same river on another fragment. If he directs his attention to a single geographic
element in his map fragments, such as rivers, he is likely to go wrong. Thus the
three sections of a river, a, a′ , and a′′ (Fig. 10.6a) could seem to be upper, middle,
and lower parts of the same river. His error becomes obvious if he considers other
elements (“characters”) of his map fragments. They remain isolated; pieces of roads
and railroad lines do not join up (Fig. 10.6a). But if all geographic elements are
satisfactorily fitted together (Fig. 10.6b) the geographer will be convinced that the
fragments have been assembled correctly, even though he did not know the original
condition of the map (Hennig 1966, pp 130–131).
Fig. 10.6 Reconstruction of a fragmented map: a one possible but not correct solution;
b voilá! the correct solution (from Hennig 1966), by permission of University of Illinois
Press
All that we can do is re-collect and creatively juxtapose, experimenting with

assertions and insertions of the spatial against the prevailing grain of time. In the
end, the interpretation of postmodern geographies can be no more than a beginning
(Soja 1989, p 2).
References
Ali JR (2006) Biogeographical and geological evidence for a smaller, completely-enclosed Pacific
Basin in the Late Cretaceous: a comment. J Biogeogr 33:1670–1674
Attwell CAM, Cotterill FPD (2000) Postmodernism and African conservation science. Biodivers
Conserv 9:559–577
Barkera PF, Thomas E (2004) Origin, signature and palaeoclimatic influence of the Antarctic
Circumpolar Current. Earth Sci Rev 66:143–162
Bortoft H (1996) The wholeness of nature: Goethe’s way of science. Floris Books, Edinburgh
Boyer SL, Giribet G (2009) Welcome back New Zealand: regional biogeography and Gondwanan
origin of three endemic genera of mite harvestmen (Arachnida, Opiliones, Cyphophthalmi).
J Biogeogr 36:1084–1099
Briggs JC (2004) The ultimate expanding earth hypothesis. J Biogeogr 31:855–857
Brockman J (1995) The third culture: beyond the scientific revolution. Simon and Schuster, New
York, NY
Carey SW (1988) Theories of the earth and the universe: a history of dogma in the earth science.
Stanford University Press, Stanford, CA
Cooper A, Cooper RA (1995) The Oligocene bottleneck and New Zealand biota: genetic record of
a past environmental crisis. Proc R Soc Lond B 261:293–302
Cox CB (1990) New geological theories and old biogeographical problems. J Biogeogr 17:
117–130
Craw R (1988) Continuing the synthesis between panbiogeography, phylogenetic systematics and
geology as illustrated by empirical studies on the biogeography of New Zealand and the
Chatham Islands. Syst Zool 37:291–310
Crisci JV, Katinas L (2009) Darwin, historical biogeography, and the importance of overcoming
binary opposites. J Biogeogr 36:1027–1032
Crisci JV, Cigliano MM, Morrone JJ, Roig-Juñent S (1991) Historical biogeography of southern
South America. Syst Zool 40:152–171
Crisci JV, Katinas L, Posadas P (2003) Historical biogeography: an introduction. Harvard
University Press, Cambridge, MA
Crisci JV, Sala OE, Katinas L, Posadas P (2006) Bridging historical and ecological approaches in
biogeography. Aust Syst Bot 19:1–10
Croizat L (1952) Manual of phytogeography, or an account of plant dispersal throughout the world.
Junk, The Hague
Croizat L (1958) Panbiogeography or an introductory synthesis of zoogeography, phytogeogra-
phy, and geology; with notes on evolution, systematics, ecology, anthropology, etc. (3 vol.)
Published by the author, Caracas, Venezuela
Davis CC, Bell CD, Mathews S, Donoghue MJ (2002) Laurasian migration explains Gondwanan
disjunctions: evidence from Malpighiaceae. Proc Natl Acad Sci USA 99:6833–6837
Dawkins R (1998) Postmodernism disrobed. Nature 394:141–143
Dear M, Wassmansdorf G (1993) Postmodern consequences. Geogr Rev 83:321–325
Demeritt D (2009) Geography and the promise of integrative environmental research. Geoforum
40:127–129
Dickins JM, Choi DR, Yeates AT (1992) Past distribution of oceans and continents. In: Chatterjee
S, Hotton N III, (eds) New concepts in global tectonics. Texas Tech. University Press, Lubbock,
TX, pp 193–199
References 287
Dobson JE (1992) Spatial logic in paleogeography and the explanation of continental drift. Ann
Assoc Am Geogr 82:187–206
Donoghue MJ, Moore BR (2003) Toward an integrative historical biogeography. Integr Comp Biol
43:261–270
Doyle JA (2008) Integrating molecular phylogenetic and paleobotanical evidence on origin of the
flower. Int J Plant Sci 169:816–843
Ebach MC (2005) Anschauung and the archetype: the role of Goethe’s delicate empiricism in
comparative biology. Janus Head 8:254–270
Ebach MC, Humphries CJ (2003) Ontology of biogeography. J Biogeogr 30:959–962
Ebach MC, Humphries CJ, Williams DM (2003) Phylogenetic biogeography deconstructed.
J Biogeogr 30:1285–1296
Ebach MC, Tangney RS (2007) Biogeography in a changing world. Systematics Association
Special Volumes. CRC Press, Boca Raton, FL
Edgecombe GD, Giribet G (2008) A New Zealand species of the trans-Tasman centipede order
Craterostigmomorpha (Arthropoda: Chilopoda) corroborated by molecular evidence. Invertebr
Syst 22:1–15
Egner H (2006) Autopoiesis, form and observation – modern systems theory and its potential
contribution to the integration of human and physical geography. Mitt der Österr Geogr Gesell
148:92–108
Ette O (2007) Towards world science? Humboldtian science, world concepts, and TransArea stud-
ies. In: Rehrmann N, Ramírez Sáinz L, (eds) Dos culturas en diálogo. Historia cultural de la
naturaleza, la técnica y las ciencias naturales en España y América Latina. Madrid – Frankfurt
am Main, Iberoamericana - Vervuert, pp. 121–150
Ezcurra C, Baccala N, Wardle P (2008) Floristic relationships among vegetation types of New
Zealand and the southern Andes: Similarities and biogeographic implications. Ann Bot
101:1401–1412
Feild TS, Arens NC (2005) Form, function and environments of the early angiosperms: merging
extant phylogeny and ecophysiology with fossils. New Phytol 166:383–408
Fernández-Armesto F (1997) Truth a history. Bantam Press, London
Ferré F (1997) The postmodern world. In: Meffe GK, Carroll CR, (eds) Principles of conservation
biology, 2nd edn. Sinauer Associates, Sunderland, MA, pp 532–53
Frohlich MW (2006) Recommendations and goals for evo-devo research: scenarios, genetic
constraint, and developmental homeostasis. Aliso 22:172–187
García Azkonobieta T (2005) Evolución, desarrollo y (auto)organización. Un estudio sobre los
principios filosóficos de la evo-devo. Tesis doctoral, Universidad del País Vasco
Gibbons M, Limoges C, Nowotny H, Schwartzman S, Scott P, Trow M (1994) The new production
of knowledge: the dynamics of science and research in contemporary societies. Sage, London
Gibbs G (2006) Ghosts of Gondwana: the history of life in New Zealand. Craig Potton Publishing,
Nelson
Goldberg J, Trewick SA, Paterson AM (2008) Evolution of New Zealand’s terrestrial fauna: a
review of molecular evidence. Philos Trans R Soc B Biol Sci 363:3319–3334
Goodwin B (2007) Nature’s due: healing our fragmented culture. Floris Books, Edinburgh
Grande L (1994) Repeating patterns in nature, predictability, and “impact” in science. In: Grande L,
Rieppel O, (eds) Interpreting the hierarchy of nature: from systematic patterns to evolutionary
process theories. Academic Press, Inc, New York, NY, pp 61–84
Greppi C (2008) A proposito di evoluzionismo e geografia. L’incontro mancato fra Humboldt e
Darwin. Quad Stor 127:1–21
Griffith MP (2004) Early cactus evolution: the postmodern view. Haseltonia 10:3–11
Haigh MJ (1985) Geography and general system theory, philosophical homologies and current
practice. Geoforum 16:191–203
Hall R (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific:
computer-based reconstructions, model and animations. J Asian Earth Sci 20:353–431
Hartshorne R (1939) The classical period: Humboldt and Ritter. Ann Assoc Am Geogr 29:48–84
Hartshorne R (1960) Perspective on the nature of geography. John Murray, London

Hay WW, De Conto RM, Wold CN, Wilson KM, Voigt S, Schulz M et al. (1999) Alternative
global Cretaceous paleogeography. In: Barrera E, Johnson C (eds) Evolution of the Cretaceous
Ocean-Climate System. GSA Special Paper 332:1–47
Heads MJ (1990) Mesozoic tectonics and the deconstruction of biogeography: a new model of
Australasian biology. J Biogeogr 17:223–225
the genus Abrotanella (Compositae). Biol J Linn Soc 67:391–432
Heads M (2002) Birds of paradise, vicariance biogeography and terrane tectonics in New Guinea.
Heads M (2005a) The history and philosophy of panbiogeography. In: Llorente Bousquets J,
Morrone JJ, (eds) Regionalización biogeográfica en Iberoamérica y tópicos afines. Las Prensas
de Ciencias, Facultad de Ciencias, UNAM, México DF, pp 67–123
Heads M (2005b) Dating nodes on molecular phylogenies: a critique of molecular biogeography.
Cladistics 21:62 –78
Heads M (2009) Globally basal centres of endemism: the Tasman-Coral Sea region (south-west
Hennig W (1966) Phylogenetic systematics. University of Illinois Press, Chicago, IL
Hettner A (1927) Die Geographie, ihre Geschichte, ihr Wesen und ihre Methoden. Ferdinand Hirt,
Breslau
Hilgenberg OC (1933) Vom wachsenden Erdball. Published by the author, Berlin
Holdrege C (2005) Doing Goethean science. Janus Head 8:27–52
Holt-Jensen A (1999) Geography history & concepts. A student’s Guide, 3rd edn. Sage, London
Jones O (2008) Stepping from the wreckage: geography, pragmatism and anti-representational
theory. Geoforum 39:1600–1612
Jones MEH, Tennyson AJD, Worthy JP, Evans SE, Worthy TH (2009) A sphenodontine
(Rhynchocephalia) from the Miocene of New Zealand and palaeobiogeography of the tuatara
(Sphenodon). Proc R Soc B 276:1385–1390
Jordan GJ, Bannister JM, Mildenhall DC, Zetter R, Lee DE (2010) Fossil Ericaceae from New
Zealand: deconstructing the use of fossil evidence in historical biogeography. Am J Bot 97:
59–70
Kamp PJJ (1980) Pacifica and New Zealand: proposed eastern elements in Gondwanaland’s
history. Nature 288:659–664
Katinas L, Morrone JJ, Crisci JV (1999) Track analysis reveals the composite nature of the Andean
biota. Aust J Bot 47:111–130
Knapp M, Mudaliar R, Havell D, Wagstaff SJ, Lockhart PJ (2007) The drowning of New Zealand
and the problem of Agathis. Syst Biol 56:862–870
Krause DW, O’Connor PM, Rogers KC, Sampson SD, Buckley GA, Rogers RR (2006) Late
Cretaceous terrestrial vertebrates from Madagascar: implications for Latin American biogeog-
raphy. Ann Mo Bot Gard 93:178–208
Larson RL, Pockalny RA, Viso RF, Erba E, Abrams LJ, Luyendyk BP, Stock JM, Clayton RW
(2002) Mid-Cretaceous tectonic evolution of the Tongareva triple junction in the southwestern
Pacific Basin. Geology 30:67–70
Lee DE, Bannister JM, Lindqvist JK (2007) Late Oligocene-Early Miocene leaf macrofossils
confirm a long history of Agathis in New Zealand. N Z J Bot 45:565–578
Lindemann B (1927) Kettengebirge, kontinentale Zerspaltung und Erdexpansion. Gustav Fischer,
Jena
Linder HP, Crisp MD (1995) Nothofagus and Pacific biogeography. Cladistics 11:5–32
Lomolino MV, Heaney LR (2004) Reticulations and reintegration of modern biogeography. In:
Lomolino MV, Heaney LR, (eds) Frontiers in biogeography: new directions in the geography
of nature. Sinauer, Sunderland, MA, pp 1–3
Marquet PA, Fernández M, Navarrete SA, Valdovinos C (2004) Diversity emerging: toward
a deconstruction of biodiversity patterns. In: Lomolino MV, Heaney LR, (eds) Frontiers
References 289
in biogeography: new directions in the geography of nature. Sinauer, Sunderland, MA, pp

191–209
McCarthy D (2003) The trans-Pacific zipper effect: disjunct sister taxa and matching geological
outlines that link the Pacific margins. J Biogeogr 30:1545–1561
McCarthy D (2005a) Biogeographical and geological evidence for a smaller, completely-enclosed
Pacific Basin in the Late Cretaceous. J Biogeogr 32:2161–2177
McCarthy D (2005b) Biogeography and scientific revolutions. Systematist 25:3–12
McCarthy D (2007) Are plate tectonic explanations for trans-pacific disjunctions plausible?
Empirical tests of radical dispersalist theories. In: Ebach MC, Tangney RS, (eds) Biogeography
in a changing world. Systematics Association special volumes. CRC Press, Boca Raton, FL, pp
171–192
McDowall RM (2008) Process and pattern in the biogeography of New Zealand – a global
microcosm? J Biogeogr 35:197–212
McGlone MS (2005) Goodbye Gondwana. J Biogeogr 32:739–740
McLoughlin S (2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic
provincialism. Aust J Bot 49:271–300
Michaux B, Leschen RAB (2005) East meets west: biogeology of the Campbell Plateau. Biol J
Linn Soc 86:95–115
Miller JM (2009 onwards) Paleobotany of Angiosperm Origin. http://www.gigantopteroid.org/
html/research.htm
Milne RI (2006) Northern Hemisphere plant disjunctions: a window on tertiary land bridges and
climate change? Ann Bot 98:465–472
Morley RJ (2001) Why are there so many primitive angiosperms in the rain forests of Asia-
Australasia? In: Metcalfe I, Smith JMB, et al.(eds) Faunal and floral migrations and evolution
in SE Asia-Australasia. Balkema Publishers, Tokyo, pp 185–199
Morley RJ (2003) Interplate dispersal paths for megathermal angiosperms. Perspect Plant Ecol
Evol Syst 6:5–20
Morrone JJ (2003) ¿Quién le teme al Darwinismo? Ciencia 54:78–88
Morrone JJ (2007) Hacia una biogeografía evolutiva. Rev Chil Hist Nat 80:509–520
Morrone JJ (2009) Evolutionary biogeography: an integrative approach with case studies.
Columbia University Press, New York, NY
Morrone JJ, Crisci JV (1995) Historical biogeography: an introduction to methods. Annu Rev Ecol
Syst 26:373–401
Myers N (1997) Exploring Goethean Science at Schumacher College. http://www.island.org/
prescience/goethe.html
Myers AA, Giller PS, (eds) (1988) Analytical biogeography: an integrated approach to the study
of animal and plant distributions. Chapman and Hall, London
Nelson G (1975) Biogeography, the vicariance paradigm, and continental drift. Syst Zool 24:
490–504
Nelson G, Ladiges PY (2001) Gondwana, vicariance biogeography and the New York School
revisited. Aust J Bot 49:389–409
Nur A, Ben-Avraham Z (1977) Lost Pacifica continent. Nature 270:41–43
Nur A, Ben-Avraham Z (1981) The lost Pacifica continent: a mobilistic speculation. In: Nelson G,
Rosen DE, (eds) Vicariance biogeography: a critique. Columbia University Press, New York,
NY, pp 341–358
Parenti LR, Ebach MC (2009) Comparative biogeography: discovering and classifying
Biogeographical patterns of a dynamic earth. University of California Press, Barkley, CA
Pole M (1994) The New Zealand flora-entirely long-distance dispersal? J Biogeogr 21:625–635
Rapalini AE (2005) The accretionary history of southern South America from the latest Proterozoic
to the Late Palaeozoic: some palaeomagnetic constraints. In: Vaughan APM, Leat PT,
Pankhurst RJ, (eds) Terrane processes at the margins of Gondwana, 246. Geological Society,
London , 305–328, Special Publications
Renner SS (2005) Relaxed molecular clocks for dating historical plant dispersal events. Trends
Plant Sci 10:550–558
Riddle BR (2005) Is biogeography emerging from its identity crisis? J Biogeogr 32:185–186
Rose S (1997) Lifelines: biology beyond determinism. Oxford University Press, Oxford
Rosen BR (1988) Biogeographic patterns: a perceptual overview. In: Myers AA, Giller P, (eds)
Analytical biogeography: an integrated approach to the study of animal and plant distributions.
Chapman and Hall, London, pp 23–56
Salomon M (2001) Evolutionary biogeography and speciation: essay on a synthesis. J Biogeogr
28:13–27
Sanders H, Rothwell GW, Wyatt S (2007) Paleontological context for the developmental mecha-
nisms of evolution. Int J Plant Sci 168:719–728
Scalera G (2006) TPW and Polar Motion as due to an asymmetrical Earth expansion. In: Lavecchia
G and Scalera G (eds) Frontiers in Earth Sciences: New Ideas and Interpretations. Ann Geophys
49(Supplement):483–500
Scalera G (2007) Fossils, frogs, floating islands and expanding Earth in changing-radius car-
tography. a comment to a discussion on Journal of Biogeography. Ann Geophys 50:
789–798
Scalera G (2008) Great and old earthquakes against great and old paradigms – paradoxes, historical
roots, alternative answers. Adv Geosci 14:41–57
Scalera G, Jacob K-H, (eds) (2003) Why expanding Earth? – a book in honour of OC Hilgenberg.
INGV, Rome
Sdrolias M, Müller RD, Gaina C (2003) Tectonic evolution of the southwest Pacific using
constraints from backarc basins. In: Hillis RR, Müller RD (eds) Evolution and dynamics
of the Australian plate. Geol Soc Aust Spec Publ 22, and Geol Soc Am Spec Pap 372:
343–359
Shields O (1998) Upper Triassic Pacific vicariance as a test of geological theories. J Biogeogr
25:203–211
Sluys R (1994) Explanations for biogeographic tracks across the Pacific Ocean: a challenge for
paleogeography and historical biogeography. Prog Phys Geogr 18:42–58
Smith CH (1989) Historical biogeography: geography as evolution, evolution as geography. N Z J
Zool 16:773–785
Smith AD (2007) A plate model for Jurassic to Recent intraplate volcanism in the Pacific Ocean
basin. GSA Spec Pap 430:471–495
Soja EW (1989) Postmodern geographies: the reassertion of space in critical social theory. Verso,
London
Spedding N (2003) Landscape and environment: biophysical processes, biophysical forms. In:
Holloway SL, Rice SP, Valentine G, (eds) Key concepts in geography. SAGE, London, pp
281–303
Stevens GR (1997) The Late Jurassic ammonite fauna of New Zealand. Institute of Geological and
Nuclear Sciences monograph 18 (New Zealand Geological Survey Paleontological Bulletin
74). Institute of Geological & Nuclear Sciences, Lower Hutt
Stöckler K, Daniel IL, Lockhart PJ (2002) New Zealand Kauri (Agathis australis (D. don) Lindl.,
Araucariaceae) survives Oligocene drowning. Syst Biol 51:827–832
Terribile LC, Felizola Diniz-Filho JA, Rodríguez MA, Rangel TFLVB (2009) Richness patterns,
species distributions and the principle of extreme deconstruction. Glob Ecol Biogeogr 18:
123–136
Turrill WB (1953) Pioneer plan geography: the phytogeographical researches of Sir Joseph Dalton
Hooker. Martinus Nijhoff, The Hague
Udvardy MDF (1981) The riddle of dispersal: dispersal theories and how they affect vicariance
biogeography. In: Nelson G, Rosen DE, (eds) Vicariance biogeography: a critique. Columbia
University Press, New York, NY, pp 6–29
Upchurch P (2006) What’s wrong with palaeobiogeography? In: Meeting Abstracts: palaeogeog-
raphy and palaeobiogeography: biodiversity in space and time, 10–13 Apr 2006, University of
Cambridge, pp 5–6
Upchurch P (2008) Gondwanan break-up: legacies of a lost world? Trends Ecol Evol 23:229–236
References 291
Upchurch P, Hunn CA, Norman DB (2002) An analysis of dinosaurian biogeography: evidence

