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91 92 Plant Geography of Chile 93

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103 Andrés Moreira-Muñoz 104 Pontiﬁcia Universidad Católica de Chile, Santiago, Chile 105

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133 134 123 135 UNCORRECTED PROOF 136 Dr. Andrés Moreira-Muñoz

137 Pontiﬁcia Universidad Católica de Chile Instituto de Geograﬁa 138 Av. Vicuna Mackenna 4860 Santiago 139 Chile 140 [email protected] 141

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167 168 ISSN 1875-1318 e-ISSN 1875-1326 169 ISBN 978-90-481-8747-8 e-ISBN 978-90-481-8748-5 170 DOI 10.1007/978-90-481-8748-5

171 Springer Dordrecht Heidelberg London New York

172 Library of Congress Control Number: xxxxx 173

174 © 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 175 any means, electronic, mechanical, photocopying, microﬁlming, recording or otherwise, without written 176 permission from the Publisher, with the exception of any material supplied speciﬁcally for the purpose 177 of being entered and executed on a computer system, for exclusive use by the purchaser of the work.

178 Printed on acid-free paper 179

180 Springer is part of SpringerUNCORRECTED Science+Business Media (www.springer.com) PROOF 01 Foreword 02

12 13 It is not just the brilliant and dramatic scenery that makes Chile such an attractive 14 part of the world. No, that country has so very much more! And certainly it has a 15 rich and beautiful flora. Chile’s plant world is strongly diversified and shows inter- 16 esting geographical and evolutionary patterns. This is due to several factors: The 17 geographical position of the country on the edge of a continental plate and stretch- 18 ing along an extremely long latitudinal gradient from the tropics to the cold, barren 19 rocks of Cape Horn, opposite Antarctica; the strong differences in altitude from sea 20 level to the icy peaks of the Andes; the inclusion of distant islands in the country’s 21 territory; the long geological and evolutionary history of the biota; and the mixture 22 of tropical and temperate floras. 23 The flora and vegetation of Chile already drew the attention of the early adven- 24 turers and explorers and as from the eighteenth century attracted naturalists and 25 collectors from Europe. In the nineteenth century famous botanists explored and 26 studied the Chilean plant world, and gradually the flora and plant geographical pat- 27 terns became subjects of scientific analyses both by European and Chilean scholars. 28 Recently, the development of new scientific techniques have allowed to reveal the 29 remarkable evolutionary pathways in many Chilean plant groups, and have provided 30 clues to the origins of intriguing plant geographical patterns in the southern hemi- 31 sphere floras. This shall be of interest for botanists, plant geographers, ecologists 32 and evolutionary biologists worldwide. 33 I was very lucky to get into contact with Dr. Andrés Moreira-Muñoz. He is an 34 enthusiastic and outstanding Chilean plant scientist with historical roots in this sub- 35 ject area. Dr. Moreira-Muñoz here presents a modern and stimulating account of 36 the Plant Geography of Chile that analyses the floristic diversity and endemism 37 of the country. He interprets the origins of the fascinating plant geographical pat- 38 terns of Chile and explains the evolutionary background of the most important plant 39 groups. I am very pleased to present this book as a volume in the series “Plant and 40 Vegetation” to the international readership. 41 42 Utrecht, The Nederlands Marinus J.A. Werger 43

45 UNCORRECTED PROOF

v 01 Preface 02

12 13 One morning in 1897 at the Quinta Normal, Santiago: the Director of the Museo 14 Nacional de Historia Natural, Federico Philippi welcomes the new German botanist 15 responsible for taken the reins of the botanical section, Dr Carl Reiche. He has 16 been committed to maintain the National Herbarium, promoting exchanges, ana- 17 lyzing, increasing and organizing the collections of the Herbarium. He will be 18 also, and this is not a trivial thing, responsible for writing the new Flora de 19 Chile; and he has already published the first volume. Chilean botanical knowl- 20 edge showed at the end of the nineteenth century still many gaps, in spite of the 21 great achievements of Claudio Gay and R.A. Philippi, this latter the father and 22 mentor of the Museum’s Director. It took Reiche more than 15 years to system- 23 atize, revise and add the necessary information that finally encompassed the six 24 volumesoftheFloradeChile(Chap. 2). In the meantime, when Reiche was 25 already well familiarized with the Chilean flora, he got a request for writing a 26 synthetic book about the Chilean plant geography for the series Die Vegetation 27 der Erde, edited by the great German botanists Adolf Engler and Oscar Drude. 28 Reiche completed the assignment successfully, and 1907 published Grundzüge der 29 Pflanzenverbreitung in Chile, encompassing 222 pages with two maps and sev- 30 eral photographs (Vegetationsbilder). This was the first (and so far the only) Plant 31 Geography of Chile. This great effort, which put the Chilean plant world in a 32 renowned world series, only got a Spanish translation 30 years later, thanks to the 33 engagement of G. Looser, himself a botanist and notable scientific communicator 34 (Chap. 2). 35 Just as Reiche once did with the previous works of Gay and the Philippi, now it 36 seems to be time for a renewal of Reiche’s Plant Geography. No few things have 37 changed in a hundred years: plants have been renamed and reclassified; taxonomy 38 and systematics have suffered far-reaching changes; biology, geography, and bio- 39 geography have undergone paradigmatic vicissitudes. I underwent the challenge 40 of writing a “New Plant Geography of Chile” as a doctoral student in Erlangen, 41 Germany. In such an exponentially dynamic field, one and a half year after the 42 publication of the thesis many things had to be revised and updated for this book. 43 Regarding the subject, the reader may ask why to use the old concept of “plant 44 geography” rather than “phytogeography” or “geobotany”? As these terms are often 45 used indistinctly, I decidedUNCORRECTED to use the oldest term “plant geography”, PROOF honouring