for the existence of vicariance and dispersal patterns caused by geological events. Proc R Soc
Lond B 269:613–621
van Steenis CGGJ (1962) The theory of land bridges in botany, with particular references to tropical
plants. Blumea 11:235–542
Varela FJ, Maturana HR, Uribe R (1974) Autopoiesis: the organization of living systems, its
characterization and a model. Biosystems 5:187–196
Vaughan APM, Livermore RA (2005) Episodicity of Mesozoic terrane accretion along the Pacific
margin of Gondwana: implications for superplume-plate interactions. Geol Soc Lond Spec Publ
246:143–178
Vogel K (1990) The expansion of the earth – an alternative model to plate tectonics theory. In:
Belousov V, (ed) Critical aspects of the plate tectonics theory, vol II. Athens, pp 19–34
von Ihering H (1907) Archhelenis und Archinotis: Gesammelte Beiträge zur Geschichte der
Neotropischen Region. W. Engelmann, Leipzig
Wanntorp L, Wanntorp H-E (2003) The biogeography of Gunnera L.: vicariance and dispersal.
Waters JM, Craw D (2006) Goodbye Gondwana? New Zealand biogeography, geology, and the
problem of circularity. Syst Biol 55:351–356
Wegener A (1915) Die Entstehung der Kontinente und Ozeane. Sammlung Vieweg Nr. 23,
Braunschweig
Williams DM (2007) Ernst Haeckel and Louis Agassiz: trees that bite and their geographical
dimension. In: Ebach MC, Tangney RS, (eds) Biogeography in a changing world. Systematics
association special volumes. CRC Press, Boca Raton, FL, pp 1–59
Williams DM, Ebach MC (2008) Foundations of systematics and biogeography. Springer, New
York, NY
Wilson GDF (2008) Gondwanan groundwater: subterranean connections of Australian phreatoi-
cidean isopods (Crustacea) to India and New Zealand. Invertebr Syst 22:301–310
Winkworth RC, Wagstaff S, Glenny D, Lockhart PJ (2002) Plant dispersal N.E.W.S. from New
Zealand. Trends Ecol Evol 17:514–520
Zhou ZK, Yang XF, Yang QS (2006) Land bridge and long-distance dispersal – old views, new
evidence. Chin Sci Bull 51:1030–1038
Chapter 11
Epilogue: The Juan Fernández Islands
and the Long-Distance Dispersal of Utopia
As a kind of “path biographies” (cfr. Daniels and Nash 2004), ideas disperse over
the globe linking concepts that seemed separate in space and time, promoting com-
mon pulses of human development. The Life and Strange Adventures of Robinson
Crusoe (1719) by Daniel Defoe was a milestone in the construction of the industrial
society (Laborde 2007). The ideal of humanity dominating nature, and finally the
possibility of a real cohabitation between both, was the central idea of Defoe’s novel
and inspired the nineteenth century ideals of modernity, including Rousseau’s edu-
cational ideal. “Robinson Crusoe became a primer for practical education. It was the
only book Rousseau would allow the child in his educational tract Emile. [“I want
him to learn in detail not from books but from things, all that must be known in such
a situation; I want him to think he is Robinson himself”]” (Rousseau as quoted by
Daniels and Nash 2004).
But nature remains us systematically that we are still far from dominating her.
Also in Robinson Crusoe’s land: the tsunami on the 27th of February 2010 erased
the town of San Juan Bautista, leaving only agony and despair. The town is being
reconstructed, together with the hope. This is an opportunity to rethink a new rela-
tionship with the unique biota that occupies the archipelago since millions of years.
This would be a natural laboratory to rebuild and heal the relationship between
nature and culture (Goodwin 2007). Since the school building does no longer exist,
this is the moment to turn our eyes towards the montane forest for the cultivation of
postmodern Robinsons as the “last children in the woods” (cfr. Louv 2008). Children
know it better: they easily and spontaneously appreciate the wonders of nature.
As we age, childhood becomes another country, a disputed territory of memory and mean-
ing. Its true geography is quickly forgotten, giving away to an adult-imagined universo
(Jones 2008).
There shall be somewhere still a place for Utopia. Intrinsic to humanity is

the constant search for new worlds; let’s revisit this time ourselves, our origin
and dispersal upon this American and insular territory called Chile. The Juan
Fernández Islands could be a good place for starting re-building our home for the
future.

DOI 10.1007/978-90-481-8748-5_11, ⃝C Springer Science+Business Media B.V. 2011
294 11 Epilogue: The Juan Fernández Islands and the Long-Distance Dispersal of Utopia
We commonly speak about the future, as if it is just one, which is obviously a mistake.
Nothing can be foretold about the future or about the many futures whose seeds await
within the womb of time, except that they will not resemble the present (J.L. Borges, my
translation).
References
Daniels S, Nash C (2004) Lifepaths: geography and biography. J Hist Geogr 30:449–458
Goodwin B (2007) Nature’s due: healing our fragmented culture. Floris Books, Edinburgh
Jones O (2008) True geography [] quickly forgotten, giving away to an adult-imagined universe.
Approaching the otherness of childhood. Child Geogr 6:195–212
Laborde M (2007) Daniel Defoe y Jean Jacques Rousseau: con los Ojos en el Archipiélago de Juan
Fernández. Rev Unive (UC) 96:76–77
Louv R (2008) Last child in the woods: saving our children from nature-deficit disorder. Algonquin
Books of Chapel Hill, Chapel Hill, NC
Appendix A
Chilean native genera, ordered by floristic elements: (1) pantropical, (2) australasiatic, (3) neotropical, (4) antitropical, (5) aouth-temperate, (6) endemic, (7)
cosmopolitan
Native Endemic
Genus Family Order Global distribution Element Track spp. spp.
Abutilon Malvaceae Malvales Pantrops 1 1 0

Acacia Fabaceae Fabales Tropics, Subtrop, esp Afr y Aus 1 3 0
Achyrocline Asteraceae Asterales Tropics Afr, Madagascar, C Am, S Am 1 1 0
Alternanthera Amaranthaceae Caryophyllales Tropics & subtrop, S Am, N Am, Aus, 1 sp 1 2 0
DOI 10.1007/978-90-481-8748-5, ⃝
Caucasus
Aristolochia Aristolochiaceae Piperales Trop, subtrop, some in temp regions 1 2 2
Axonopus Poaceae Poales Pantrops, IP 1 1 1
Bacopa Veronicaceae Lamiales Trop & Subtrop, Hawaii, Madag, Aus 1 1 0
Beilschmiedia Lauraceae Laurales Pantropics, Mex to S Bras, Chile, Aus, 1 2 2
NZ, Afr, Madag
Boehmeria Urticaceae Rosales Pantrop, JF 1 1 1
Buddleja Scrophulariaceae Lamiales E Asia, tro & subtrop AM, Afr 1 3 1
Bulbostylis Cyperaceae Poales Tropics & subtropics 1 1 0
Caesalpinia Fabaceae Fabales Pantropics, subtropics, Arg, Madag, S Afr, 1 4 2
Arabia
Calliandra Fabaceae Fabales C & S Am, Afr, Madag, trop Asia 1 1 1
A. Moreira-Muñoz, Plant Geography of Chile, Plant and Vegetation 5,

Carpha Cyperaceae Poales Trop & S Afr, Madag, japan, New Guinea, 1 1 0
Aus, Tasm, NZ, temp S Am
Centella Apiaceae Apiales Pantrop to Chile, NZ, JF 1 1 0
Centipeda Asteraceae Asterales Aus, NZ, New Cale, Tahiti, Chile, Madag, 1 1 0

Mascarene Is, SE Asia
Cissus Vitaceae Vitales Tropics incl Australia 1 1 0
Cleome Cleomaceae Brassicales Tropics & subtropics, spec. Am 1 1 0
Conyza Asteraceae Asterales Tropics & subtropics 1 20 15
Cordia Boraginaceae Unplaced Trop & subtropics 1 1 1
295
296
Native Endemic
Cotula Asteraceae Asterales S Afr, NZ, New Guinea, Aus, S Am 1 1 0

Cressa Convolvulaceae Solanales Trop & subtrop 1 1 0
Croton Euphorbiaceae Malpighiales Tropics & subtropics 1 1 1
Cryptocarya Lauraceae Laurales Tropics & subtropics, Madag, S Afr, E Asia, 1 1 1
N Australia
Cynodon Poaceae Poales Tropics & subtropics 1 2 1
Davallia Davalliaceae Polypodiales W Medit, Himal, Japan, Aus, Tahiti, Afr, 1 1 0
Madag, IP
Dennstaedtia Dennstaedtiaceae Polypodiales Trop to warm temp, Madag 1 1 0
Dichondra Convolvulaceae Solanales Tropics & subtropics, JF 1 2 0
Dicliptera Acanthaceae Lamiales Tropics & subtropics 1 1 1
Dioscorea Dioscoreaceae Dioscoreales Trop, subtrop, warm-temp regions 1 36 34
Dodonaea Sapindaceae Sapindales Tropics & subtrop, espec Australia 1 1 0
Drymaria Caryophyllaceae Caryophyllales Pantrop 1 3 1
Elaphoglossum Dryopteridaceae Polypodiales Pantrops & subtrop, Madag, IP 1 6 2
Evolvulus Convolvulaceae Solanales Trops & subtrops 1 1 0
Glinus Molluginaceae Caryophyllales Pantrop 1 1 0
Gomphrena Amaranthaceae Caryophyllales Trop & subtrop Am, Afr, Indochina, Aus 1 2 0
Grammitis Polypodiaceae Polypodiales New Guinea, Asia, Am, Aus, Madag, 1 3 0
warm & temp, JF
Gunnera Gunneraceae Gunnerales Am from Mexic to Chile, trop & S Afr, 1 11 6
Madag, Malesia, Tasm, NZ, Hawaii, JF
Habenaria Orchidaceae Asparagales Pantrop & subtrop 1 1 0
Histiopteris Dennstaedtiaceae Polypodiales Pantrop, Madag 1 1 0
Hybanthus Violaceae Malpighiales Trops & subtrops 1 1 0
Hypolepis Dennstaedtiaceae Polypodiales Trop & subtrop, Madag 1 1 0
Ipomoea Convolvulaceae Solanales Pantrop 1 1 0
Kyllinga Cyperaceae Poales Pantrops 1 1 0
Appendix A
Native Endemic
Appendix A
Leptochloa Poaceae Poales Tropics & subtrop 1 1 0

Lippia Verbenaceae Lamiales Am trop, many in Afr trop 1 2 1
Lobelia Campanulaceae Asterales Trops & subtrops 1 5 4
Machaerina Cyperaceae Poales Trops esp Australia, Hawaii, JF 1 1 1
Maytenus Celastraceae Celastrales Trops & subtrops 1 4 0
Megalastrum Dryopteridaceae Polypodiales Trop Am, 1 Afr to Mascarenes 1 2 1
(no Madag), JF
Microlepia Dennstaedtiaceae Polypodiales Paleotrop, 1 Afr, Madag, Japan, NZ, IP 1 1 0
Microsorum Polypodiaceae Polypodiales Trop & warm, Madag, IP 1 1 0
Mikania Asteraceae Asterales Pantrops, esp Neotrops 1 2 1
Morella Myricaceae Fagales Sudam Andes Peru Ecuador, Africa, E Asia, 1 1 0
Filipinas y Malasia
Nicotiana Solanaceae Solanales Am, USA to Chile, Aust, New Caledonia, 1 14 6
Namibia, JF
Notholaena Pteridaceae Polypodiales Warm to trop Am, SW US, Mex, Madag, JF 1 1 0
Oldenlandia Rubiaceae Gentianales Tropics espec Asia & Malesia, JF 1 2 1
Otholobium Fabaceae Fabales E & S Afr, temp S Am 1 2 1
Passiflora Passifloraceae Malpighiales Trop & subtrop Am, 1 Madag, 1 1 0
20 trop E Asia, Malesia, 3 Aus, NZ
Pellaea Pteridaceae Polypodiales Trop & warm-temp, Madag 1 2 1
Pelletiera Primulaceae Ericales 1 subtrop S Am, 1 Macaronesia 1 1 0
Peperomia Piperaceae Piperales Trop & warm esp Am, JF, Hawaii 1 7 7
Phytolacca Phytolaccaceae Caryophyllales Trop & warm 1 2 1
Pilea Urticaceae Rosales Trop & warm excl Australasia 1 3 2
Pilostyles Rafflesiaceae Malpighiales Am US to Magallanes, Asia, 2 W Aus 1 1 0
Pityrogramma Pteridaceae Polypodiales Am trop desde S US, Afr, Madag 1 1 0
Pleopeltis Polypodiaceae Polypodiales Trop Am, 1 Afr-Madag, India, Sri Lanka 1 2 0
Pluchea Asteraceae Asterales Pantrops 1 1 0
297
298
Native Endemic
Podocarpus Podocarpaceae Pinales Afr, Madag, Asia, Aus, Tasm, NZ, Mex, 1 2 1
C & S Am, WI
Pouteria Sapotaceae Ericales Trop. Am, Afr, Asia, Aus, NZ, New Caled 1 1 1
Prosopis Fabaceae Fabales Trop & subtrop Am, Afr, W Asia 1 7 1
Psilotum Psilotaceae Psilotales Pantrop, Hawaii, Australia, NZ, Madag, IP 1 1 1
Pycreus Cyperaceae Poales Pantrops 1 3 1
Schizaea Schizaeceae Schizaeales Trop & austral, N Am, Madag 1 1 0
Senna Fabaceae Fabales Pantrops 1 10 7
Sigesbeckia Asteraceae Asterales Pantrops 1 1 0
Spilanthes Asteraceae Asterales Pantrops 1 1 0
Sticherus Gleicheniaceae Gleicheniales Pantrop, Madag, JF 1 5 2
Tecoma Bignoniaceae Lamiales Am tropical Arizona to Chile, 1 S Afr, Aus? 1 1 0
Thelypteris Thelypteridaceae Polypodiales Trops & subtrops, Madag 1 2 0
Trichomanes Hymenophyllaceae Hymenophyllales Patrop, Madag, JF 1 3 3
Trichoneura Poaceae Poales Arabia, trop Afr, S US, Peru 1 1 0
Tripogon Poaceae Poales Trops & subtrops 1 2 0
Triumfetta Malvaceae Malvales Pantrop 1 1 0
Vittaria Pteridaceae Polypodiales Pantrop, IP 1 1 0
Wedelia Asteraceae Asterales Pantrops, W Australia 1 1 0
Wolffiella Araceae Alismatales Trop & warm Am, 1 S Afr 1 1 0
Fagara Rutaceae Sapindales Am, Afr, Asia, JF 1 2 2
Alonsoa Scrophulariaceae Lamiales Trop Am Mexico, C Am, Chile, Bol, 1 2 1
2 sp S Afr
Amphibromus Poaceae Poales Aus, NZ, S Am 2 AUS 1 0
Aristotelia Elaeocarpaceae Oxalidales E Aus, Tasman, NZ, Arg, Chile 2 AUS 1 0
Azorella Apiaceae Apiales Andes to temp S Am, Malvinas, Antarctic 2 AUS 17 2
Islands
Appendix A
Native Endemic
Appendix A
Berberidopsis Berberidopsidaceae Berberidopsidales B. beckleri en Aus: Queensland & 2 AUS 1 1

New S Wales, B. corallina: Chile
Cortaderia Poaceae Poales S Am, 4 NZ, 1 New Guinea 2 AUS 6 0
Discaria Rhamnaceae Rosales Aus, NZ, S Am 2 AUS 2 0
Donatia Stylidiaceae Asterales Tasman, NZ, subantarctic S Am 2 AUS 1 0
Eucryphia Cunoniaceae Oxalidales SE Aus, Tasm, Chile 2 AUS 2 1
Fuchsia Onagraceae Myrtales C & S Am, NZ, Tahiti 2 AUS 2 1
Griselinia Griseliniaceae Apiales NZ, Chile, Paraguay, S Brasil 2 AUS 5 3
Haloragis Haloragaceae Saxifragales Aus, N. Caled, NZ, Rapa, JF 2 AUS 2 2
Heterozostera Zosteraceae Alismatales Coastal temp Aus, Tasm, N Chile 2 AUS 1 0
Jovellana Calceolariaceae Lamiales NZ, Chile 2 AUS 3 3
Laurelia Atherospermataceae Laurales NZ & Chile 2 AUS 1 1
Lepidothamnus Podocarpaceae Pinales 2 NZ, 1 S Chile/Arg 2 AUS 1 0
Lomatia Proteaceae Proteales Aus, Tas, S Am 2 AUS 3 0
Luzuriaga Luzuriagaceae Liliales 1 NZ, 3 Chile, Arg to T Fuego 2 AUS 3 1
Maireana Amaranthaceae Caryophyllales Australia, Chile 2 AUS 1 0
Marsippospermum Juncaceae Poales NZ, S S Am, Tfuego, Falkland 2 AUS 3 0
Orites Proteaceae Proteales 6 temp E Aus, 3 Andes S Am 2 AUS 1 0
Ourisia Veronicaceae Lamiales Tasm, NZ, Andes S Am 2 AUS 11 3
Phyllachne Stylidiaceae Asterales Tasm, NZ, temp S Am 2 AUS 1 0
Prumnopitys Podocarpaceae Pinales Costa Rica to Chile, New Caled, NZ, E Aus 2 AUS 1 0
Raukaua Araliaceae Apiales Tasm, NZ, Chile 2 AUS 2 1
Rostkovia Juncaceae Poales NZ, S S Am, Tfuego, Falkland 2 AUS 1 0
Rytidosperma Poaceae Poales Australia, NZ, Arg, Chile, IP 2 AUS 6 2
Schizeilema Apiaceae Apiales 11 NZ, 1 Aus, 1 S S Am 2 AUS 1 0
Selliera Goodeniaceae Asterales Aus, NZ, Chile 2 AUS 1 0
Tetrachondra Tetrachondraceae Lamiales 1 NZ, 1 Patagonia, Tfuego 2 AUS 1 0
Trichocline Asteraceae Asterales S Am, 1 W Aus 2 AUS 5 2
299
300
Native Endemic
Ficinia Cyperaceae Poales Cape Region S Afr, 1 subantarctic circumpolar 2 C-AUS 1 0

Nertera Rubiaceae Gentianales Madag, SE Asia, Malesia, Aus, NZ, Hawaii, 2 C-AUS 1 0
S Am, JF
Rumohra Dryopteridaceae Polypodiales Circumaustral, J Fernandez, Madag 2 C-AUS 2 1
Wahlenbergia Campanulaceae Asterales Circumaustral 2 C-AUS 6 5
Abrotanella Asteraceae Asterales New Guinea, Aus, Tasmania, NZ, SAm 2 TPAC 6 3
(TFuego), JF, Falkland Is.
Apodasmia Restionaceae Poales SE Asia, Malesia, New Guinea, Aus, Tasm, 2 TPAC 1 1
NZ, Chile, Arg
Araucaria Araucariaceae Pinales New Guinea, Aus, New Caledonia, S Chile, 2 TPAC 1 0
SE Brasil
Arthropteris Tectarinaceae Polypodiales New Guinea, NZ, Juan Fernandez, Madag 2 TPAC 1 1
Astelia Asteliaceae Asparagales Masc, New Guinea, Aus, NZ, Polinesia, Chile 2 TPAC 1 0
Caldcluvia Cunoniaceae Oxalidales Philippines, Sulawesi, Moluccas, New Guinea, 2 TPAC 1 0
trp Aus, S Chile Arg
Citronella Cardiopteridaceae Aquifoliales Malesia, E Aus, New Cale, Fiji, Samoa, C & 2 TPAC 1 1
trop S Am
Colobanthus Caryophyllaceae Caryophyllales S Pacific, Aus, NZ, temp S Am, Ecuador 2 TPAC 3 0
Coprosma Rubiaceae Gentianales E Malesia, Aus, NZ, Pacific, JF, 12 sp Hawaii 2 TPAC 2 2
Dicksonia Dicksoniaceae Cyatheales Trop Am, Malesia, New Guinea, Aus, New 2 TPAC 2 2
Caled, NZ
Doodia Blechnaceae Polypodiales Australia, NZ, Pacific islands, Hawaii, N 2 TPAC 1 0
Guinea, IP
Gaimardia Centrolepidaceae Poales New Guinea, NZ, Tasmania, TFuego, Falkland 2 TPAC 1 0
Gevuina Proteaceae Proteales New Guinea, Aus, Chile, Arg 2 TPAC 1 0
Hebe Veronicaceae Lamiales New Guinea, Aus, Tasm, NZ, temp S Am, 2 TPAC 2 0
Falkland
Appendix A
Native Endemic
Appendix A
Lagenophora Asteraceae Asterales SE Asia, Malesia, Australia, Tasmania, NZ, 2 TPAC 3 0

C & S Am, JF, Falkland, Tristan da Cunha.
Leptinella Asteraceae Asterales New Guinea, Aus, NZ, Subantarctic Is, S Am 2 TPAC 1 0
Leptostigma Rubiaceae Gentianales SE Asia, NZ, W S Am 2 TPAC 1 1
Libertia Iridaceae Asparagales New Guinea, E Aus, NZ, Andes S Am, JF 2 TPAC 6 5
Muehlenbeckia Polygonaceae Caryophyllales New Guinea, Aus, Tasm, NZ, W S Am 2 TPAC 2 0
Nothofagus Nothofagaceae Fagales New Guinea, Aus, Tasm, NZ, New Caled, 2 TPAC 10 4
temp S Am
Oreobolus Cyperaceae Poales Malesia, Aus, Tasm, NZ, Hawaii, JF, trop & 2 TPAC 1 0
temp Am, Chile, Tfuego, Falkland
Oreomyrrhis Apiaceae Apiales E Asia, Borneo, N Guinea, SE Aus, NZ, 2 TPAC 1 0
Mex, C Am, Andes, Tfuego, Falkland
Santalum Santalaceae Santalales Indomalasia to Aus, Hawaii, JF (ex) 2 TPAC 1 1
Uncinia Cyperaceae Poales Malesia, Pacific, Aus, SE Asia, C & S Am, JF 2 TPAC 18 7
Acantholippia Verbenaceae Lamiales Arid S Am, Arg, Chile, Bol 3 C-AND 3 1
Anatherostipa Poaceae Poales Arg, Bol, Chile, Ecuador, Peru 3 C-AND 4 1
Andeimalva Malvaceae Malvales Chile, Peru, Bol 3 C-AND 1 1
Anthochloa Poaceae Poales Andes Chile Peru 3 C-AND 1 0
Aphyllocladus Asteraceae Asterales Andes S Bolivia, N Chile, NW Arg 3 C-AND 1 1
Arachnitis Corsiaceae Liliales Chile, Arg, Bol 3 C-AND 1 0
Argylia Bignoniaceae Lamiales S Peru, N C Chile & Arg, 3 C-AND 11 8
A. uspallatensis S Bol
Balbisia Vivianiaceae Geraniales Peru, Bol, Chile, Arg 3 C-AND 4 2
Browningia Cactaceae Caryophyllales Peru, Bol, Chile, Paraguay 3 C-AND 1 0
Chaetanthera Asteraceae Asterales S Peru, Bol, Chile, Arg 3 C-AND 38 16
Chersodoma Asteraceae Asterales S Peru, Bol, Chile, Arg 3 C-AND 3 0
Clinanthus Amaryllidaceae Asparagales Perú, Chile 3 C-AND 1 0
Corryocactus Cactaceae Caryophyllales Bol, S Peru, N Chile 3 C-AND 1 0
301
302
Native Endemic
Cristaria Malvaceae Malvales Chile, Arg, Bol, Perú, ID 3 C-AND 19 16

Cumulopuntia Cactaceae Caryophyllales Bol, S Peru, N Chile 3 C-AND 2 0
Dielsiochloa Poaceae Poales Peru, Bol, Arg, Chile 3 C-AND 1 0
Domeykoa Apiaceae Apiales Peru Chile 3 C-AND 2 2
Eccremocarpus Bignoniaceae Lamiales Peru, Chile, Arg 3 C-AND 1 0
Eremocharis Apiaceae Apiales Andes Chile Peru 3 C-AND 1 1
Eremodraba Brassicaceae Brassicales Andes Chile Peru 3 C-AND 1 0
Fabiana Solanaceae Solanales S Peru, Bol, Chile, Arg 3 C-AND 8 1
Haageocereus Cactaceae Caryophyllales Peru, Chile 3 C-AND 2 1
Haplopappus Asteraceae Asterales S Am Chile, Arg, Peru 3 C-AND 54 42
Haplorhus Anacardiaceae Sapindales Chile, Peru 3 C-AND 1 0
Helogyne Asteraceae Asterales Peru, Chile, Bol, Arg 3 C-AND 2 1
Islaya Cactaceae Caryophyllales Peru, Chile 3 C-AND 4 3
Junellia Verbenaceae Lamiales Peru, Bol, Chile, Arg 3 C-AND 24 5
Kageneckia Rosaceae Rosales Chile, Peru, Bolivia, Arg 3 C-AND 2 2
Laennecia Asteraceae Asterales Bol, Arg, Chile 3 C-AND 1 0
Lampaya Verbenaceae Lamiales Bol, Chile, Arg 3 C-AND 2 1
Lepidoceras Santalaceae Santalales S Peru, Chile 3 C-AND 1 1
Leucheria Asteraceae Asterales Peru, Bol, Arg, Chile 3 C-AND 42 16
Loasa Loasaceae Cornales Chile, Arg, 1 sp coastal Peru 3 C-AND 31 18
Lobivia Cactaceae Caryophyllales Andes Peru, Bol, Chile, Arg 3 C-AND 2 0
Lophopappus Asteraceae Asterales Peru, Bol, Chile, Arg 3 C-AND 3 1
Maihueniopsis Cactaceae Caryophyllales Andes Peru, Bol, Chile, Arg 3 C-AND 13 9
Malesherbia Malesherbiaceae Malpighiales Andes Peru, Chile, Arg 3 C-AND 16 13
Mastigostyla Iridaceae Asparagales Peru, Arg 3 C-AND 1 0
Mathewsia Brassicaceae Brassicales Peru, Chile 3 C-AND 5 5
Mniodes Asteraceae Asterales Peru, Chile 3 C-AND 2 0
Montiopsis Portulacaceae Caryophyllales Chile, Argentina, Bolivia y Perú 3 C-AND 18 5
Appendix A
Native Endemic
Appendix A
Moschopsis Calyceraceae Asterales Perú, Chile, Arg, Patagonia 3 C-AND 5 0

Mostacillastrum Brassicaceae Brassicales S Peru, Bol, Chile, Arg 3 C-AND 8 1
Nardophyllum Asteraceae Asterales Arg, Chile 3 C-AND 4 2
Nassauvia Asteraceae Asterales Bol, Arg, Chile, Falklands 3 C-AND 23 3
Neowerdermannia Cactaceae Caryophyllales Peru, Bol, Chile, Arg 3 C-AND 1 1
Neuontobothrys Brassicaceae Brassicales S Peru, Chile, Arg 3 C-AND 8 3
Nicandra Solanaceae Solanales Peru, Arg, Chile 3 C-AND 1 0
Nolana Solanaceae Solanales Peru to Chile, Galapagos 3 C-AND 44 40
Ocyroe Asteraceae Asterales Bol, Arg, Chile 3 C-AND 1 0
Oreocereus Cactaceae Caryophyllales Andes Peru, Bol, Chile, Arg 3 C-AND 4 1
Oxychloë Juncaceae Poales Andes S Am 3 C-AND 3 0
Oziroe Asparagaceae Asparagales Chile, Arg, Perú, Bol, Paraguay 3 C-AND 4 1
Pachylaena Asteraceae Asterales Andes N Chile Arg 3 C-AND 1 0
Palaua Malvaceae Malvales Coastal Peru and Chile 3 C-AND 4 2
Parastrephia Asteraceae Asterales Andes Bol, Chile, Perú, Arg 3 C-AND 3 0
Pasithea Xanthorrhoeaceae Asparagales Peru, Chile 3 C-AND 1 0
Patosia Juncaceae Poales Andes Chile Arg Bol 3 C-AND 1 0
Pennellia Brassicaceae Brassicales Andes Peru, Chile, Arg, Bol 3 C-AND 2 0
Philibertia Apocynaceae Gentianales Andes Perú, Chile, Bolivia, Argentina 3 C-AND 2 0
Phylloscirpus Cyperaceae Poales Arg, Chile, Bol 3 C-AND 3 0
Pitraea Verbenaceae Lamiales S Am, Chile, Peru, Arg 3 C-AND 1 0
Plazia Asteraceae Asterales Peru, Bol, Arg, Chile 3 C-AND 2 1
Polyachyrus Asteraceae Asterales Peru, Chile 3 C-AND 7 4
Proustia Asteraceae Asterales Bol, Chile, Arg, Peru 3 C-AND 3 2
Pycnophyllum Caryophyllaceae Caryophyllales Andes Peru, Chile, Arg, Bol 3 C-AND 5 0
Quinchamalium Santalaceae Santalales Andes Bol, Chile, Perú, Arg 3 C-AND 14 13
Salpichroa Solanaceae Solanales S Am: Ecuad, Peru, Bol, Bra, Parag, Uru, 3 C-AND 2 0
Chile, Arg
303
304
Native Endemic
Stangea Valerianaceae Dipsacales Arg,Chil,Perú 3 C-AND 1 0

Tetraglochin Rosaceae Rosales Arg, Chil, Perú 3 C-AND 3 0
Trichocereus Cactaceae Caryophyllales Chile, Arg, S Peru 3 C-AND 7 6
Tunilla Cactaceae Caryophyllales Andes Peru, Bol, Chile, Arg 3 C-AND 2 0
Urmenetea Asteraceae Asterales N Chile, NW Arg 3 C-AND 1 0
Urocarpidium Malvaceae Malvales Perú, Chile 3 C-AND 1 0
Weberbauera Brassicaceae Brassicales Andes Peru, Chile, Arg, Bol 3 C-AND 6 3
Zameioscirpus Cyperaceae Poales Arg, Chile, Bol 3 C-AND 3 0
Aa Orchidaceae Asparagales Andes Colombia to Argentina 3 W-AND 1 0
Aristeguietia Asteraceae Asterales Colombia, Ecuador, Peru, Chile 3 W-AND 1 1
Blepharocalyx Myrtaceae Myrtales Trop & subtro S Am, WI 3 W-AND 1 1
Bulnesia Zygophyllaceae Zygophyllales S Am: Venezuela Colombia disjunct to 3 W-AND 1 1
Chile-Arg, Brazil
Chuquiraga Asteraceae Asterales Andes S Am, Patagonia 3 W-AND 5 2
Cremolobus Brassicaceae Brassicales Andes Colombia to Chile 3 W-AND 1 0
Cuatrecasasiella Asteraceae Asterales Ecuador, Peru, Chile, Arg 3 W-AND 1 0
Desfontainia Desfontainiaceae Bruniales Costa Rica to Cape Horn 3 W-AND 1 0
Diplostephium Asteraceae Asterales Trop Andes from Colombia to Bol & 3 W-AND 2 0
N Chile, 1 sp Costa Rica
Distichia Juncaceae Poales Andes S Am 3 W-AND 2 0
Dunalia Solanaceae Solanales Andes Colombia to Argentina 3 W-AND 1 0
Dysopsis Euphorbiaceae Malpighiales Andes Costa Rica, Ecuador, Chile, Arg, JF 3 W-AND 2 1
Escallonia Escalloniaceae Escalloniales Andes S Am, SE Brasil, JF 3 W-AND 14 6
Eudema Brassicaceae Brassicales Andes Ecuador to Arg 3 W-AND 3 0
Exodeconus Solanaceae Solanales S Am, Peru, Ecuador, Galapagos, N Chile 3 W-AND 3 0
Geoffroea Fabaceae Fabales S Am from Colombia & Venez to Arg, Brazil 3 W-AND 1 0
Llagunoa Sapindaceae Sapindales Andean trop S Am 3 W-AND 1 1
Luciliocline Asteraceae Asterales Andes S Am, Venezuela to Peru, Chile, Arg 3 W-AND 3 0
Appendix A
Native Endemic
Appendix A
Mutisia Asteraceae Asterales Andes Colombia to Chile, Arg, Parag, 3 W-AND 23 11

SE Brasil, Urug
Myrcianthes Myrtaceae Myrtales Trop Am, Andes, Brazil, WI 3 W-AND 1 1
Myrteola Myrtaceae Myrtales Colombia to Chile, JF, Brazil 3 W-AND 1 0
Nototriche Malvaceae Malvales Ecuador, Peru, Bol, Chile, Arg 3 W-AND 24 6
Ombrophytum Balanophoraceae Santalales Peru, W Brasil, N Arg, Chile 3 W-AND 1 0
Perezia Asteraceae Asterales Andes from Colombia to Chile, SE Brazil 3 W-AND 20 1
Polylepis Rosaceae Rosales Andes Venezuela to Arg, Chile 3 W-AND 2 0
Polypsecadium Brassicaceae Brassicales Colombia to Chile/Arg 3 W-AND 3 2
Tristerix Loranthaceae Santalales Andes Colombia to Chile 3 W-AND 3 1
Werneria Asteraceae Asterales Andes Ecuador to Chile 3 W-AND 8 1
Xenophyllum Asteraceae Asterales Andes Colombia to N Arg, Chile 3 W-AND 5 0
Adesmia Fabaceae Fabales Peru, Chile, Arg, S Brasil, TFuego 3 S-AMZ 130 85
Alstroemeria Alstroemeriaceae Liliales Chile, S Brasil, Perú, Arg 3 S-AMZ 35 26
Arjona Santalaceae Santalales Temp S Am, Brazil 3 S-AMZ 3 0
Azara Salicaceae Malpighiales 8 temp Chile-Arg, s subtrop NW Arg, Bol, SE 3 S-AMZ 8 5
Bra, Uruguay, JF
Blumenbachia Loasaceae Cornales Brasil, Uruguay, Paraguay, Argentina, Chile 3 S-AMZ 3 0
Boopis Calyceraceae Asterales Andes, Arg, S Brasil 3 S-AMZ 7 2
Brachystele Orchidaceae Asparagales Trop Am esp. Brasil 3 S-AMZ 1 0
Caiophora Loasaceae Cornales Peru, Bol, Arg, Chile, 1 sp Urug, Bras, Ecuad 3 S-AMZ 5 0
Calotheca Poaceae Poales Arg, Bras, Chile, Uruguay 3 S-AMZ 1 0
Calycera Calyceraceae Asterales Temp S Am, Brazil 3 S-AMZ 7 4
Chevreulia Asteraceae Asterales S Brasil, Bol, Paraguay, Urug, Chile, N C Arg, 3 S-AMZ 4 0
Falkland
Chloraea Orchidaceae Asparagales Andes Chile Perú, Arg, Brazil 3 S-AMZ 32 18
Ciclospermum Apiaceae Apiales Arg, Bol, Brasil, Chile, Paraguay, Uruguay 3 S-AMZ 2 0
Cliococca Linaceae Malpighiales Brasil, Uruguay, Paraguay, Argentina, Chile 3 S-AMZ 1 0
305
306
Native Endemic
Codonorchis Orchidaceae Asparagales S trop & temp S Am, Brazil 3 S-AMZ 1 0

Colletia Rhamnaceae Rosales Subtrop & temp S Am, Brazil, JF 3 S-AMZ 3 2
Colliguaja Euphorbiaceae Malpighiales Temp S Am, S Brasil 3 S-AMZ 3 2
Crinodendron Elaeocarpaceae Oxalidales Bol, N Arg, C Chile, Brasil 3 S-AMZ 2 2
Dasyphyllum Asteraceae Asterales Chile, Andes, SE Brazil 3 S-AMZ 2 1
Deuterocohnia Bromeliaceae Poales Peru, Bol, SW Bra, N Chile, N Arg 3 S-AMZ 1 1
Facelis Asteraceae Asterales S Brasil, Peru, Bol, Paraguay, Urug, Chile, Arg 3 S-AMZ 2 0
Herbertia Iridaceae Asparagales Temp S Am, Chile, Brazil 3 S-AMZ 1 0
Herreria Asparagaceae Asparagales Brasil, Uru, Parag, Arg, Chile 3 S-AMZ 1 1
Jaborosa Solanaceae Solanales Temp S Am. S Peru, Bol, Bras, Parag, Chile, 3 S-AMZ 11 1
Arg, Uru
Ligaria Loranthaceae Santalales C Peru, E Bol, C Chile, Urug, n Arg, S Brasil 3 S-AMZ 1 0
Lithrea Anacardiaceae Sapindales S Am: Arg, Bol, Paraguay, SE Brazil, Chile. 3 S-AMZ 1 1
Lucilia Asteraceae Asterales Andes S Am, Chile, Arg, Peru 3 S-AMZ 3 1
Margyricarpus Rosaceae Rosales S Andes to Chile, JF, Brazil 3 S-AMZ 2 1
Micropsis Asteraceae Asterales Brazil, Arg, Urug, Chile 3 S-AMZ 1 1
Myrceugenia Myrtaceae Myrtales Chile, Arg, SE Brasil, JF 3 S-AMZ 14 10
Noticastrum Asteraceae Asterales Peru, Bol, Chile, Arg, Ecuador, Brazil 3 S-AMZ 5 4
Ophryosporus Asteraceae Asterales Trop & subtrop S Am, Brazil 3 S-AMZ 8 3
Picrosia Asteraceae Asterales Subtrop S Am, Chile, Peru, Arg, Brazil 3 S-AMZ 1 0
Pleurophora Lythraceae Myrtales S Am, Brazil 3 S-AMZ 3 3
Quillaja Quillajaceae Fabales Temp S Am, Arg, Chile, Brazil 3 S-AMZ 1 1
Rhodophiala Amaryllidaceae Asparagales Chile, Arg, Bol, Bras 3 S-AMZ 27 24
Rhombolytrum Poaceae Poales S Brasil, Urug, Chile 3 S-AMZ 1 1
Skytanthus Apocynaceae Gentianales Brasil, Chile 3 S-AMZ 1 1
Tessaria Asteraceae Asterales S Am, Brazil 3 S-AMZ 1 0
Tweedia Apocynaceae Gentianales Bol, Uru, Chile, Arg 3 S-AMZ 3 3
Viviania Vivianiaceae Geraniales S Brasil, Chile, Arg 3 S-AMZ 3 1
Appendix A
Native Endemic
Appendix A
Cruckshanksia Rubiaceae Gentianales Chile, Arg 3 C-AND 7 4

Gymnophyton Apiaceae Apiales Andes Chile Arg 3 C-AND 5 5
Kurzamra Lamiaceae Lamiales Chile/Arg 3 C-AND 1 0
Lenzia Portulacaceae Caryophyllales Chile Arg 3 C-AND 1 0
Monttea Veronicaceae Lamiales Chile 1, Arg. 2 3 C-AND 1 1
Pyrrhocactus Cactaceae Caryophyllales Arg, Chile 3 C-AND 20 20
Reyesia Solanaceae Solanales Chile Arg 3 C-AND 4 3
Schizopetalon Brassicaceae Brassicales C N Chile, Arg 3 C-AND 10 9
Mulguraea Verbenaceae Lamiales Chile, Arg, Peru 3 C-AND 2 1
Werdermannia Brassicaceae Brassicales N Chile, Arg 3 C-AND 3 3
Ageratina Asteraceae Asterales E USA, C & W S Am, West Indies 3 W-NT 1 0
Aloysia Verbenaceae Lamiales Am: SW US to Chile & Arg 3 W-NT 1 1
Ambrosia Asteraceae Asterales Canada, Mexico, S Am, West Indies 3 W-NT 1 0
Antidaphne Santalaceae Santalales W trop SA: S Mexico, S & S Am 3 W-NT 1 1
Argemone Papaveraceae Ranunculales N & S AM, WI, Hawaii 3 W-NT 4 2
Argyrochosma Pteridaceae Polypodiales Trop Am, USA to Chile 3 W-NT 2 1
Asteriscium Apiaceae Apiales Mexico to Patagonia 3 W-NT 4 4
Astrolepis Pteridaceae Polypodiales Trop Am, USA to Chile 3 W-NT 1 0
Baccharis Asteraceae Asterales Trop AM, W N Am, esp S Am 3 W-NT 40 11
Bomarea Alstroemeriaceae Liliales Mex to trop Am 3 W-NT 3 1
Bouteloua Poaceae Poales Am from Canada to Argentina 3 W-NT 1 0
Bowlesia Apiaceae Apiales N & S Am, B. incana S US 3 W-NT 7 3
Calandrinia Portulacaceae Caryophyllales Am W USA to Chile 3 W-NT 10 2
Calceolaria Calceolariaceae Lamiales AM from Mex to TFuego 3 W-NT 59 45
Calydorea Iridaceae Asparagales Trop & Subtrop Am 3 W-NT 1 1
Cestrum Solanaceae Solanales Trop Am, esp Brasil & Andes 3 W-NT 1 0
Chaptalia Asteraceae Asterales Trop & subtrop Am, S USA 3 W-NT 1 0
Chusquea Poaceae Poales C & S Am, WI, JF 3 W-NT 10 7
307
308
Native Endemic
Coreopsis Asteraceae Asterales N & S Am 3 W-NT 1 1