vii viii Preface

46 the seminal works from A. von Humboldt: Géographie des plantes, and A.P. de 47 Candolle’s Géographie Botanique (Chap. 4). The present book also takes inspiration 48 from Stanley Cain’s words in his book Foundations of Plant Geography: “This is 49 not a descriptive plant geography, but rather an inquiry into the foundations of the 50 science of plant geography” (Cain 1944, p xi) (Chap. 3).

53 What Is This Book Not About?

55 This book is not a traditional geobotanical textbook. It rather attempts to enter

56 into the discussion on the challenges that shape (post)modern biogeography in the

57 twenty-ﬁrst century. A detailed vegetation description, which is sometimes mis-

58 understood as a main task of “plant geography”, is very far from the goal of the

59 book. The reader is redirected to recent advances in this speciﬁc ﬁeld (Chap. 1).

60 Many new concepts and methods are currently emerging in biogeography. This

61 book doesn’t offer new conceptual or methodological advances; it rather wants to

62 be a “ﬁeld guide” to the possibilities for the development of the discipline in Chile.

63 Consequently, several conﬂicting approaches that have been proposed for explaining

64 current biogeographic patterns are confronted throughout the text (e.g. vicariance

65 versus dispersal). The result is mostly not deﬁnitive, suggesting that a dichotomy is

66 just a too simple problem design of a much more complex problem.

67 68 What Is This Book Then About? 69 70 The present book intends to reflect the “state of the art” or a synthesis of the plant 71 geographical discipline in Chile. The challenge is seemingly overwhelming, since 72 in such a composite discipline like biogeography, today any intend to integrate the 73 different views that shape it, must confront the differences inherent to the diverse 74 approaches involved in the discipline. To what extent biogeography assumes and 75 reflects the conflicts, assumptions and challenges inherent to (post)modern science 76 must then be kept in mind while analysing the Chilean plant geography. 77 This approach leaves us the theoretical basis and practical lines of direction for 78 the endeavour of doing plant geography in the twenty-first century, in the constantly 79 “changing world” of biogeography (sensu Ebach and Tangney 2007) (Chap. 10). 80 Most efforts at the regional level concentrate rather on the descriptive or on the ana- 81 lytical. I would like to do both and also to present the few results in a more general 82 interpretative framework. I would like to accept the challenge posted by Morrone 83 (2009) (Chap. 10), touching methodological as well as more theoretical aspects that 84 will help the student build an own “road map” towards a future development of the 85 discipline in Chile, integrating methods, data, concepts, and interpretations from 86 different fields. 87 Applying one of the basic principles of geography, for a better comprehension 88 of the subject I have often put the eye beyond the Pacific and beyond the Andes, 89 touching aspects of the New Zealand biota, the Antarctic palaeobiomes, Argentinian 90 UNCORRECTED PROOF Patagonia... I apologize if I have mentioned these aspects in a superficial form. Preface ix

91 Nevertheless, I suspect that several aspects of the book are applicable or of interest 92 for biogeographers in the other (once united) southern hemisphere territories; if so, 93 I will be deeply satisﬁed.