Dalea Fabaceae Fabales Am, Canada to Arg 3 W-NT 3 2
Dichanthelium Poaceae Poales Toda America 3 W-NT 1 0
Drimys Winteraceae Canellales Mexico, C & S Am, JF 3 W-NT 3 2
Echinodorus Alismataceae Alismatales USA to Argentina and Chile 3 W-NT 1 0
Elodea Hydrocharitaceae Alismatales Am temp 3 W-NT 1 0
Erechtites Asteraceae Asterales N & S Am 3 W-NT 1 1
Fuertesimalva Malvaceae Malvales Mexico, C Andes, Galapagos, ID 3 W-NT 3 1
Gamochaeta Asteraceae Asterales N & S Am, Brazil, JF 3 W-NT 24 10
Grabowskia Solanaceae Solanales Mexico, Galapagos, temp S Am 3 W-NT 1 0
Greigia Bromeliaceae Poales C & S Am, JF 3 W-NT 4 4
Grindelia Asteraceae Asterales W N Am & S Am, Brazil 3 W-NT 3 0
Habranthus Amaryllidaceae Asparagales USA, Mexico to Argentina and Chile 3 W-NT 1 0
Helenium Asteraceae Asterales W N Am, Mexico, C America, Andes 3 W-NT 7 6
Heterosperma Asteraceae Asterales SW USA, Mexico, C Am, Peru, Arg, Chile 3 W-NT 2 0
Jarava Poaceae Poales Arg, Bol, Chile, Colombia, CosRica, Ecuador, 3 W-NT 19 1
El Salvador, Guatemala, Mexico, Peru, USA,
Uruguay, Venezuela
Krameria Krameriaceae Zygophyllales SW US, Mexico, Ecuador, Chile, Arg, Brazil 3 W-NT 2 1
Lachemilla Rosaceae Rosales Mexico to Andes Chile Arg 3 W-NT 3 0
Lepechinia Lamiaceae Lamiales California, Andes, Mexico to S Brasil, 3 W-NT 3 3
Arg, Chile
Lophosoria Dicksoniaceae Cyatheales Trop Am 3 W-NT 1 0
Mecardonia Veronicaceae Lamiales Am S US to Arg 3 W-NT 1 0
Mentzelia Loasaceae Cornales SW Nam, WI, Canada, 3 W-NT 3 1
neotropics to Chile, Arg
Monnina Polygalaceae Fabales Am SW US to Chile 3 W-NT 2 2
Appendix A
Native Endemic
Appendix A
Nama Boraginaceae Unplaced N Am from California, Mex, C Am to Brasil, 3 W-NT 2 0

WI, Hawaii
Nasa Loasaceae Cornales Colombia, Ecuador, Peru, few in Chile, 3 W-NT 1 0
Bolivia, Venezuela and C Am (to S Mexico).
Nassella Poaceae Poales C & S Am 3 W-NT 28 11
Nierembergia Solanaceae Solanales Mex to Arg, Bras 3 W-NT 1 0
Nothoscordum Alliaceae Asparagales Am 3 W-NT 5 3
Oenothera Onagraceae Myrtales Temp & subtrop N C & S Am 3 W-NT 14 3
Olsynium Iridaceae Asparagales Andean and temp S Am and NW N Am 3 W-NT 10 3
Piptochaetium Poaceae Poales Am US to Arg, JF 3 W-NT 7 3
Puya Bromeliaceae Poales C & S Am, Brazil 3 W-NT 8 8
Rostraria Poaceae Poales Arg, Caribe, Chil, Mexico, Peru, USA,Urug 3 W-NT 1 0
Schinus Anacardiaceae Sapindales Mexico to Arg 3 W-NT 11 4
Schkuhria Asteraceae Asterales Trop & subtrop Am 3 W-NT 2 0
Sisyrinchium Iridaceae Asparagales N, C & S Am 3 W-NT 18 10
Stenandrium Acanthaceae Lamiales S US to C Chile 3 W-NT 1 0
Stevia Asteraceae Asterales S US to Chile 3 W-NT 2 1
Tagetes Asteraceae Asterales N & S Am 3 W-NT 3 0
Tillandsia Bromeliaceae Poales Trop & subtrop Am, WI 3 W-NT 5 1
Trixis Asteraceae Asterales SW N Am to Chile 3 W-NT 1 0
Tropaeolum Tropaeolaceae Brassicales Mexico to Chile 3 W-NT 18 15
Ugni Myrtaceae Myrtales Am from Mex to Chile, JF 3 W-NT 3 2
Vasconcellea Caricaceae Brassicales Mexico to N Arg & S Brasil 3 W-NT 1 1
Verbesina Asteraceae Asterales Trop & subtrop Am, WI 3 W-NT 3 1
Viguiera Asteraceae Asterales Trop & subtrop Am, WI 3 W-NT 4 2
Villanova Asteraceae Asterales Mex to Chile 3 W-NT 2 1
Agalinis Orobanchaceae Lamiales Trop, warm C & S Am, USA 4 ANT-P 2 1
Agoseris Asteraceae Asterales 9 sp W N Am, Mexico, 2 sp Chile, Arg 4 ANT-P 2 1
309
310
Native Endemic
Allionia Nyctaginaceae Caryophyllales SW & C US, Mexico to Chile & Arg 4 ANT-P 1 0
Amblyopappus Asteraceae Asterales Calif, NW Mexico, Chile 4 ANT-P 1 0
Amphiscirpus Cyperaceae Poales Canada, USA, Arg, Chile 4 ANT-P 1 0
Amsinckia Boraginaceae Unplaced W US, W temp S Am 4 ANT-P 2 0
Bahia Asteraceae Asterales SW US, Mex, Chile 4 ANT-P 1 1
Blennosperma Asteraceae Asterales 2 California, 1 Chile 4 ANT-P 1 1
Bromidium Poaceae Poales S US & subtrop S Am 4 ANT-P 2 1
Bryantiella Polemoniaceae Ericales W N Am, Chile 4 ANT-P 1 0
Camissonia Onagraceae Myrtales W N Am, Mex, S Am temp 4 ANT-P 1 0
Cardionema Caryophyllaceae Caryophyllales W N Am to Chile 4 ANT-P 3 1
Chascolytrum Poaceae Poales Central & S Am, USA 4 ANT-P 1 0
Chiropetalum Euphorbiaceae Malpighiales Mexico 2, Peru, S Bras, Arg, Chile 4 ANT-P 4 3
Chorizanthe Polygonaceae Caryophyllales Arid & semiarid W Am, Chile 4 ANT-P 13 12
Cistanthe Portulacaceae Caryophyllales N Am, Calif, Arizona, N México & 4 ANT-P 24 15
S Am, Peru, Bol, Arg, Chile
Clarkia Onagraceae Myrtales N Am spec. California, 1 sp Chile/Arg 4 ANT-P 1 0
Collomia Polemoniaceae Ericales W N Am, Bol to Patagonia 4 ANT-P 1 0
Cryptantha Boraginaceae Unplaced W US, W temp S Am 4 ANT-P 40 31
Dissanthelium Poaceae Poales Peru, Chile, Bol to California 4 ANT-P 1 0
Downingia Campanulaceae Asterales W N Am, 1 Chile 4 ANT-P 1 0
Encelia Asteraceae Asterales SW USA, Mexico, Chile, Peru, Galapagos 4 ANT-P 1 0
Errazurizia Fabaceae Fabales SW US, Mex, 1 sp Chile 4 ANT-P 1 1
Flourensia Asteraceae Asterales SW USA, Mexico, Peru, Chile, Arg 4 ANT-P 1 1
Gayophytum Onagraceae Myrtales Temp W N Am & W S Am 4 ANT-P 2 0
Gilia Polemoniaceae Ericales W N Am & S Am 4 ANT-P 3 0
Glandularia Verbenaceae Lamiales Disjunct N Am S Am 4 ANT-P 12 3
Gutierrezia Asteraceae Asterales W N Am & South S Am 4 ANT-P 6 5
Hoffmannseggia Fabaceae Fabales 25 SW US, Chile, Arg 4 ANT-P 8 0
Appendix A
Native Endemic
Appendix A
Ipomopsis Polemoniaceae Ericales W N Am & Florida, 1 Chile Arg 4 ANT-P 1 0

Larrea Zygophyllaceae Zygophyllales SW N Am, S Am 4 ANT-P 2 0
Lastarriaea Polygonaceae Caryophyllales California, Chile 4 ANT-P 1 1
Lasthenia Asteraceae Asterales SW US, Chile-Arg 4 ANT-P 1 0
Legenere Campanulaceae Asterales 1 Calif, 1 Chile 4 ANT-P 1 0
Leptosiphon Polemoniaceae Ericales W N Am, Chile 4 ANT-P 1 1
Lepuropetalon Celastraceae Celastrales SE USA, Mexico, Chile, Arg Uruguay 4 ANT-P 1 0
Lilaea Juncaginaceae Alismatales Pacific Am Canada to Chile 4 ANT-P 1 0
Madia Asteraceae Asterales W N Am, Chile-Arg 4 ANT-P 2 1
Malacothrix Asteraceae Asterales W N Am, Chile 4 ANT-P 1 0
Mancoa Brassicaceae Brassicales Mexico, Andes Peru, Bol, Arg, Chile 4 ANT-P 1 0
Microsteris Polemoniaceae Ericales W N Am & S Am 4 ANT-P 1 0
Munroa Poaceae Poales 1 W US, 4 SW S Am 4 ANT-P 3 0
Navarretia Polemoniaceae Ericales W N Am, Chile, Arg 4 ANT-P 1 0
Nitrophila Amaranthaceae Caryophyllales US, Mex, Chile, Arg 4 ANT-P 1 1
Oxytheca Polygonaceae Caryophyllales W N Am, Chile, Arg 4 ANT-P 1 0
Pectocarya Boraginaceae Unplaced W N Am, S Am 4 ANT-P 3 1
Perityle Asteraceae Asterales SW N Am, C Am, 1 Chile Peru 4 ANT-P 1 0
Phacelia Boraginaceae Unplaced W N Am, S Am 4 ANT-P 6 0
Podagrostis Poaceae Poales Arg, Chile, Canadá, USA 4 ANT-P 1 0
Porlieria Zygophyllaceae Zygophyllales S USA, Mexico, S Am 4 ANT-P 1 1
Psilocarphus Asteraceae Asterales W US, temp S Am 4 ANT-P 2 0
Sphaeralcea Malvaceae Malvales W N Am, Mex, S Am 4 ANT-P 2 1
Symphyotrichum Asteraceae Asterales N Am, Mexico, S Am 4 ANT-P 4 0
Tarasa Malvaceae Malvales Mexico (2), disjunct to Peru, Chile, Arg 4 ANT-P 10 4
Tigridia Iridaceae Asparagales Mex, Guatemala disjunct to Peru, Chile, Bol, 4 ANT-P 1 0
Brazil
Tiquilia Boraginaceae Unplaced Arid N & S Am 4 ANT-P 5 1
311
312
Native Endemic
Tropidocarpum Brassicaceae Brassicales California, Chile 4 ANT-P 1 1

Vahlodea Poaceae Poales Arg, Chile, Canadá, USA 4 ANT-P 1 0
Distichlis Poaceae Poales Am from Canada to Argentina, 1 sp Australia 4 C-PAC 3 0
Flaveria Asteraceae Asterales W N Am, Mexico, S Am, 1 sp Aus 4 C-PAC 1 0
Gaultheria Ericaceae Ericales E & SE Asia, Malesia, Aus, NZ, N, C, S Am, 4 C-PAC 10 1
WI, JF
Gochnatia Asteraceae Asterales Trop & Subtrop Am, S USA, SE Asia 4 C-PAC 1 1
Lilaeopsis Apiaceae Apiales Am Canada to TFuego, Aus, NZ 4 C-PAC 1 0
Microseris Asteraceae Asterales W N Am, 1 Chile, 1 Aus, NZ 4 C-PAC 1 0
Plagiobothrys Boraginaceae Unplaced W N Am, South Am, E Asia, Aus 4 C-PAC 17 11
Sicyos Cucurbitaceae Cucurbitales Aus, Tasm, NZ, Hawaii, N & S Am, ID 4 C-PAC 1 0
Soliva Asteraceae Asterales S Am, N Am, Aus 4 C-PAC 2 0
Acaena Rosaceae Rosales S Afr (1), New Guinea, Aus, Tasmania, NZ, 4 C-PAC 19 2
subantarctic Is, Polynesia, Hawaii,
California to SA, JF
Carpobrotus Aizoaceae Caryophyllales S Afr, Aus, Tasman, NZ, Calif, Chile 4 C-PAC 1 0
Weinmannia Cunoniaceae Oxalidales Madag, Australasia, Mex, C & S Am 4 C-PAC 1 0
Adenocaulon Asteraceae Asterales W NA, Mexico, Hawaii, Guatemala, Chile, 4 W-ANT 1 0
Arg, Himalayas, E Asia
Agrostis Poaceae Poales Temp & warm regions, montane tropics, JF, IP 4 W-ANT 22 4
Alopecurus Poaceae Poales Warm temp & subtrop regions & 4 W-ANT 6 2
montane tropics
Amelichloa Poaceae Poales S Am, Mexico 4 W-ANT 2 0
Anaphalis Asteraceae Asterales Asia, North and South America 4 W-ANT 1 1
Androsace Primulaceae Ericales N Hemisphere, China, few sp SA (TFuego) 4 W-ANT 1 0
Anemone Ranunculaceae Ranunculales Euras, Sumatra, S & E Afr, NZ, 4 W-ANT 7 3
A multifida disj N Am to Chile
Antennaria Asteraceae Asterales Arctic & temp Euras, Andes S Am 4 W-ANT 1 0
Appendix A
Native Endemic
Appendix A
Arenaria Caryophyllaceae Caryophyllales America, temp Eurasia, N Afr 4 W-ANT 7 3

Armeria Plumbaginaceae Caryophyllales SAm Andes to Tfuego, Euro, Russia 4 W-ANT 1 0
Artemisia Asteraceae Asterales Temp Eurasia & W N Am, S Afr, S Am, 4 W-ANT 2 0
Pacific Is
Astragalus Fabaceae Fabales N Am, Andes, temp Eurasia, montane trop Afr 4 W-ANT 49 23
Bartsia Orobanchaceae Lamiales Circumboreal, Eur, Medit, Afr, Andes 4 W-ANT 3 1
Berberis Berberidaceae Ranunculales Euras, N Afr, N & S AM 4 W-ANT 19 11
Botrychium Ophioglossaceae Ophioglossales Temp, polar, trop montane 4 W-ANT 2 0
Bromus Poaceae Poales Eurasia, N & S Afr, montane trop Asia, Am, JF 4 W-ANT 10 0
Caltha Ranunculaceae Ranunculales Arctic & temp N Hemis, NZ, temp S AM 4 W-ANT 3 0
Cardamine Brassicaceae Brassicales Cosmop, temp N hemis & montane tropics, JF 4 W-ANT 15 4
Castilleja Orobanchaceae Lamiales Euras, E N Am, C & S Am 4 W-ANT 3 0
Catabrosa Poaceae Poales Temp Eurasia, N Am, Chile 4 W-ANT 2 0
Chrysosplenium Saxifragaceae Saxifragales N & temp S Am, Euro, N Afr, Asia, 4 W-ANT 2 0
Cicendia Gentianaceae Gentianales 1 Euro & Medit, 1 Calif, W S Am 4 W-ANT 1 0
Cryptogramma Pteridaceae Polypodiales Temperate N & S 4 W-ANT 1 0
Cystopteris Woodsiaceae Polypodiales Temp & warm regions, montane tropics 4 W-ANT 1 0
Danthonia Poaceae Poales N & S Am, Euro, Caucasus 4 W-ANT 4 2
Deschampsia Poaceae Poales Temp & cool, espec N Euro, montane tropics 4 W-ANT 15 3
Descurainia Brassicaceae Brassicales S Am, N Am, Canary Is. 4 W-ANT 12 4
Draba Brassicaceae Brassicales Arctic & temp n Hemis, Asia, montane C & 4 W-ANT 8 1
S Am
Elatine Elatinaceae Malpighiales Temp, trop & subtrop regions 4 W-ANT 1 0
Empetrum Ericaceae Ericales N temp & Arctic, Europa, S Andes, 4 W-ANT 1 0
Falkland, JF
Ephedra Ephedraceae Ephedrales Eurasia, AM 4 W-ANT 7 2
Epilobium Onagraceae Myrtales Temp Euro, N Am, arctic, montane trop 4 W-ANT 12 2
Erigeron Asteraceae Asterales Eurasia, N & S Am, Aus, JF 4 W-ANT 17 8
313
314
Native Endemic
Euphrasia Orobanchaceae Lamiales Temp N Hemishp, Euro, Middle Asia, 4 W-ANT 16 12

Australia, NZ, temp S Am, JF
Fagonia Zygophyllaceae Zygophyllales Medit, Euro, SW Asia, NE Afr, SW N Am, 4 W-ANT 1 0
Peru, Chile
Gentiana Gentianaceae Gentianales Euro, NW Afr, Asia, E Aus, Am 4 W-ANT 1 0
Glycyrrhiza Fabaceae Fabales Eurasia, Aus, N Am & temp S Am 4 W-ANT 1 0
Halerpestes Ranunculaceae Ranunculales Temperate Asia, N & S Am 4 W-ANT 2 0
Hieracium Asteraceae Asterales Temp regions & montane tropics, excl 4 W-ANT 4 0
Australasia
Hippuris Veronicaceae Lamiales Circumboreal, S Am, Australia 4 W-ANT 1 0
Hordeum Poaceae Poales Temp & subtrop regs, montane tropics 4 W-ANT 9 1
Hydrangea Hydrangeaceae Cornales Himalayas, Asia, Philippines, N, C Am, 4 W-ANT 1 0
Andes to Chile
Hypochaeris Asteraceae Asterales Temp S Am, Eurasia Medit, N Afr 4 W-ANT 31 17
Isolepis Cyperaceae Poales Cape Region S Afr, Australasia, S Am, north 4 W-ANT 4 0
temperate
Lathyrus Fabaceae Fabales Temp N hemis, Medit, montane tropics, 4 W-ANT 13 3
E Afr, S Am
Leymus Poaceae Poales Temp regs 4 W-ANT 1 0
Limonium Plumbaginaceae Caryophyllales N temperate, circumboreal, austral 4 W-ANT 1 1
Littorella Veronicaceae Lamiales 1 N Am, 1 S Am, 1 Euro 4 W-ANT 1 0
Lupinus Fabaceae Fabales N & S Am, Medit, montane trops, E Afr 4 W-ANT 6 2
Malvella Malvaceae Malvales 3 sp W USA, Mexico, 1 sp Peru to Uruguay, 1 4 W-ANT 1 0
sp. Medit
Melica Poaceae Poales Temp regs & montane trops 4 W-ANT 8 7
Minuartia Caryophyllaceae Caryophyllales Arctic & temp regions, Mexico, Ethiopia, 4 W-ANT 1 1
Himalayas, Chile
Mirabilis Nyctaginaceae Caryophyllales Am trop, Himal, China 4 W-ANT 5 3
Appendix A
Native Endemic
Appendix A
Muhlenbergia Poaceae Poales Temp & subtrop Am, Asia 4 W-ANT 3 0

Myosotis Boraginaceae Unplaced Eurasia, montane trops, S Afr, N Guinea, Aus, 4 W-ANT 2 0
NZ
Myosurus Ranunculaceae Ranunculales Temp regs hemis N & S, M apetalus disj N Am 4 W-ANT 2 0
Noccaea Brassicaceae Brassicales N temp, few S Am 4 W-ANT 1 0
Osmorhiza Apiaceae Apiales N Am, E Asia, temp S Am 4 W-ANT 4 0
Paronychia Caryophyllaceae Caryophyllales North temperate, austral 4 W-ANT 6 3
Pennisetum Poaceae Poales Trop & temp regs 4 W-ANT 1 0
Persea Lauraceae Laurales Trops & subtrops Am & Asia, Macaronesia 4 W-ANT 1 0
Phalaris Poaceae Poales Temperate Eurasia, America 4 W-ANT 2 1
Phleum Poaceae Poales Temperate Eurasia, America 4 W-ANT 1 0
Pinguicula Lentibulariaceae Lamiales Am, Medit, circumboreal 4 W-ANT 2 0
Poa Poaceae Poales Temp & cold & montane trops 4 W-ANT 42 7
Polemonium Polemoniaceae Ericales Temp Eurasia, N Am, Chile 4 W-ANT 1 0
Potentilla Rosaceae Rosales N & S Am, temp Euras 4 W-ANT 1 0
Primula Primulaceae Ericales N Hemisp, trop Asia, temp S Am 4 W-ANT 1 0
Rhamnus Rhamnaceae Rosales N Hemisp, S Afr, S Am 4 W-ANT 1 1
Ribes Grossulariaceae Saxifragales N Hemis, N Afr, S Am Canada to Tfuego 4 W-ANT 11 6
Salix Salicaceae Malpighiales Arctic temp, few in trops, temp S Am, 1 S Afr 4 W-ANT 1 0
Saxifraga Saxifragaceae Saxifragales N temp, Eurasia, W N Am, Andes S Am 4 W-ANT 1 0
Solidago Asteraceae Asterales Eurasia, N Am, Mexico, 1 S Am 4 W-ANT 2 0
Spartina Poaceae Poales Am, W Euro, NW Afr 4 W-ANT 1 0
Spergula Caryophyllaceae Caryophyllales Europe, Patagonia 4 W-ANT 4 1
Stemodia Veronicaceae Lamiales Trops and Subtrops, Madag 4 W-ANT 1 0
Taraxacum Asteraceae Asterales Temp N Hemis & temp S Am, NZ, JF 4 W-ANT 3 1
Tetragonia Aizoaceae Caryophyllales N & S Afr, SE Asia, Aus, Tasm, NZ, Polinesia, 4 W-ANT 8 7
Peru, Chile
Trifolium Fabaceae Fabales Temp & subtrop exc Aus 4 W-ANT 10 5
315
316
Native Endemic
Triglochin Juncaginaceae Alismatales N temperate, circumboreal, austral 4 W-ANT 3 0

Triodanis Campanulaceae Asterales N Am, Mex, Guatem, Medit 4 W-ANT 1 0
Valeriana Valerianaceae Dipsacales N temp Eurasia, S Afr, Andes 4 W-ANT 44 21
Verbena Verbenaceae Lamiales Temp & trop Am, few OW 4 W-ANT 10 4
Vicia Fabaceae Fabales Temp N Hemis, Medit, few trop Afr, S Am, 4 W-ANT 19 12
Hawaii
Woodsia Woodsiaceae Polypodiales Temp & cool temp Eurasia, Afr, Am, exc Aus 4 W-ANT 1 0
Menodora Oleaceae Lamiales Trop & subtrop Am, S Afr 4 1 1
Acrisione Asteraceae Asterales Central Chile, Arg 5 2 1
Aextoxicon Aextoxicaceae Berberidopsidales Chile, Arg 5 1 0
Amomyrtus Myrtaceae Myrtales Temp S Am: Arg, Chile 5 2 1
Anarthrophyllum Fabaceae Fabales Andes Chile Arg 5 6 2
Asteranthera Gesneriaceae Lamiales Chile, Arg 5 1 0
Austrocactus Cactaceae Caryophyllales Arg, Chile 5 3 1
Austrocedrus Cupressaceae Pinales Arg, Chile 5 1 0
Barneoudia Ranunculaceae Ranunculales Arg, Chile 5 3 0
Belloa Asteraceae Asterales Andes S Arg, Chile 5 1 0
Benthamiella Solanaceae Solanales S Chile-Arg, Patagonia 5 4 0
Bipinnula Orchidaceae Asparagales Temp S Am: Chile, Arg 5 4 4
Bolax Apiaceae Apiales Temp SA 5 2 0
Boquila Lardizabalaceae Ranunculales Arg, Chile 5 1 0
Brachyclados Asteraceae Asterales Temp Chile, Arg 5 1 0
Callianthemoides Ranunculaceae Ranunculales Arg, Chile 5 1 0
Campsidium Bignoniaceae Lamiales Arg, Chile: C. valdivianum 5 1 0
Chiliotrichum Asteraceae Asterales S Andes Chile, Argentina 5 2 0
Combera Solanaceae Solanales Andes Chile Arg 5 2 1
Corynabutilon Malvaceae Malvales Chile, Arg 5 7 6
Diostea Verbenaceae Lamiales Temperate S Am, Arg, Chile 5 1 0
Appendix A
Native Endemic
Appendix A
Diplolepis Apocynaceae Gentianales Chile, Arg 5 6 3

Diposis Apiaceae Apiales temp S Am 5 1 1
Doniophyton Asteraceae Asterales Andes Chile Arg 5 2 0
Drapetes Thymelaeaceae Malvales temp S Am 5 1 0
Elytropus Apocynaceae Gentianales Chile, Arg 5 1 0
Embothrium Proteaceae Proteales Chile y Argentina, 1 spp. 5 1 0
Eriachaenium Asteraceae Asterales Tierra del Fuego 5 1 0
Famatina Amaryllidaceae Asparagales Chile, Arg 5 2 2
Fitzroya Cupressaceae Pinales Chile Arg 5 1 0
Fonkia Veronicaceae Lamiales Chile, Arg 5 1 0
Gamocarpha Calyceraceae Asterales Temp S Am 5 6 2
Gamochaetopsis Asteraceae Asterales S Arg, Chile 5 1 0
Gavilea Orchidaceae Asparagales Temp S Am, JF 5 12 6
Gilliesia Alliaceae Asparagales Chile, Arg 5 4 3
Guindilia Sapindaceae Sapindales Chile Arg 5 1 0
Hamadryas Ranunculaceae Ranunculales Antarctic S Am 5 4 0
Huanaca Apiaceae Apiales Chile, Arg 5 4 0
Hymenoglossum Hymenophyllaceae Hymenophyllales Chile, Arg 5 1 0
Lardizabala Lardizabalaceae Ranunculales Chile and adjacent Argentina 5 1 1
Laretia Apiaceae Apiales Chile, Argentina 5 1 0
Laureliopsis Atherospermataceae Laurales Chile Arg 5 1 0
Lebetanthus Ericaceae Ericales Patagonia 5 1 0
Lecanophora Malvaceae Malvales Arg, Chile 5 1 0
Lepidophyllum Asteraceae Asterales Patagonia Chile-Arg 5 1 0
Leptophyllochloa Poaceae Poales Arg Chile, JF 5 1 0
Luma Myrtaceae Myrtales Chile Arg 5 2 1
Macrachaenium Asteraceae Asterales Patagonia & TFuego 5 1 0
Maihuenia Cactaceae Caryophyllales Arg, Chile 5 2 0
317
318
Native Endemic
Melosperma Veronicaceae Lamiales Chile, Arg 5 2 0

Menonvillea Brassicaceae Brassicales Chile, Arg 5 19 13
Misodendrum Misodendraceae Santalales S Am, south of 33 5 8 1
Mitraria Gesneriaceae Lamiales Chile, Arg 5 1 0
Mulinum Apiaceae Apiales S Andes 5 5 1
Myoschilos Santalaceae Santalales Chile, Arg 5 1 0
Nanodea Santalaceae Santalales Temp S Am 5 1 0
Nastanthus Calyceraceae Asterales Chile, Arg, Falkland Is. 5 5 1
Notopappus Asteraceae Asterales Chile Arg 5 4 0
Ochetophila Rhamnaceae Rosales Chile, Arg 5 2 0
Odontorrhynchus Orchidaceae Asparagales Temp S Am: Chile, Arg 5 4 4
Onuris Brassicaceae Brassicales Chile, Patagonia 5 5 0
Oreopolus Rubiaceae Gentianales Chile, Arg, Patagonia 5 1 0
Ortachne Poaceae Poales Arg, Chile 5 2 0
Ovidia Thymelaeaceae Malvales temp S Am 5 1 0
Philesia Philesiaceae Liliales Chile, Arg 5 1 0
Philippiella Caryophyllaceae Caryophyllales Patagonia Chile-Arg. 5 1 0
Phycella Amaryllidaceae Asparagales Chile, Arg 5 6 5
Pilgerodendron Cupressaceae Pinales Chile Arg 5 1 0
Pozoa Apiaceae Apiales Andes Chile Arg 5 2 0
Pterocactus Cactaceae Caryophyllales Patagonia Arg, Chile 5 2 0
Relchela Poaceae Poales Chile Arg 5 1 0
Retanilla Rhamnaceae Rosales Chile, Arg 5 3 3
Rhaphithamnus Verbenaceae Lamiales Chile, Arg, JF 5 2 1
Salpiglossis Solanaceae Solanales Chile, Arg 5 2 1
Sarcodraba Brassicaceae Brassicales Chile, Arg 5 1 0
Saxegothaea Podocarpaceae Pinales Chile, Arg 5 1 0
Saxifragella Saxifragaceae Saxifragales Antarctic S Am 5 1 0
Appendix A
Native Endemic
Appendix A
Saxifragodes Saxifragaceae Saxifragales Tfuego 5 1 0

Schizanthus Solanaceae Solanales Chile, Arg 5 12 10
Serpyllopsis Hymenophyllaceae Hymenophyllales Chile, Arg 5 1 0
Solaria Alliaceae Asparagales Chile, Arg 5 1 0
Solenomelus Iridaceae Asparagales Chile, Arg 5 2 1
Synammia Polypodiaceae Polypodiales Arg, Chile, JF 5 3 2
Tapeinia Iridaceae Asparagales Chile, Arg 5 1 0
Tepualia Myrtaceae Myrtales Chile, Arg 5 1 0
Tetroncium Juncaginaceae Alismatales Temp S Am, Magallanes 5 1 0
Tribeles Escalloniaceae Escalloniales Chile & TFuego 5 1 0
Trichopetalum Asparagaceae Asparagales Chile, Arg 5 1 1
Triptilion Asteraceae Asterales Chile, Arg 5 7 5
Tristagma Alliaceae Asparagales Chile, Arg, Uru 5 6 5
Xerodraba Brassicaceae Brassicales S Arg, Chile 5 2 0
Zoellnerallium Alliaceae Asparagales Chile, Arg 5 2 1
Adenopeltis Euphorbiaceae Malpighiales Chile 6 1 1
Anisomeria Phytolaccaceae Caryophyllales Chile 6 2 2
Araeoandra Vivianiaceae Geraniales Chile 6 1 1
Avellanita Euphorbiaceae Malpighiales Chile 6 1 1
Bakerolimon Plumbaginaceae Caryophyllales Chile 6 1 1
Balsamocarpon Fabaceae Fabales Chile 6 1 1
Bridgesia Sapindaceae Sapindales Chile 6 1 1
Calopappus Asteraceae Asterales Chile 6 1 1
Cissarobryon Vivianiaceae Geraniales Chile 6 1 1
Conanthera Tecophilaeaceae Asparagales Chile 6 5 5
Copiapoa Cactaceae Caryophyllales N Chile 6 22 22
Cyphocarpus Campanulaceae Asterales Chile 6 3 3
Desmaria Loranthaceae Santalales Chile 6 1 1
319
320
Native Endemic
Dinemagonum Malpighiaceae Malpighiales Chile 6 1 1

Dinemandra Malpighiaceae Malpighiales Chile 6 1 1
Epipetrum Dioscoreaceae Dioscoreales Chile 6 3 3
Ercilla Phytolaccaceae Caryophyllales Chile 6 2 2
Eriosyce Cactaceae Caryophyllales Chile 6 5 5
Eulychnia Cactaceae Caryophyllales Chile 6 6 6
Fascicularia Bromeliaceae Poales Chile 6 1 1
Francoa Francoaceae Geraniales Chile 6 1 1
Gethyum Alliaceae Asparagales Chile 6 2 2
Gomortega Gomortegaceae Laurales Chile 6 1 1
Guynesomia Asteraceae Asterales Chile 6 1 1
Gymnachne Poaceae Poales Chile 6 1 1
Gypothamnium Asteraceae Asterales N Chile 6 1 1
Hollermayera Brassicaceae Brassicales Chile 6 1 1
Homalocarpus Apiaceae Apiales Chile 6 6 6
Huidobria Loasaceae Cornales Chile 6 2 2
Ivania Brassicaceae Brassicales N Chile 6 1 1
Jubaea Arecaceae Arecales Chile 6 1 1
Lapageria Philesiaceae Liliales Chile 6 1 1
Latua Solanaceae Solanales Chile 6 1 1
Legrandia Myrtaceae Myrtales Chile 6 1 1
Leontochir Alstroemeriaceae Liliales Chile 6 1 1
Leptocarpha Asteraceae Asterales Chile 6 1 1
Leucocoryne Alliaceae Asparagales Chile 6 14 14
Leunisia Asteraceae Asterales C Chile 6 1 1
Marticorenia Asteraceae Asterales Chile 6 1 1
Metharme Zygophyllaceae Zygophyllales N Chile 6 1 1
Microphyes Caryophyllaceae Caryophyllales Chile 6 3 3
Appendix A
Native Endemic
Appendix A
Miersia Alliaceae Asparagales Chile 6 4 4

Miqueliopuntia Cactaceae Caryophyllales Chile 6 1 1
Moscharia Asteraceae Asterales Chile 6 2 2
Neoporteria Cactaceae Caryophyllales Chile 6 6 6
Notanthera Loranthaceae Santalales Chile temperate, JF 6 1 1
Ochagavia Bromeliaceae Poales Chile, JF 6 4 4
Oxyphyllum Asteraceae Asterales N Chile 6 1 1
Peumus Monimiaceae Laurales Chile 6 1 1
Phrodus Solanaceae Solanales Chile (Atacama & Coquimbo) 6 1 1
Pintoa Zygophyllaceae Zygophyllales Chile 6 1 1
Pitavia Rutaceae Sapindales Chile 6 1 1
Placea Amaryllidaceae Asparagales Chile 6 5 5
Pleocarphus Asteraceae Asterales N Chile 6 1 1
Podanthus Asteraceae Asterales Chile 6 2 2
Reicheella Caryophyllaceae Caryophyllales Chile 6 1 1
Sarmienta Gesneriaceae Lamiales Chile 6 1 1
Scyphanthus Loasaceae Cornales Chile 6 1 1
Speea Alliaceae Asparagales Chile 6 1 1
Tecophilaea Tecophilaeaceae Asparagales Chile 6 2 2
Tetilla Francoaceae Geraniales Chile 6 1 1
Thelocephala Cactaceae Caryophyllales Chile 6 6 6
Traubia Amaryllidaceae Asparagales Chile 6 1 1
Trevoa Rhamnaceae Rosales Chile 6 1 1
Valdivia Escalloniaceae Escalloniales Chile 6 1 1
Vestia Solanaceae Solanales Chile 6 1 1
Zephyra Tecophilaeaceae Asparagales Chile 6 1 1
Centaurodendron Asteraceae Asterales JF 6 2 2
Cuminia Lamiaceae Lamiales JF 6 1 1
321
322
Native Endemic
Dendroseris Asteraceae Asterales JF 6 11 11

Juania Arecaceae Arecales JF 6 1 1
Lactoris Lactoridaceae Piperales JF 6 1 1
Lycapsus Asteraceae Asterales Desventuradas Islands (Chile) 6 1 1
Margyracaena Rosaceae Rosales JF 6 1 1
Megalachne Poaceae Poales JF 6 2 2
Nesocaryum Boraginaceae Unplaced Desventuradas Islands (Chile) 6 1 1
Podophorus Poaceae Poales JF 6 1 1
Robinsonia Asteraceae Asterales JF 6 8 8
Sanctambrosia Caryophyllaceae Caryophyllales Desventuradas Islands (Chile) 6 1 1
Selkirkia Boraginaceae Unplaced JF 6 1 1
Thamnoseris Asteraceae Asterales Desventuradas Islands (Chile) 6 1 1
Thyrsopteris Thyrsopteridaceae Cyatheales JF 6 1 1
Yunquea Asteraceae Asterales JF 6 1 1
Adiantum Pteridaceae Polypodiales Cosmop, esp Neotrops, Madag 7 5 3
Amaranthus Amaranthaceae Caryophyllales Subtrop & temp, Euro, S Am, N Am, 16 sp 7 2 1
Russia,
Anagallis Primulaceae Ericales Euro, Russia, N & S Afr, Madag, W Asia, 7 1 0
Himalayas, N Aus, 2 sp. S Am
Anthoxanthum Poaceae Poales Temp & warm Eurasia, N Afr, montane tropics 7 7 2
Aphanes Rosaceae Rosales Am, Euro, Medit, Ethiop, Australia, Caucasus 7 3 2
Apium Apiaceae Apiales Euro, Medit, N & S Afr, Madag, W & E Asia, 7 5 3
Malesia, Aus, N, C & S Am, JF, IP
Aristida Poaceae Poales N Am, Medit, temp Asia, tropics & subtropics 7 3 0
Asplenium Aspleniaceae Polypodiales Subcosmop, Madag, JF 7 10 3
Atriplex Amaranthaceae Caryophyllales Cosmop, Eurasia, Aus, ID 7 21 15
Azolla Salviniaceae Salviniales Trop & warm, Euro, Asia, Madag, Tristan da 7 1 0
Cunha, Hawaii
Appendix A
Native Endemic
Appendix A
Bidens Asteraceae Asterales Cosmop, mostly Am 7 2 0

Blechnum Blechnaceae Polypodiales Subcosmop, Madag, JF 7 13 6
Bothriochloa Poaceae Poales Temp & subtrop, montane tropics 7 2 0
Callitriche Veronicaceae Lamiales Cosmop 7 6 1
Calystegia Convolvulaceae Solanales Cosmop, temp & trop hemis N & S, JF 7 2 0
Carex Cyperaceae Poales Cosmop, JF 7 67 16
Cenchrus Poaceae Poales Temp & subtrop, montane tropics 7 3 0
Centaurea Asteraceae Asterales Eurasia, N China, N & tro Afr, Aus, 7 7 7
NZ, N Am
Centaurium Gentianaceae Gentianales Cosmop exc trop & S Afr, N hemisp 1 ext. to 7 1 0
Aus, 1 to Chile
Cerastium Caryophyllaceae Caryophyllales Cosmop 7 1 0
Ceratophyllum Ceratophyllaceae Ceratophyllales Cosmop 7 1 0
Cheilanthes Pteridaceae Polypodiales Cosmop, esp Neotrops & S Afr, Madag 7 9 0
Chenopodium Amaranthaceae Caryophyllales Cosmop, JF, ID 7 16 4
Chloris Poaceae Poales Cosmop 7 2 0
Clinopodium Lamiaceae Lamiales Temp & subtrop, Medit 7 4 2
Convolvulus Convolvulaceae Solanales Cosmop esp. temp, trop & subtrops 7 6 1
Coriaria Coriariaceae Cucurbitales Mex to Chile, W Medit, Himal to Japan, 7 1 0
New Guinea, NZ. C ruscifolia disj NZ
Corrigiola Caryophyllaceae Caryophyllales S Am Andes Colom to Chile, Euro, Medit, 7 3 3
Afr, Madag
Crassula Crassulaceae Saxifragales Cosmop, Euro, A & S Afr, Madag, Aus, Am 7 7 1
Cuscuta Convolvulaceae Solanales Cosmop 7 11 5
Cynanchum Apocynaceae Gentianales Trops & subtrops 7 4 2
Cynoglossum Boraginaceae Unplaced Temp & subtrop reg 7 1 1
Cyperus Cyperaceae Poales Trop, subtrop, temp regions, JF 7 14 3
323
324
Native Endemic
Daucus Apiaceae Apiales Euro, Medit, SW & S Asia, trop Afr. Aus, 7 2 0
NZ, & Am
Deyeuxia Poaceae Poales Temp & subtrop reg & montane tropics 7 22 0
Digitaria Poaceae Poales Temp & subtrop reg & montane tropics 7 2 0
Diplazium Woodsiaceae Polypodiales Cosmop, IP 7 1 0
Drosera Droseraceae Caryophyllales Cosmop, espec Aus, Tasm & NZ 7 1 0
Dryopteris Dryopteridaceae Polypodiales Cosmop, IP 7 1 0
Eleocharis Cyperaceae Poales Cosmop, JF 7 18 1
Elymus Poaceae Poales Temp & subtrop, montane tropics 7 4 0
Enneapogon Poaceae Poales Temp Eurasia, Am, trop & subtrop 7 1 0
Equisetum Equisetaceae Equisetales Almost cosmop, Madag 7 2 0
Eragrostis Poaceae Poales Temp & trop, ID 7 8 2
Eriochloa Poaceae Poales Temp & subtrop 7 1 0
Erodium Geraniaceae Geraniales Cosmop 7 1 0
Eryngium Apiaceae Apiales Trop & temp, Eurasia, Am, Afr, Malesia, 7 15 13
Aus, JF
Euphorbia Euphorbiaceae Malpighiales Subcosmop 7 23 16
Festuca Poaceae Poales Cosmop 7 25 5
Frankenia Frankeniaceae Caryophyllales Temp Euro, Afr, Aus, Am, ID 7 6 1
Galium Rubiaceae Gentianales Cosmop, JF 7 20 9
Gentianella Gentianaceae Gentianales Eurasia, NW Afr, N, C & S Am, Aus, NZ 7 5 2
Geranium Geraniaceae Geraniales Cosmop, espec tem regs & tropic montane 7 5 1
Geum Rosaceae Rosales Arctic & temp regions 7 7 4
Glyceria Poaceae Poales Temp & subtrop, montane tropics 7 1 0
Gnaphalium Asteraceae Asterales Cosmop 7 27 14
Gratiola Veronicaceae Lamiales Temp & subtrop, montane tropics 7 1 0
Heliotropium Boraginaceae Unplaced Temp, subtrop, trop regs, arid 7 22 16
Huperzia Lycopodiaceae Lycopodiales Subcosmop, Madag 7 2 0
Appendix A
Native Endemic
Appendix A
Hydrocotyle Apiaceae Apiales Cosmop 7 8 1