94 95 Structure of the Book 96

97 The book is divided in five parts that organize the different chapters. 98 99 The 1st part presents an overview of the geographical and botanical scenar- 100 ios that shape the Chilean vascular plant world, in the present as well as in the 101 geologic past. In chapter one, the main physical characteristics of the Chilean ter- 102 ritory are briefly exposed, especially the geological and tectonic origins of Chile 103 and their effects on the palaeogeography and the evolution of the Southern Cone 104 biomes. This contributes to a better understanding of the current climate and vege- 105 tation. The 2nd chapter makes a succinct revision of the historical development of 106 Chilean botany, and synthesizes the current knowledge regarding the composition of 107 the flora. 108 The 2nd part deals with Chilean plant geographical relationships, oriented to a 109 synthesis of the floristic elements of the extant flora. The classification of Chilean 110 genera into floristic elements in Chap. 3, will be the basis for the discussion of the 111 disjunct patterns that shape the Chilean flora. This analysis will be further comple- 112 mented with the task undertaken in the 4th chapter, regarding the biogeographical 113 regionalization of the Chilean territory. 114 The 3rd part provides an analysis of two close related subdisciplines: island bio- 115 geography and conservation biogeography. Chapter 5 presents a synthesis of the 116 plant world of the Chilean Pacific offshore islands, emphasizing their uniqueness 117 and threats, while the 6th chapter analyses the fragmentation in the mainland, related 118 to the impacts of human activities on the Chilean ecosystems. Concepts and tools 119 developed within the field of conservation biogeography are analyzed in relation to 120 current global changes. 121 The 4th part moves into the case studies, regarding specific groups that deserve 122 special attention in biogeography. Chapter 7 gets into the biogeography of one 123 of the most charismatic American families, the Cactaceae, of course regarding its 124 Chilean representatives. Chapter 8 turns to another not less interesting family, the 125 Asteraceae, the most genus/species-rich family in Chile. The last case study is pre- 126 sented in Chap. 9, devoted to a monogeneric family also called the “key genus in 127 plant geography”: Nothofagus. 128 The 5th and last part of the book announces several ways in which Chilean plant 129 geography can further develop; maybe more rapidly and effectively than during the 130 last 100 years? Chapter 10 is in this sense rather speculative, in an attempt to put 131 Chilean plant geography in a more general context of modern biogeography. Finally, 132 the 11th chapter only adds several digressions about the scientific endeavour and the 133 artificial distinction between nature and culture. 134 135 Santiago, ChileUNCORRECTED Andrés PROOF Moreira-Muñoz 01 Acknowledgments 02

12 13 The book was initially developed as a doctoral study at the Geographical Institute 14 of Erlangen-Nürnberg University, Germany. Support in form of a grant was fortu- 15 nately provided by the German Academic Exchange Service (DAAD). I am much 16 indebted to Prof. Dr. Michael Richter, who was from the first moment the main sup- 17 porter of the idea. He and his family, together with all the colleagues and workers 18 at the Geographical Institute in Erlangen made our family’s stay in Germany a great 19 life experience. From the Geography to the Botanical Garden in Erlangen there are 20 just several blocks, and the support and friendship we found there in the person of 21 Dr. Walter Welss and his family was also a foothold in our stay. Prof. Dr.Werner 22 Nezadal (Erlangen) and Prof. Dr. Tod Stuessy (Vienna) gently assumed the revision 23 of the thesis. 24 The thesis was improved by the attendance of several conferences thanks 25 to grants from the Zantner-Busch Stiftung (Erlangen). At the conference 26 “Palaeogeography and Palaeobiogeography: Biodiversity in Space and Time”, 27 NIEeS, Cambridge, UK, 10th–11th April, I attended the workshop for using the 28 program TimeTrek for plate tectonic reconstructions. I also could attend the XVII 29 International Botanical Congress in Vienna, 17th–23rd July 2005. 30 The idea of transforming the thesis into a book found absolute support in the 31 person of Prof. Dr. Marinus Werger. He acted not just as a language editor but as a 32 very patient reviewer guiding the editing process in all its stages. The early intention 33 was also promoted by Dr. Leslie R. Landrum and Dr. Juan J. Morrone. 34 Crucial for the positive development of the book has been Springer’s produc- 35 tion and editing team: first Inga Wilde and Ria Kanters, and lately Ineke Ravesloot. 36 Several colleagues and friends graciously read and commented on draft chap- 37 ters: Federico Luebert (Berlin), Hermann Manríquez (Santiago), Patrick Griffith 38 (Florida), Malte Ebach (Arizona), Michael Heads (Wellington), Michael Dillon (Tal 39 Tal), Carlos Lehnebach (Wellington), and Patricio Pliscoff (Lausanne). Of course 40 the errors and misconceptions that may still exist are exclusively my responsibility. 41 In Chile, the project found early support in Dr. Belisario Andrade (Pontificia 42 Universidad Católica de Chile) and Dr. Roberto Rodríguez (Universidad de 43 Concepción). Once back in Chile, I can only express gratitude to the colleagues 44 at the Pontificia Universidad Católica de Chile, which facilitated my incorporation 45 UNCORRECTED PROOF