Hymenophyllum Hymenophyllaceae Hymenophyllales Trop & S temp, Euro, Japan, Madag, JF 7 19 3
Hypericum Hypericaceae Malpighiales Temp regs, montane tropics 7 2 1
Imperata Poaceae Poales Trop, subtrop & temp regions 7 2 1
Isoetes Isoetaceae Isoetales Cosmop, Madag 7 3 0
Juncus Juncaceae Poales Cosmop, JF, IP 7 22 2
Koeleria Poaceae Poales Temp & subtrops, montane trops 7 3 0
Landoltia Araceae Alismatales Cosmop 7 1 0
Lemna Araceae Alismatales Cosmop 7 3 0
Lepidium Brassicaceae Brassicales Cosmop, ID 7 25 14
Limosella Scrophulariaceae Lamiales Cosmop, Madag 7 1 0
Lindernia Linderniaceae Lamiales Trops, subtrops, temp, Madag 7 1 0
Linum Linaceae Malpighiales Temp & subtrops 7 5 4
Ludwigia Onagraceae Myrtales Cosmop 7 4 0
Luzula Juncaceae Poales Cosmop, JF 7 7 1
Lycium Solanaceae Solanales N Am, tem S Am, Eurasia, S Afr, Aus, IP 7 11 2
Lycopodium Lycopodiaceae Lycopodiales Trop & temp 7 6 0
Lysimachia Primulaceae Ericales Cosmop 7 1 0
Marsilea Marsileaceae Salviniales Trop & temp, Madag 7 1 0
Mimulus Phrymaceae Lamiales Am, S Afr, Madag, E Asia, JF 7 7 2
Montia Portulacaceae Caryophyllales Temp Eurasia, Medit, montane tropics, Afr, 7 1 0
A Aus, A & S Am,
Myriophyllum Haloragaceae Saxifragales Subcosmop 7 2 0
Ophioglossum Ophioglossaceae Ophioglossales Subcosmop, Madag, JF 7 5 1
Orobanche Orobanchaceae Lamiales Temp, subtrop, N Hemis 7 2 1
Oxalis Oxalidaceae Oxalidales Cosmop 7 53 26
Panicum Poaceae Poales Temp, subtrop, trop regs 7 3 0
Parietaria Urticaceae Rosales Subcosmop, JF, ID 7 1 0
325
326
Native Endemic
Paspalum Poaceae Poales Temp & trop regs, IP 7 8 0

Phragmites Poaceae Poales Cosmop 7 1 0
Pilularia Marsileaceae Salviniales Euro, Aus, NZ, Ethiopia, W S Am 7 1 0
Plantago Veronicaceae Lamiales Cosmop, JF, ID 7 23 8
Pleurosorus Aspleniaceae Polypodiales España, Marruecos, Aus, Tasm, NZ, Chile, Arg 7 1 0
Plumbago Plumbaginaceae Caryophyllales Trop & warm-temp regs 7 1 0
Polycarpon Caryophyllaceae Caryophyllales Cosmop 7 1 1
Polygala Polygalaceae Fabales Subcosmop 7 7 2
Polygonum Polygonaceae Caryophyllales Cosmop 7 5 2
Polypogon Poaceae Poales Cosmop 7 8 1
Polystichum Dryopteridaceae Polypodiales Cosmop, Madag, IP 7 6 2
Portulaca Portulacaceae Caryophyllales Trop, subtrop & temp regions 7 1 0
Potamogeton Potamogetonaceae Alismatales Cosmop 7 7 0
Pteris Pteridaceae Polypodiales Cosmop, Madag, JF 7 3 2
Puccinellia Poaceae Poales N temp: Asia, N Am, S temp: S Afr, 7 8 0
Aus, S Am
Ranunculus Ranunculaceae Ranunculales Cosmop, R acaulis disj NZ, Falkland, 7 18 1
R bonariensis disj N Am S Am
Rhynchospora Cyperaceae Poales Cosmp 7 1 0
Rorippa Brassicaceae Brassicales Cosmop 7 3 1
Rubus Rosaceae Rosales Cosmop, JF 7 2 0
Rumex Polygonaceae Caryophyllales Cosmop 7 10 3
Ruppia Ruppiaceae Alismatales Cosmop 7 2 0
Sagina Caryophyllaceae Caryophyllales Temp N Hemis, E Afr, Asia, N Guinea, 7 2 0
Aus, NZ, Andes
Sagittaria Alismataceae Alismatales Cosmop 7 2 0
Salvia Lamiaceae Lamiales Cosmop 7 5 0
Samolus Primulaceae Ericales Cosmop, IP 7 3 1
Appendix A
Appendix A
Native Endemic
Sanicula Apiaceae Apiales Subcosmop, exc Aus & NZ 7 2 0

Sarcocornia Amaranthaceae Caryophyllales Eurasia, S Afr (15), Aus, Tasm, NZ, 7 4 0
N & S Am (8), JF
Schoenoplectus Cyperaceae Poales Cosmp 7 3 0
Schoenus Cyperaceae Poales Subcosmop 7 4 0
Scirpus Cyperaceae Poales North temperate, austral, JF, IP 7 5 3
Scutellaria Lamiaceae Lamiales Cosmop 7 3 1
Senecio Asteraceae Asterales Cosmop 7 224 103
Setaria Poaceae Poales Trop, subtrop & temp regions 7 1 0
Silene Caryophyllaceae Caryophyllales Temp N hemis, Medit, S Afr 7 8 0
Solanum Solanaceae Solanales Subcosmop, JF, ID, IP 7 44 19
Sophora Fabaceae Fabales Trop & most N temp, JF, IP 7 5 4
Spergularia Caryophyllaceae Caryophyllales Cosmop, JF, ID 7 10 9
Sporobolus Poaceae Poales Am, Asia, Afr, 1 Euro 7 2 0
Stachys Lamiaceae Lamiales Trop, subtrop, temp regions exc. Australasia 7 11 9
Stellaria Caryophyllaceae Caryophyllales Cosmop 7 9 4
Stipa Poaceae Poales Temp N Hemis, montane trops, IP 7 1 0
Stuckenia Potamogetonaceae Alismatales Subcosmop 7 3 0
Suaeda Amaranthaceae Caryophyllales Cosmop, ID 7 5 2
Teucrium Lamiaceae Lamiales Cosmop 7 2 2
Trisetum Poaceae Poales Cold & temp, montane tropics, JF 7 15 2
327
328
Native Endemic
Typha Typhaceae Poales Cosmop 7 2 0

Urtica Urticaceae Rosales Subcosmop, temperate 7 10 4
Utricularia Lentibulariaceae Lamiales Cosmop 7 1 0
Veronica Veronicaceae Lamiales Cosmop 7 1 0
Viola Violaceae Malpighiales Cosmop 7 72 50
Vulpia Poaceae Poales N & S Am, Euro, N Afr, M Asia 7 3 0
Wolffia Araceae Alismatales Cosmop, warm temperate and tropical regions 7 1 0
Zannichellia Potamogetonaceae Alismatales Cosmop 7 1 0
Tracks: Austral-antarctic (AUS), Tropical Pacific (TPAC), Circum-austral (C-AUS), Wide Neotropical (W-NT), Wide Andean (W-AND), Central Andean
(C-AND), South-Amazonian (S-AMZ), Wide antitropical (W-ANT), Antitropical Pacific (ANT-P), Circum-Pacific track (C-PAC).
Islands: Juan Fernández (JF), Isla de Pascua (IP), Islas Desventuradas (ID).
Appendix A
General Index
A Australasia, 22, 27, 75, 89–95, 101, 110, 113,

Acantilado, 7 118, 121, 130, 137, 139, 141, 154, 156,
Aconcagua (Monte), 3–4 158, 161, 168, 224–225, 227–229, 238,
Africa-first model, 279–280 249, 252, 254, 257, 259, 261–262, 271,
Agathis, 272–273 275–276
Albert, F., 182 Australasiatic (floristic element), 89–95, 110,
Allochtonous terranes, 12, 261 113, 139, 141, 154, 156, 158, 161, 225,
Altiplano, 3, 25, 32–33, 208, 227, 230, 233 227–229, 238
Amphitropical (element), 13, 123 Austral floristic realm, 93, 136–137, 141, 278
Andean Australia, 10, 16, 22, 27, 63–64, 73, 75, 93,
Cordillera, 8, 210 95, 116–118, 133, 135–137, 157–158,
Dry Diagonal, 111–113 169, 226, 234, 238, 253, 255, 257, 260,
uplift, 25–26, 111–112, 216, 237 262–263, 273, 278–279
vegetation, 32, 34–37
Angiosperms, 17, 19, 22, 57, 66–70, 73–74, B
76–79 Barnadesieae, 223, 225–226, 235–237, 240
Antarctica, 3, 10–11, 16–17, 19, 21–25, 32, Bennettitales, 15–16, 210
35, 38, 60, 73, 116, 121, 169, 221, 234, Bertero, C. G., 55–56
250, 253–260, 275–276, 278–279 Biobío (región, Río), 31, 139–140, 166, 184
Antarctic Bioclimates, 33–35
circumpolar current (ACC), 17, 32, 259, Biodiversity, 185–189, 213
279–280 Biogeographical composite areas, 122
Peninsula, 4, 11, 17, 131, 257–258, 278 Biogeographic
Plate, 6 map, 131–132
Antitropical (floristic element), 89–90, 98–101, regionalization, 129–147, 229–234, 281
110, 113–114, 120, 130, 138–139, 141, Biomes, 11, 13–15, 19, 21, 23–26, 78
155–156, 158, 161, 224, 227–229, 238, Biosphere reserve, 176, 184, 188
295, 328 Blooming desert, 211
Antofagasta, 139, 144, 166, 184, 201 Bofedal, 37, 182
Antuco (volcano), 8, 72, 94, 222–223 Bonpland, A., 51
Apomorphic, 251 Botrychiopsis, 14
Area cladogram, 115, 255 Broad-leaved forest, 38, 272
Area of endemism, 144, 200, 202, 233 Brundin, L., 116, 136
Areagram, 115, 255
Area homology, 284 C
Arealkunde, 87 Cabo de Hornos [Cape Horn], 3, 38, 138, 159,
Asterids, 74–75 188, 230, 304
Atacama Desert, 5, 24, 33, 37, 77, 112, Cabrera, A. L., 130, 142, 146, 225, 236, 242
210–213, 231, 237 Cactoideae, 198, 200–201, 203, 214–216
329
330 General Index
Cain, S., 114, 120 Coring site, 28

Camanchaca, 211 Cosmopolitan (floristic element), 106–109
Cambrian, 12, 66 Cretaceous, 11–12, 16–20, 66–67, 73–75, 78,
Campanian, 19, 169, 257, 261, 263 169, 214–215, 234, 257–261, 263, 273,
Campanulids, 74, 77 278–280
Campos de Hielo, 5, 29, 31, 34 Croizat, L., 78, 89, 113–115, 214, 269, 274,
Cape Floristic Region, 63–64, 137 276–277
Carboniferous, 11–14, 66–67, 70, 78 Cycads, 15–16, 70
Carrán (volcano), 8 Cyttaria, 251, 262–263
Cenozoic, 9, 11–12, 20–31, 66–67, 73–74,
111, 113, 117–118, 167–168, 216, 239, D
257–258, 261, 276 Darwin, Ch., 49–50, 120
Central Chile, 19, 22–27, 29–30, 32–33, de Bibar, G., 47
37–38, 50, 55, 75–76, 95, 98, 101, de Candolle, A., 131–132
112–113, 117, 136, 138–139, 144, de Candolle, A. P., 144
146–147, 166, 181, 183, 185, 190, de Ovalle, A., 48
200–201, 213, 224, 229–231, 237, de Rosales, D., 48
240–241 Deciduous forests, 34–35, 38–39, 138, 181
Central depression, 7, 29–31, 35, 38, 183 Deconstruction, 271, 283
Centres of origin, 114–117, 214, 260–261 Desert, 3, 5, 13–15, 19, 21, 23–24, 26, 33–35,
Cerro Centinela, 27 37, 53, 77, 100, 112–113, 176, 208,
Cerro La Campana, 36, 50, 60 210–213, 231, 237
Cerro Las Vizcachas, 5 Desierto florido, 37, 211
Chaco Domain, 215 Desventuradas Islands, 3, 58, 157–158, 169,
Chaitén (volcano), 8–9 225, 238, 322
Chile Rise, 6 Devonian, 12, 66–67
Chile Triple Junction, 6 Dicots, 21, 57–58, 74–75, 119, 156, 158, 162,
Chiloé, 7–8, 28, 30, 38, 50, 72, 144 216
Chorology, 87, 284 Dicroidium, 15, 70
Cipreses (Cajón de), 187 Diels, L., 57, 131, 135–136, 141
Circum-Pacific ring of fire, 7 Disjunct, 22, 30, 35, 89, 91, 95, 98, 100–101,
Cladistic(s), 57, 65, 114–115, 169, 281, 285 109–118, 137, 224–225, 236, 239, 255,
Cladograms, 68, 74, 115, 145, 207, 215, 235, 260–261, 275–276
255, 260 Disjunction
Climate change, 12–13, 111, 186, 211 antitropical, 113, 118
Clubmosses, 67–68 bipolar, 121
Coastal cordillera, 7, 26, 28, 30, 38, 73, 189, Dispersal, 114–122, 134–135, 159, 166, 168,
211 170, 214–215, 231, 237–239, 242,
Comparative biogeography, 87, 282, 284 260–263, 272, 274–276, 279–282,
Concón, 5, 62, 97, 103, 107, 109, 183 293–294
Conifer forests, 36, 38 Dispersal capacity, 238–239, 261–262
Conifers, 15–16, 20, 22, 30, 34, 36–38, 66, Dispersalism, 114, 270, 275
70–72, 101, 117, 210, 258–259 Dispersalist biogeography, 114–115
Consensus area, 145–147, 207, 233 Domeyko, I., 51
Conservation biogeography, 181–190 Dominion (floristic), 130, 142
Continent of Ice, 3 Drake Passage, 16–17, 259
Continental drift, 11, 14, 274 Drude, O., 131–135, 138, 141
Copiapó, 8, 50, 68, 70 Dry Diagonal, 111–113
Coquimbo, 139–140, 145–146, 166, 184–185,
209, 231, 240 E
Cordaites, 13 Early Eocene Climatic Optimum, 20–21, 23
Cordillera de Darwin, 5, 28–29 Earthquakes, 7
Cordillera Pelada, 10 Easter Island, 3, 53–54, 153–154
General Index 331
El Niño Southern Oscillation [ENSO], 211, Ginkgoales, 70

213 Ginkgos, 15, 18, 70
Endemism, 30, 63–65, 111, 132, 144–147, Glaciar Los Perros, 5
154, 158, 168, 190, 200, 202–209, 221, Glaciers, 28, 30–31, 182, 186
230–231, 233, 239, 281 Glaciation, 9, 13, 21, 23, 27–31, 78, 112, 237,
Endemic (floristic element), 89–90, 103, 106, 258–259
138–139, 157, 225 Glossopteris, 13–115
Engler, A., 57, 87, 113, 130, 133–136, 139, Gnetales, 15–16, 66, 70–71
141 Goethean approach to science, 283
Eocene, 11, 16, 20–23, 111, 117, 234, Gomortegoxylon, 116
237–238, 257–260, 276, 278 Gondwana, 11–17, 19, 73, 93, 113, 121, 137,
Espinal, 38 210, 214–215, 234, 238, 257–261,
Eudicots, 74–75, 119 274–276, 279–282
Euphyllophytes, 66, 68 Gondwanan element, 93
Evolutionary biogeography, 234–239, 281–284 Good, R., 87, 89, 106, 131, 135
Expanding Earth, 275, 278 Grisebach, A., 131, 133, 139
Gymnosperms, 13, 15, 19, 21, 57–58, 67–70,
F 78
Fabids, 74, 76
Falkland Islands, 137
Fern allies, 57–58, 67–68, 107 H
Fernandezian flora, 159–161, 168–171, 176 Hennig, W., 285
Ferns, 14–16, 20, 53, 57–58, 63–64, 66–69, 79, Heusser, C. J., 27–30, 32, 112
113, 120, 135, 154, 156, 162, 164, 170, Hierarchical classification, 129
210, 272 Holantarctic, 131, 136
Filmy ferns, 69 Holocene, 27, 31, 211
Fjords, 5, 9–10, 35, 39 Hooker, J. D., 49, 60, 115, 133, 250, 260, 273
Flat-slab zone, 8 Humboldt, A. v., 32–33, 51–52, 112, 133, 210,
Flora de Chile, 47, 52, 55–56 270, 283–284
Floristic, 11–14, 27, 33, 37, 49, 63–65, 87–109, Humboldt current, 32–33, 112, 210
113, 120–121, 130–144, 155–158, Hyperarid, 25–26, 35, 112, 210
161–168, 176, 223–229, 231–234, 239,
273–274, 276, 278 I
elements, 91–92, 95–109, 130, 139, Illapel, 4, 100, 199
156–158, 161–164 Interandean arid track, 210
regions, 12, 63–64 Intertropical Convergence Zone (ITCZ), 32
Forests, 28, 34–39, 75, 110, 112–113, 138, Invasive species, 173, 176, 241
144–146, 166, 170, 172–174, 182, 184, Isla de Pascua, 4, 58, 68, 95, 153–155
190, 213, 215, 240, 258, 272, 293 Isla Masafuera, 159
Fray Jorge forest, 112–113, 144, 146, 213 Isla Masatierra, 159
Fuentes, F., 53, 154–155 Island biogeography, 181
Islas Desventuradas, 4, 157–159
G
Islas Diego Ramírez, 4
Gay, C., 51, 159, 182
Isla Salas y Gómez, 4
Generalized tracks, 89, 93, 95, 99, 109–110,
Isla San Ambrosio, 157
167, 223–225, 227, 229
Geobotany, 47 Isla San Félix, 157
Geodispersal, 281
Geographic Information Systems (GIS), 33, J
188, 242 Jaccard similarity, 139–140, 166
Géographie botanique, 131 Johnston, I. M., 113–114, 157
Geoparks, 188 Johow, F., 52, 173
Geosystems, 186, 283–284 Juan Fernández (archipelago, islands), 3–4, 69,
Geodiversity, 111 159–176, 225
332 General Index
Juan Fernández Ridge, 8, 159, 167 Middle Miocene, 21, 23–24, 210, 215, 237,
Jurassic, 9, 12, 15–18, 66–67, 70, 73, 78–79, 259
210, 276, 278–279 Mid-Miocene climatic optimum, 259
Miocene, 8, 21, 23–26, 110–113, 116–117,
K 159, 169, 210–211, 215, 234–235, 237,
Kerguelen Islands, 49, 216 257, 259–260, 273
Kingdom (floristic), 133, 135 Molecular dating, 66, 73, 78, 119, 263
Kontingente, 88–89, 138 Molina, J. I., 48, 130
Köppen, W., 33, 135 Monilophytes, 66–69
K/T boundary, 20 Monocots, 20, 53, 57–58, 74–75, 78, 156, 158,
163
L Monophyletic, 66, 68, 70, 78, 115, 252–253
La Campana (cerro), 36, 50, 60 Monte Tronador, 31
Laguna del Hunco, 22
Morphostructural zones, 10
Laguna del Inca, 5
Muñoz Pizarro, C., 50, 56–57, 159, 182, 224
Lamiids, 74, 77
Muñoz-Schick, M., 48, 52, 55, 75, 98, 185,
Land-bridges, 114, 260, 276–277
211, 213, 226–227, 229, 239–241, 263
Láscar (volcano), 9
Museo Nacional de Historia Natural, 51–52,
Last Glacial Maximum (LGM), 28, 31
54–56
Late Oligocene Warming, 21, 23
La Ternera formation, 68–70
Laurifolious forest, 30, 38 N
Lepidodendron, 13, 67 Nahuelbuta (cordillera de), 5, 10, 28–30, 54,
Leptosporangiate ferns, 66, 68–69 72–73
Linnean shortfall, 189 National Herbarium, 29, 53, 55, 107, 200, 202,
Llanquihue glaciation, 27, 29 207, 229
Lomas formation, 37 National Park, 147, 157, 176, 182, 184–185,
Long-distance dispersal (LDD), 114, 117–122, 202
159, 166, 168, 170, 199, 214, 238–239, Nazca Plate, 6–7, 153, 159
261, 272, 276, 281, 293–294 NDM/VNDM optimality method, 144–146,
Looser, G., 53 207–209, 230–231
Los Molles, 70, 100 Neogene, 111, 258–259
Lycophytes, 15, 66–68
Neotropical (floristic element), 95–98
Lycopod trees, 13
Neotropics, 22, 91, 226, 308
Lycopsids, 13, 67
Nevados de Chillán, 9
M New biogeography, 282–283
Maastrichtian, 19–20, 169, 257 New Caledonia, 27, 121, 137, 253, 255,
Macrobioclimates, 33, 35, 38 260–261, 274, 297, 300
Magallanes, 9, 20, 35, 47, 55, 100, 108, New Guinea, 27, 95, 137, 253–254, 257, 261,
138–141, 144, 166–167, 184, 202, 231, 274–275, 295–296, 299–301, 312, 323
234, 297 New Zealand, 10, 22, 49, 63–64, 93, 95, 98,
Maihuenioideae, 198, 201, 205, 214 116, 118–119, 121–122, 130, 133,
Main massings, 95, 139, 261, 263 135–137, 139, 170, 216, 234, 238, 253,
Malesia, 27, 95, 118, 135, 137, 296–297 255, 257–258, 260–263, 271–275, 278
Malvids, 74, 76 Niche modeling, 242–243
Malvinas Islands, 49, 131, 142, 167 Norte Chico, 8
Matorral, 34–38, 182 Norte Grande, 52
sclerophyllous, 34–35 Northpatagonian forests, 28, 34, 38
Maule (region), 183 Nothofagoxylon, 257
Maulino forest, 34, 38 Nothofagus, 249–263
Mediterranean-type climate, 37–38, 112 alessandrii, 38, 190, 251, 253–254, 256,
Mejillones, 4 261, 263
Mesozoic, 7, 14–20, 73, 78, 210, 239, 271 Nunatak, 31
General Index 333
O Province (floristic), 20, 27, 68–69, 130, 133,

Ojos del Salado (volcán) 4 138, 142–147, 215, 281
Oligocene, 17, 21, 23–26, 110–111, 116–117, Pteridophyta [Pteridophytes], 22, 57, 67–68
217, 235, 256, 258–261, 272–273, 276 Pteridosperms, 13
Optimality method (NDM/VNDM) 144, 146,
202, 207, 231 Q
Opuntioideae, 198, 201, 205, 214–216 Quaternary, 7, 12, 27, 30–31
Orchids, 75, 120, 262 Queñoales, 37
Ordovician, 12, 66 Quillworts, 67–68
P
R
Pacifica continent, 167, 276, 278
Rapa Nui, 4, 6, 69, 153–157
Pacific Ocean, 3, 21, 278
Realm (floristic), 93, 132, 134–142, 278
Parsimony Analysis of Endemicity (PAE),
Refugia, 22, 30–31, 263
144–145, 202, 206–208, 231
Region (floristic)
Palaeocene, 16, 20–22, 111, 234, 238, 257, 276
Antarctic, 133
Palaeoclimate, 25
neotropical, 136
Palaeoflora
Red lists, 185, 209, 239
mixed, 22, 24, 113
Reiche, C., 33, 47, 52–54, 56–57, 88–89, 103,
neotropical, 20
138, 141–142, 144, 146–147, 231–234
subantarctic, 22–24
Richness, 20, 22, 61, 67, 69, 139, 157, 190,
tropical, 20, 24
201–202, 209, 223, 230–231, 234,
Palaeogene, 12, 22, 38, 111, 113, 215, 239, 263
239–240, 243, 255–256
Palaeozoic, 7, 9, 11–12, 14
Rosids, 74–76
Panbiogeography, 115, 281
Ruiz, H., 48, 49, 73
Pangaea, 12, 15, 276
Pantropical (floristic element), 91–92
Pan-Gondwana model, 279 S
Paraphyletic, 66–68 Samafrica model, 279
Parinacota (volcano, region), 9, 184 Sanmiguelia, 78
Parsimony Analysis of Endemism (PAE), Santa Juana formation, 70
144–145, 197, 202, 206–208, 231 Schmithüsen, J., 33–35, 39, 138–139
Paschalococos, 154–155 Schouw, J. F., 131–132
Patagonia, 10–12, 20, 22, 24, 26, 28, 30, Sclerophyllous vegetation, 25, 34–36, 38–39,
39, 54–55, 101, 103, 112, 145, 169, 76, 112–113, 138, 181, 240
197–198, 201, 234–236, 299, 303–304, Seed ferns, 15, 20, 79
307, 310, 315–318 Seed plants, 13, 66, 68–69, 78
Pavón, J., 48, 49 Sigillaria, 13
Permian, 12–15, 66–67, 78, 258 Silurian, 12, 66–67
Peru-Chile trench, 6 Similarity analysis, 139–140, 165
Philippi, F., 52, 56–57, 113, 157 Skottsberg, C., 113, 136, 141–142, 154–155,
Philippi, R. A., 51–52, 157, 159 157–160, 168, 170, 174
Phylogeography, 242, 281–282 South American Plate, 6
Pisano, E., 33, 113, 181 South Pacific, 32, 92–93, 110, 254
Pissis, A., 51 South Pole, 49, 54
Pliocene, 25–26, 111–112, 210, 237, 257, 259 South-temperate (floristic element), 101–102
Poeppig, E., 49 Southern Cone Checklist, 57, 101, 198–199,
Polyphyletic, 78 202, 222
Polytopic origin, 114 Spermatophytes, 68–70
Postmodern biogeography, 270–273 Sphenopsids, 13, 16, 210
Precordillera, 8, 37 Spikemosses, 67–68
Protected áreas, 182, 184–185, 188–189, 239 Subtropical Anticyclone, 32
Proterozoic, 12 Synapomorphies, 78, 145, 207
334 General Index
Systematic biogeography, 281–282, 284 V

Systematic conservation planning, 181, Valdivian forest, 35, 38, 113, 145–146, 272
188–189 Valle de la Luna, 5
Vascular plants, 12, 55, 57, 66–70, 140, 259
T Vegetation formations, 20, 33–39
Taiquemó, 28 Vicariance, 114–122, 214–215, 260–261, 270,
Takhtajan, A., 78, 87, 130–131, 135–136 274–276, 279–281
Tamarugal, 37, 182 Vicariance biogeography, 115–117, 276
Tasmania, 20, 22, 49, 93, 95, 116–118, 125, Vicariant form-making, 239
135–137, 252–253, 255, 257, 259, 262, Vicariant speciation, 114, 213
275, 300–301, 312 Volcanic hotspot, 168
Tierra de Juan Fernández, 167–168
Volcanic zone, 6–7
Tierra del Fuego, 3, 9, 19–20, 23, 35, 39, 49,
Volcanoes, 7, 9
130, 231, 317
von Humboldt, A., 51–52, 133, 270, 283
TimeTrek models, 11, 16
von Ihering, H., 115, 121, 276–277
Torres del Paine, 5, 202
Track analysis, 89
Tracks (biogeographic), 89, 109–110, 142, W
167, 210, 229, 275 Wallacean shortfall, 189, 209, 242
Transoceanic dispersal, 117, 261 Wegener, A., 274
Trans-Pacific track, 275 Winclada, 206–207
Tree-ferns, 14, 69, 170 Woody savanna, 24–26, 34, 38
Treviranus, G., 98–99, 130–131
Triassic, 12, 14–15, 20, 66–70, 78, 210, 276
Triple Junction, 6, 276 Z
Tropic of Capricorn, 33 Zöllner, O., 55
Vascular Chilean Plant Genera Index
A Anemone, 312
Aa, 304 Anisomeria, 104, 319
Abrotanella, 95, 124, 162, 225, 227, 237, 238, Antennaria, 99, 227, 312
239, 300 Anthochloa, 301
Abutilon, 295 Anthoxanthum, 322
Acacia, 27, 38, 91, 273, 295 Antidaphne, 307
Acaena, 101, 163, 312 Aphanes, 108, 322
Acantholippia, 37, 98, 301 Aphyllocladus, 227, 301
Achyrocline, 225, 227, 295 Apium, 47, 108, 156, 162, 322
Acrisione, 146, 225, 228, 316 Apodasmia, 300
Adenocaulon, 100, 224, 227, 231, 312 Arachnitis, 301
Adenopeltis, 103, 104, 105, 146, 319 Araeoandra, 104, 319
Adesmia, 37, 61, 62, 63, 76, 89, 305 Araucaria, 15, 21, 27, 30, 36, 38, 48, 70, 71,
Adiantum, 53, 107, 162, 322 72, 73, 95, 111, 137, 258, 273, 300
Aextoxicon, 38, 64, 102, 103, 113, 190, 316 Arenaria, 313
Agalinis, 77, 101, 309 Argemone, 307
Ageratina, 95, 110, 227, 233, 307 Argylia, 301
Agoseris, 97, 227, 309 Argyrochosma, 307
Agrostis, 155, 156, 164, 312 Aristeguietia, 227, 304
Allionia, 310 Aristida, 322
Alonsoa, 77, 91, 298 Aristolochia, 73, 92, 169, 295
Alopecurus, 312 Aristotelia, 172, 173, 298
Aloysia, 307 Arjona, 305
Alstroemeria, 63, 75, 96, 110, 117, 305 Armeria, 313
Alternanthera, 295 Artemisia, 99, 227, 231, 313
Amaranthus, 322 Arthropteris, 58, 95, 164, 168, 172, 300
Amblyopappus, 100, 227, 231, 310 Asplenium, 108, 113, 156, 162, 322
Ambrosia, 227, 240, 307 Astelia, 30, 39, 137, 300
Amelichloa, 312 Asteranthera, 316
Amomyrtus, 316 Asteriscium, 307
Amphibromus, 298 Astragalus, 61, 63, 76, 100, 127, 313
Amphiscirpus, 310 Astrolepis, 307
Amsinckia, 310 Atriplex, 37, 158, 322
Anagallis, 322 Austrocactus, 197, 201–202, 316
Anaphalis, 227, 233, 312 Austrocedrus, 30, 31, 35, 37, 71, 72, 73, 101,
Anarthrophyllum, 61, 316 117, 316
Anatherostipa, 301 Avellanita, 104, 146, 319
Andeimalva, 301 Axonopus, 58, 155, 156, 295
Androsace, 99, 100, 312 Azara, 96, 98, 110, 113, 163, 305
335
336 Vascular Chilean Plant Genera Index
Azolla, 322 Callitriche, 323

Azorella, 36, 37, 95, 298 Calopappus, 104, 225, 228, 230, 233, 319
Calotheca, 305
B Caltha, 88, 121, 127, 313
Baccharis, 37, 63, 95, 223, 224, 227, 230, 231, Calycera, 77, 225, 234, 235, 305
233, 307 Calydorea, 97, 307
Bacopa, 91, 295 Calystegia, 154, 162, 323
Bahia, 100, 224, 228, 310 Camissonia, 101, 310
Bakerolimon, 104, 146, 319 Campsidium, 316
Balbisia, 37, 98, 301 Cardamine, 47, 162, 313
Balsamocarpon, 61, 103, 104, 105, 146, 319 Cardionema, 310
Barneoudia, 316 Carex, 61, 62, 63, 163, 323
Bartsia, 77, 313 Carpha, 295
Beilschmiedia, 38, 91, 295 Carpobrotus, 101, 312
Belloa, 225, 228, 233, 316 Castilleja, 77, 313
Benthamiella, 316 Catabrosa, 313
Berberidopsis, 64, 93, 94, 137, 190, 299 Cenchrus, 323
Berberis, 162, 313 Centaurea, 228, 229, 240, 241, 323
Bidens, 108, 228, 229, 240, 241, 323 Centaurium, 323
Bipinnula, 316 Centaurodendron, 58, 162, 225, 228, 321
Blechnum, 162, 323 Centella, 162, 295
Blennosperma, 101, 224, 228, 233, 239, 310 Centipeda, 225, 227, 239, 295
Blepharocalyx, 98, 304
Cerastium, 323
Blumenbachia, 305
Ceratophyllum, 75, 107, 108, 323
Boehmeria, 58, 163, 295
Cestrum, 307
Bolax, 316
Chaetanthera, 63, 222, 223, 227, 237, 301
Bomarea, 110, 111, 307
Chaptalia, 97, 227, 231, 233, 307
Boopis, 234, 235, 305
Chascolytrum, 310
Boquila, 101, 316
Cheilanthes, 323
Bothriochloa, 323
Chenopodium, 108, 158, 162, 171, 323
Botrychium, 68, 100, 313
Chersodoma, 227, 233, 301
Bouteloua, 95, 307
Bowlesia, 307 Chevreulia, 227, 233, 305
Brachyclados, 228, 233, 316 Chiliotrichum, 102, 222, 228, 316
Brachystele, 305 Chiropetalum, 310
Bridgesia, 104, 106, 146, 319 Chloraea, 63, 97, 110, 305
Bromidium, 310 Chloris, 108, 323
Bromus, 100, 164, 313 Chorizanthe, 310
Browningia, 35, 37, 199, 201, 203, 207, 208, Chrysosplenium, 100, 313
301 Chuquiraga, 97, 213, 223, 227, 236, 237, 304
Bryantiella, 310 Chusquea, 60, 97, 164, 307
Buddleja, 77, 295 Cicendia, 313
Bulbostylis, 295 Ciclospermum, 305
Bulnesia, 37, 304 Cissarobryon, 104, 319
Cissus, 76, 91, 110, 295
C Cistanthe, 37, 101, 310
Caesalpinia, 154, 295 Citronella, 77, 95, 300
Caiophora, 97, 305 Clarkia, 100, 310
Calandrinia, 95, 307 Cleome, 110, 295
Calceolaria, 59, 60, 61, 62, 63, 77, 93, 95, 96, Clinanthus, 301
97, 137, 299, 307 Clinopodium, 323
Caldcluvia, 300 Cliococca, 305
Calliandra, 295 Codonorchis, 306
Callianthemoides, 316 Colletia, 97, 163, 306
Vascular Chilean Plant Genera Index 337
Colliguaja, 37, 98, 306 Deuterocohnia, 306

Collomia, 113, 310 Deyeuxia, 314
Colobanthus, 3, 95, 259, 300 Dichanthelium, 308
Combera, 316 Dichondra, 162, 296
Conanthera, 75, 104, 319 Dicksonia, 15, 27, 58, 69, 95, 162, 164, 172,
Convolvulus, 323 300, 308
Conyza, 91, 227, 230, 240, 295 Dicliptera, 296
Copiapoa, 101, 104, 146, 200, 201, 203, 208, Dielsiochloa, 302
319 Digitaria, 324
Coprosma, 58, 93, 95, 118, 163, 168, 300 Dinemagonum, 76, 104, 146, 320
Cordia, 21, 295 Dinemandra, 76, 104, 320
Coreopsis, 95, 227, 233, 308 Dioscorea, 63, 75, 91, 104, 296, 320
Coriaria, 110, 323 Diostea, 316
Corrigiola, 323 Diplazium, 58, 69, 156, 324
Corryocactus, 201, 203, 208, 301 Diplolepis, 317
Cortaderia, 60, 95, 299 Diplostephium, 37, 227, 231, 233, 304
Corynabutilon, 316 Diposis, 317
Cotula, 91, 227, 233, 240, 296 Discaria, 299
Crassula, 323 Dissanthelium, 310
Cremolobus, 96, 98, 304 Distichia, 304
Cressa, 296 Distichlis, 101, 312
Crinodendron, 38, 111, 306 Dodonaea, 91, 296
Cristaria, 158, 212, 302 Domeykoa, 302
Croton, 92, 110, 296 Donatia, 30, 39, 77, 137, 299
Cruckshanksia, 98, 103, 307 Doniophyton, 228, 236, 317
Cryptantha, 63, 310 Doodia, 58, 95, 154, 155, 156, 300
Cryptocarya, 38, 91, 110, 296 Downingia, 310
Cryptogramma, 313 Draba, 313
Cuatrecasasiella, 227, 304 Drapetes, 29, 30, 101, 317
Cuminia, 58, 103, 163, 321 Drimys, 38, 47, 163, 170, 172, 308
Cumulopuntia, 200, 201, 205, 302 Drosera, 324
Cuscuta, 323 Drymaria, 296
Cynanchum, 323 Dryopteris, 58, 156, 324
Cynodon, 296 Dunalia, 98, 304
Cynoglossum, 323 Dysopsis, 98, 163, 172, 304
Cyperus, 107, 163, 323
Cyphocarpus, 77, 104, 105, 319 E
Cystopteris, 100, 162, 313 Eccremocarpus, 98, 302
Echinodorus, 308
D Elaphoglossum, 156, 162, 296
Dalea, 95, 110, 308 Elatine, 313
Danthonia, 164, 313 Eleocharis, 163, 324
Dasyphyllum, 38, 98, 223, 227, 231, 233, 236, Elodea, 308
239, 240, 306 Elymus, 324
Daucus, 324 Elytropus, 317
Davallia, 58, 69, 154, 156, 296 Embothrium, 94, 101, 102, 121, 317
Dendroseris, 58, 101, 103, 162, 169, 225, 228, Empetrum, 113, 163, 171, 313
322 Encelia, 228, 310
Dennstaedtia, 91, 296 Enneapogon, 324
Deschampsia, 3, 60, 259, 313 Ephedra, 37, 70, 313
Descurainia, 313 Epilobium, 313
Desfontainia, 77, 96, 98, 304 Epipetrum, 104, 146, 320
Desmaria, 104, 319 Equisetum, 68, 69, 324
Eragrostis, 158, 324 Geoffroea, 61, 97, 304

Ercilla, 104, 320 Geranium, 324
Erechtites, 227, 233, 308 Gethyum, 104, 320
Eremocharis, 302 Geum, 107, 108, 324
Eremodraba, 96, 302 Gevuina, 121, 300
Eriachaenium, 101, 225, 228, 317 Gilia, 101, 310
Erigeron, 162, 227, 313 Gilliesia, 317
Eriochloa, 324 Glandularia, 310
Eriosyce, 104, 146, 199, 200, 201, 203, 208, Glinus, 91, 296
209, 320 Glyceria, 108, 324
Erodium, 324 Glycyrrhiza, 314
Errazurizia, 101, 113, 310 Gnaphalium, 107, 108, 228, 229, 324
Eryngium, 108, 118, 162, 171, 324 Gochnatia, 101, 224, 228, 233, 312
Escallonia, 22, 74, 77, 96, 97, 98, 104, 137, Gomortega, 64, 73, 104, 107, 116, 190, 320
163, 165, 304, 319, 321 Gomphrena, 91, 296
Eucryphia, 22, 38, 94, 95, 258, 299 Grabowskia, 308
Eudema, 96, 304 Grammitis, 162, 296
Eulychnia, 37, 104, 199, 200, 201, 203, 208, Gratiola, 324
209, 320 Greigia, 163, 308
Euphorbia, 37, 109, 144, 324 Grindelia, 227, 308
Euphrasia, 77, 99, 113, 163, 314 Griselinia, 77, 113, 137, 299
Evolvulus, 296 Guindilia, 37, 317
Exodeconus, 98, 304 Gunnera, 19, 75, 121, 163, 165, 273, 296
Gutierrezia, 224, 228, 310
F Guynesomia, 104, 228, 229, 233, 320
Fabiana, 37, 302 Gymnachne, 104, 146, 320
Facelis, 227, 306 Gymnophyton, 307
Fagara, 58, 163, 298 Gypothamnium, 104, 228, 229, 233, 237, 320
Fagonia, 100, 119, 314
Famatina, 317
Fascicularia, 104, 105, 146, 320 H
Festuca, 37, 39, 60, 324 Haageocereus, 201, 203, 208, 302
Ficinia, 95, 300 Habenaria, 296
Fitzroya, 30, 31, 35, 38, 71, 72, 73, 101, 182, Habranthus, 308
185, 190, 317 Halerpestes, 314
Flaveria, 101, 224, 228, 239, 312 Haloragis, 58, 95, 163, 168, 299
Flourensia, 37, 228, 233, 310 Hamadryas, 317
Fonkia, 317 Haplopappus, 37, 61, 63, 222, 223, 227, 302
Francoa, 64, 65, 104, 105, 146, 320 Haplorhus, 302
Frankenia, 99, 158, 324 Hebe, 93, 95, 300
Fuchsia, 94, 95, 299 Helenium, 227, 308
Fuertesimalva, 158, 308 Heliotropium, 37, 213, 324
Helogyne, 227, 233, 302
G Herbertia, 164, 306
Gaimardia, 300 Herreria, 306
Galium, 163, 324 Heterosperma, 227, 233, 308
Gamocarpha, 234, 317 Heterozostera, 299
Gamochaeta, 95, 162, 227, 230, 308 Hieracium, 227, 240, 241, 314
Gamochaetopsis, 228, 317 Hippuris, 314
Gaultheria, 101, 163, 312 Histiopteris, 162, 296
Gavilea, 164, 317 Hoffmannseggia, 99, 101, 114, 117, 118, 120,
Gayophytum, 310 127, 310
Gentiana, 118, 314 Hollermayera, 104, 320
Gentianella, 118, 324 Homalocarpus, 104, 320
Hordeum, 60, 314 Lasthenia, 228, 239, 311

Huanaca, 317 Lathyrus, 314
Huidobria, 103, 104, 105, 320 Latua, 104, 320
Huperzia, 29, 30, 67, 68, 107, 108, 324 Laurelia, 19, 38, 56, 73, 116, 121, 299
Hybanthus, 296 Laureliopsis, 38, 101, 102, 116, 317
Hydrangea, 314 Lebetanthus, 317
Hydrocotyle, 108, 325 Lecanophora, 101, 317
Hymenoglossum, 69, 162, 317 Legenere, 311
Hymenophyllum, 69, 113, 162, 325 Legrandia, 104, 190, 320
Hypericum, 108, 325 Lemna, 325
Hypochaeris, 31, 100, 118, 224, 227, 237, 238, Lenzia, 98, 307
240, 314 Leontochir, 103, 104, 105, 106, 146, 320
Hypolepis, 162, 296 Lepechinia, 308
Lepidium, 118, 158, 325
I Lepidoceras, 302
Imperata, 325 Lepidophyllum, 101, 228, 317
Ipomoea, 58, 156, 296 Lepidothamnus, 30, 71, 73, 299
Ipomopsis, 101, 311 Leptinella, 93, 225, 227, 239, 301
Islaya, 201, 203, 302 Leptocarpha, 104, 228, 229, 230, 320
Isoetes, 67, 68, 107, 325 Leptochloa, 297
Isolepis, 314 Leptophyllochloa, 164, 317
Ivania, 104, 320 Leptosiphon, 101, 311
Leptostigma, 301
J
Lepuropetalon, 311
Jaborosa, 306
Leucheria, 63, 223, 227, 230, 231, 237, 302
Jarava, 37, 308
Leucocoryne, 101, 104, 320
Jovellana, 77, 93, 95, 299
Leunisia, 104, 225, 228, 230, 320
Juania, 58, 103, 161, 163, 322
Leymus, 314
Jubaea, 48, 104, 107, 146, 155, 320
Libertia, 164, 301
Juncus, 107, 156, 164, 325
Ligaria, 306
Junellia, 302
Lilaea, 311
K Lilaeopsis, 101, 312
Kageneckia, 37, 98, 302 Limonium, 314
Koeleria, 325 Limosella, 77, 325
Krameria, 76, 308 Lindernia, 77, 325
Kurzamra, 98, 307 Linum, 325
Kyllinga, 58, 154, 156, 296 Lippia, 91, 297
Lithrea, 111, 306
L Littorella, 314
Lachemilla, 308 Llagunoa, 98, 304
Lactoris, 58, 64, 103, 163, 165, 169, 170, 173, Loasa, 63, 98, 302
190, 322 Lobelia, 77, 91, 92, 162, 297
Laennecia, 227, 302 Lobivia, 201, 204, 302
Lagenophora, 93, 95, 162, 225, 227, 231, 239, Lomatia, 22, 95, 121, 299
301 Lophopappus, 227, 233, 302
Lampaya, 302 Lophosoria, 69, 162, 308
Landoltia, 106, 108, 325 Lucilia, 227, 233, 306
Lapageria, 48, 49, 64, 75, 104, 105, 107, 113, Luciliocline, 227, 233, 304
116, 117, 146, 320 Ludwigia, 325
Lardizabala, 317 Luma, 38, 172, 317
Laretia, 37, 102, 317 Lupinus, 314
Larrea, 99, 100, 113, 122, 311 Luzula, 164, 325
Lastarriaea, 101, 311 Luzuriaga, 75, 95, 117, 137, 299
Lycapsus, 58, 157, 158, 169, 225, 228, 238, Montiopsis, 302
322 Monttea, 98, 307
Lycium, 62, 108, 156, 325 Morella, 297
Lycopodium, 67, 68, 107, 162, 325 Moscharia, 104, 146, 228, 229, 230, 231, 233,
Lysimachia, 325 237, 321
Moschopsis, 234, 303
M Mostacillastrum, 303
Machaerina, 58, 163, 297 Muehlenbeckia, 95, 301
Macrachaenium, 228, 317 Muhlenbergia, 315
Madia, 228, 311 Mulguraea, 307
Maihuenia, 197, 198, 200, 201, 202, 205, 215, Mulinum, 37, 101, 318
216, 317 Munroa, 311
Maihueniopsis, 197, 199, 201, 205, 208, 209, Mutisia, 111, 222, 227, 231, 242, 243, 305
216, 302 Myoschilos, 318
Maireana, 157, 158, 299 Myosotis, 315
Malacothrix, 228, 311 Myosurus, 315
Malesherbia, 63, 64, 76, 96, 213, 302 Myrceugenia, 98, 110, 111, 124, 163, 172, 306
Malvella, 314 Myrcianthes, 98, 305
Mancoa, 99, 101, 311 Myriophyllum, 325
Margyracaena, 58, 163, 171, 322 Myrteola, 97, 98, 163, 305
Margyricarpus, 163, 306
N
Marsilea, 69, 325
Nama, 309
Marsippospermum, 299
Nanodea, 318
Marticorenia, 104, 225, 228, 230, 233, 320
Nardophyllum, 102, 227, 230, 303
Mastigostyla, 302
Nasa, 95, 309
Mathewsia, 302
Nassauvia, 51, 98, 227, 230, 231, 303
Maytenus, 38, 297
Nassella, 95, 164, 309
Mecardonia, 96, 308 Nastanthus, 101, 234, 235, 318
Megalachne, 58, 164, 322 Navarretia, 101, 311
Megalastrum, 91, 162, 297 Neoporteria, 104, 146, 200, 201, 204, 208, 321
Melica, 55, 60, 314 Neowerdermannia, 98, 201, 204, 303
Melosperma, 318 Nertera, 95, 113, 163, 300
Menodora, 98, 316 Nesocaryum, 58, 157, 158, 322
Menonvillea, 318 Neuontobothrys, 303
Mentzelia, 97, 308 Nicandra, 303
Metharme, 104, 320 Nicotiana, 91, 163, 297
Microlepia, 58, 156, 297 Nierembergia, 309
Microphyes, 104, 146, 320 Nitrophila, 311
Micropsis, 227, 233, 306 Noccaea, 315
Microseris, 101, 224, 228, 238, 239, 312 Nolana, 37, 62, 63, 77, 80, 212, 303
Microsorum, 58, 156, 297 Notanthera, 104, 161, 163, 171, 321
Microsteris, 101, 311 Nothofagus, 19, 21, 22, 27, 29, 30, 36, 38, 39,
Miersia, 104, 146, 321 76, 95, 116, 132, 133, 147, 190, 249–263,
Mikania, 91, 225, 227, 233, 297 275, 301
Mimulus, 77, 108, 163, 325 Notholaena, 91, 162, 297
Minuartia, 314 Nothoscordum, 309
Miqueliopuntia, 104, 198, 201, 205, 208, 321 Noticastrum, 227, 233, 306
Mirabilis, 314 Notopappus, 102, 228, 318
Misodendrum, 64, 263, 318 Nototriche, 96, 98, 305
Mitraria, 113, 318
Mniodes, 227, 233, 302 O
Monnina, 308 Ochagavia, 104, 146, 161, 163, 321
Montia, 325 Ochetophila, 318
Ocyroe, 222, 227, 303 Phragmites, 326

Odontorrhynchus, 318 Phrodus, 104, 146, 321
Oenothera, 309 Phycella, 318
Oldenlandia, 163, 297 Phyllachne, 299
Olsynium, 309 Phylloscirpus, 303
Ombrophytum, 49, 305 Phytolacca, 104, 297, 319, 320
Onuris, 318 Picrosia, 227, 306
Ophioglossum, 68, 156, 162, 325 Pilea, 297
Ophryosporus, 222, 227, 306 Pilgerodendron, 30, 38, 71, 73, 101, 318
Oreobolus, 39, 95, 118, 163, 301 Pilostyles, 297
Oreocereus, 98, 200, 201, 204, 208, 303 Pilularia, 326
Oreomyrrhis, 93, 118, 123, 301 Pinguicula, 99, 130, 315
Oreopolus, 318 Pintoa, 104, 146, 321
Orites, 11, 22, 94, 121, 299 Piptochaetium, 97, 164, 309
Orobanche, 77, 325 Pitavia, 103, 104, 105, 190, 321
Ortachne, 318 Pitraea, 303
Osmorhiza, 113, 118, 315 Pityrogramma, 297
Otholobium, 48, 297 Placea, 104, 106, 146, 321
Ourisia, 88, 117, 118, 299 Plagiobothrys, 101, 312
Ovidia, 318 Plantago, 158, 163, 326
Oxalis, 63, 109, 325 Plazia, 227, 231, 240, 303
Oxychloë, 98, 303 Pleocarphus, 104, 106, 146, 224, 228, 229,
Oxyphyllum, 104, 228, 229, 231, 233, 237, 321 233, 321
Oxytheca, 311 Pleopeltis, 162, 297
Oziroe, 303 Pleurophora, 306
Pleurosorus, 326
P Pluchea, 227, 233, 297
Pachylaena, 227, 303 Plumbago, 326
Palaua, 303 Poa, 63, 315
Panicum, 108, 325 Podagrostis, 311
Parastrephia, 37, 227, 233, 303 Podanthus, 104, 228, 229, 233, 321
Parietaria, 158, 163, 325 Podocarpus, 21, 38, 71, 72, 73, 111, 298
Paronychia, 315 Podophorus, 58, 164, 171, 322
Pasithea, 75, 303 Polemonium, 315
Paspalum, 156, 326 Polyachyrus, 213, 227, 231, 237, 303
Passiflora, 64, 76, 91, 92, 110, 297 Polycarpon, 326
Patosia, 303 Polygala, 109, 326
Pectocarya, 311 Polygonum, 326
Pellaea, 297 Polylepis, 37, 97, 213, 305
Pelletiera, 91, 297 Polypogon, 326
Pennellia, 303 Polypsecadium, 305
Pennisetum, 315 Polystichum, 156, 162, 326
Peperomia, 163, 297 Porlieria, 37, 311
Perezia, 98, 111, 227, 230, 231, 233, 237, 345 Portulaca, 108, 215, 326
Perityle, 228, 311 Potamogeton, 326
Persea, 38, 111, 315 Potentilla, 100, 315
Peumus, 19, 38, 73, 104, 105, 116, 146, 321 Pouteria, 91, 298
Phacelia, 101, 113, 311 Pozoa, 318
Phalaris, 315 Primula, 315
Philesia, 64, 102, 103, 116, 117, 318 Prosopis, 37, 38, 120, 298
Philibertia, 98, 303 Proustia, 227, 303
Philippiella, 318 Prumnopitys, 29, 30, 71, 73, 95, 299
Phleum, 315 Psilocarphus, 228, 233, 239, 311
Psilotum, 58, 68, 154, 156, 298 Saxifragodes, 319

Pteris, 162, 326 Schinus, 27, 309
Pterocactus, 197, 199, 201, 205, 216, 318 Schizaea, 16, 69, 298
Puccinellia, 326 Schizanthus, 62, 319
Puya, 36, 37, 38, 209, 309 Schizeilema, 299
Pycnophyllum, 98, 303 Schizopetalon, 103, 307
Pycreus, 58, 156, 298 Schkuhria, 227, 231, 309
Pyrrhocactus, 199, 204, 208, 209, 307 Schoenoplectus, 327
Schoenus, 327
Q Scirpus, 107, 156, 163, 327
Quillaja, 38, 63, 64, 76, 98, 111, 306 Scutellaria, 327
Quinchamalium, 303 Scyphanthus, 104, 107, 321
Selkirkia, 58, 162, 322
R Selliera, 234, 299
Ranunculus, 107, 108, 163, 326 Senecio, 61, 62, 63, 107, 223, 228, 229, 230,
Raukaua, 95, 121, 299 233, 240, 242, 327
Reicheella, 104, 321 Senna, 298
Relchela, 318 Serpyllopsis, 69, 162, 319
Retanilla, 318 Setaria, 327
Reyesia, 307 Sicyos, 101, 158, 312
Rhamnus, 100, 315 Sigesbeckia, 91, 227, 298
Rhaphithamnus, 163, 165, 318 Silene, 108, 327
Rhodophiala, 306 Sisyrinchium, 309
Rhombolytrum, 306 Skytanthus, 306
Rhynchospora, 326 Solanum, 63, 156, 158, 163, 327
Ribes, 99, 100, 130, 315 Solaria, 319
Robinsonia, 58, 101, 162, 169, 171, 225, 228,
Solenomelus, 319
238, 240, 322
Solidago, 227, 315
Rorippa, 118, 326
Soliva, 101, 224, 228, 239, 312
Rostkovia, 299
Sophora, 155, 156, 157, 163, 165, 327
Rostraria, 309
Spartina, 315
Rubus, 107, 163, 172, 173, 326
Speea, 104, 106, 146, 321
Rumex, 326
Spergula, 315
Rumohra, 95, 113, 162, 172, 300
Spergularia, 158, 162, 327
Ruppia, 326
Sphaeralcea, 311
Rytidosperma, 93, 155, 156, 299
Spilanthes, 91, 227, 298
S Sporobolus, 327
Sagina, 326 Stachys, 327
Sagittaria, 106, 326 Stangea, 77, 304
Salix, 99, 130, 315 Stellaria, 327
Salpichroa, 303 Stemodia, 315
Salpiglossis, 62, 318 Stenandrium, 97, 309
Salvia, 108, 109, 326 Stevia, 227, 309
Samolus, 108, 156, 326 Sticherus, 162, 298
Sanctambrosia, 58, 157, 158, 322 Stipa, 58, 156, 327
Sanicula, 110, 327 Stuckenia, 327
Santalum, 58, 95, 163, 168, 171, 173, 174, 301 Suaeda, 158, 327
Sarcocornia, 162, 327 Symphyotrichum, 228, 311
Sarcodraba, 318 Synammia, 162, 319
Sarmienta, 104, 113, 321
Saxegothaea, 71, 73, 101, 318 T
Saxifraga, 100, 315 Tagetes, 227, 309
Saxifragella, 101, 318 Tapeinia, 319
Tarasa, 101, 311 Urmenetea, 98, 227, 304

Taraxacum, 162, 224, 227, 315 Urocarpidium, 304
Tecoma, 298 Urtica, 163, 328
Tecophilaea, 75, 104, 146, 321 Utricularia, 328
Tepualia, 102, 319
Tessaria, 227, 306 V
Tetilla, 64, 104, 146, 321 Vahlodea, 312
Tetrachondra, 119, 299 Valdivia, 77, 104, 321
Tetraglochin, 37, 98, 304 Valeriana, 63, 77, 100, 316
Tetragonia, 37, 158, 315 Vasconcellea, 96, 309
Tetroncium, 319 Verbena, 316
Teucrium, 327 Verbesina, 227, 309
Thamnoseris, 58, 157, 158, 169, 225, 228, 238, Veronica, 328
322 Vestia, 104, 146, 321
Thelocephala, 104, 146, 200, 201, 204, 208, Vicia, 100, 316
321 Viguiera, 227, 309
Thelypteris, 156, 298 Villanova, 227, 309
Thyrsopteris, 58, 64, 69, 162, 164, 170, 322 Viola, 49, 61, 62, 63, 76, 108, 109,
Tigridia, 311 328
Tillandsia, 309 Vittaria, 58, 69, 154, 156, 298
Tiquilia, 101, 113, 114, 117, 118, 311 Viviania, 76, 98, 111, 301, 306, 319
Traubia, 104, 146, 321 Vulpia, 328
Trevoa, 104, 146, 321
Tribeles, 77, 319 W
Trichocereus, 37, 38, 201, 204, 208, 216, 304 Wahlenbergia, 77, 95, 162, 170, 300
Trichocline, 225, 227, 229, 299 Weberbauera, 98, 304
Trichomanes, 162, 298 Wedelia, 227, 298
Trichoneura, 298 Weinmannia, 38, 101, 111, 312
Trichopetalum, 75, 101, 319
Werdermannia, 307
Trifolium, 315
Werneria, 227, 231, 305
Triglochin, 316
Wolffia, 106, 328
Triodanis, 316
Wolffiella, 91, 298
Tripogon, 298
Woodsia, 100, 316
Triptilion, 101, 222, 225, 228, 231, 319
Trisetum, 164, 327
Tristagma, 319 X
Tristerix, 305 Xenophyllum, 227, 233, 305
Triumfetta, 58, 154, 156, 298 Xerodraba, 101, 319
Trixis, 227, 233, 309
Tropaeolum, 309 Y
Tropidocarpum, 99, 101, 312 Yunquea, 58, 162, 169, 225, 228,
Tunilla, 98, 201, 205, 208, 304 322
Tweedia, 97, 98, 306
Typha, 328 Z
Zameioscirpus, 304
U Zannichellia, 328
Ugni, 97, 163, 172, 173, 309 Zephyra, 75, 104, 106, 321
Uncinia, 95, 163, 301 Zoellnerallium, 319

Plant Geography of Chile

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Plant Geography of Chile

Uploaded by

Copyright:

Available Formats

Plant Geography of Chile

PLANT AND VEGETATION

Series Editor: M.J.A. Werger

For further volumes:

ISSN 1875-1318 e-ISSN 1875-1326

© Springer Science+Business Media B.V. 2011

Printed on acid-free paper

Springer is part of Springer Science+Business Media (www.springer.com)

Utrecht, The Nederlands Marinus J.A. Werger

What Is This Book Not About?

What Is This Book Then About?

Structure of the Book

Santiago, Chile Andrés Moreira-Muñoz

at the Pontificia Universidad Católica de Chile, which facilitated my incorporation

1 May 2010 Limache

Part I Geobotanical Scenario

Part II Chorology of Chilean Plants

3.2 To Be or Not To Be Disjunct? . . . . . . . . . . . . . . . . . . 109

Part III Islands Biogeography

Part IV Case Studies on Selected Families

7.5 Cactus Postmodern Evolution . . . . . . . . . . . . . . . . . . 215

Part V Where to from Here? Projections of Chilean Plant Geography

AAO Antarctic Oscillation

Chile has been characterized as “a geographic extravaganza” (Subercaseaux 1940)

A. Moreira-Muñoz, Plant Geography of Chile, Plant and Vegetation 5, 3

1.1 Tectonics and Physiography

Box 1.1 Living Under the Volcano

1.1.1 Morphostructural Macrozones

2,000 Campo de Hielo Sur

Box 1.2 Patagonian Vicissitudes

1.2 Past Climate and Vegetation

COOL WARM COOL WARM COOL WARM COOL WARM COOL

1.2.1 The Palaeozoic (542–251 mya)

By the Late Carboniferous (330–299 mya) the southern flora consisted

Mid latitude desert

1.2.2 The Mesozoic (251–65.5 mya)

to that of today’s Mediterranean-type one. The relatively increased proportion of

Box 1.3 Gondwana Breaks-Up

Major separation Palaeoreconstructions on a TimeTrek

(W Gondwana / During Middle to Late Jurassic

Africa–S Early Cretaceous (119–105

West Early Oligocene (ca 30 mya):

Mid latitude desert

(Fig. 1.14), the latter being characterized by Araucariaceae, Nothofagaceae,

Box 1.4 Floral Extinction at the K/T Boundary ?

1.2.3 The Cenozoic (65.5 mya Onwards)

0 Glaciation Glaciation East Antarctic

et al. 2001). This was followed by a gradual cooling and reestablishment of a

Box 1.5 Slow or Rapid Andean Uplift?

Desert (hyperarid / arid)

Box 1.6 Six Steps for Palaeoenvironmental Reconstruction

5. Once in the laboratory, identify the pollen grains under a microscope, at

LGM ice extension

xeric Amaranthaceae and Asteraceae (Heusser 1997). The presence of Nothofagus

(1) Existence of disjuct populations of conifers (Fitzroya, Araucaria, Prumnopitys,

or had deeper evolutionary implications is still a matter of discussion (Box 1.7).

Box 1.7 How to Survive a Glaciation? The Refugial Debate

1.3 Current Climate and Vegetation

January (southern summer) July (southern winter)

The Humboldt Current is characterised by a predominant northward flow of sur-

1.3.1 Bioclimatic Zones

1.3.2 Vegetation Formations

North Snow limit Upper vegetation limit South

Andean vegetation Desert scrub Valdivian forest North Patagonian rainforest