xi xii Acknowledgments

46 as an assistant professor by means of a grant for young doctors. I am especially 47 indebted to the Director of the Institute of Geography, Dr. Federico Arenas and the 48 dean of the Faculty, Dr. José Ignacio González. 49 Field work in Chile during 2008–010, especially for research on Asteraceae ◦ 50 (Chap. 8), was supported by project Fondecyt Iniciación (2008) n 11085016. 51 Speaking about field work, long ago I learned from Calvin and Linda Heusser the 52 “dirty side” of scientific field work. I will be always indebted to my old friends. 53 Vanezza Morales was a crucial helper in the final editing of most maps, and 54 with computer programs like NDM/VNDM. I gratefully mention also the important 55 advice provided by Tania Escalante (UNAM) and Claudia Szumik (U. de Tucumán). 56 Giancarlo Scalera (Roma), and Carlos Le Quesne (Valdivia) kindly provided articles 57 and figures. Sergio Elórtegui generously acceded to draw several original illustra- 58 tions for this work and also contributed many photographs. Carlos Jaña helped 59 finishing the most complicated figures. Sergio Moreira, Walter Welss, Hendrik 60 Wagenseil, Jeff Marso, María Castro, Francisco Casado, and Carlo Sabaini kindly 61 provided photos for illustrating this book. 62 Last but not least, I must acknowledge the life-long support of Mélica Muñoz- 63 Schick and Sergio Moreira, who could transfer to me their passion for nature 64 and beauty. Mélica, as ever, helped with the identification of species. Sergio also 65 helped providing scanned images of botanical specimens, thanks to a grant to the 66 National Herbarium provided by the Andrew W. Mellon Foundation trough the 67 Latin American Plants Initiative (LAPI). 68 When the doctoral thesis was still a draft project, my way crossed the one of 69 Paola, who soon turned to become my life companion. I would not have reached this 70 goal without her continuous support. I could also not imagine that the relationship 71 would be so fruitful: Sayén, Silene, Coyán, and Relmu remind me every evening 72 that there are other important things in life than just writing books... there is also 73 the possibility to read them!...especially when they deal not just with flowers but 74 also with rabbits, bears, elves and fairies.

76 1 May 2010 Limache

90 UNCORRECTED PROOF 01 Contents 02

14 Part I Geobotanical Scenario 15 1 The Extravagant Physical Geography of Chile ...... 3 16 1.1 Tectonics and Physiography ...... 6 17 1.2 Past Climate and Vegetation ...... 11 18 1.2.1 The Palaeozoic (542–251 mya) ...... 12 19 1.2.2 The Mesozoic (251–65.5 mya) ...... 15 20 1.2.3 The Cenozoic (65.5 mya Onwards) ...... 20 21 1.3 Current Climate and Vegetation ...... 31 22 References ...... 39 23

24 2 Getting Geobotanical Knowledge ...... 47

25 2.1 Romancing the South: The Discovery of a Virgin World .... 47

26 2.2 Classiﬁcation and Phylogeny of the Chilean Vascular Flora ... 55

27 2.2.1 Rich Families and Genera ...... 57

28 2.2.2 Endemic Families ...... 63

29 2.2.3 Phylogenetic Groups ...... 64

30 References ...... 79

32 Part II Chorology of Chilean Plants 33 3 Geographical Relations of the Chilean Flora ...... 87 34 3.1 Floristic Elements ...... 87 35 3.1.1 Pantropical Floristic Element ...... 91 36 3.1.2 Australasiatic Floristic Element ...... 92 37 3.1.3 Neotropical (American) Floristic Element ...... 95 38 3.1.4 Antitropical Floristic Element ...... 98 39 3.1.5 South-Temperate Floristic Element ...... 101 40 3.1.6 Endemic Floristic Element ...... 101 41 3.1.7 Cosmopolitan Floristic Element ...... 104 42 3.2 To Be or Not To Be Disjunct? ...... 105 43 3.2.1 Paciﬁc-Atlantic Disjunctions ...... 108 44 3.2.2 Antitropical (Paciﬁc) Disjunctions ...... 113 45 UNCORRECTED PROOF

xiii xiv Contents

46 3.3 In the Search for Centres of Origin: Dispersal v/s 47 Vicariance in the Chilean Flora ...... 114 48 3.3.1 Revitalizing Long-Distance Dispersal ...... 117 49 3.3.2 Relativising Long-Distance Dispersal ...... 120 50 References ...... 122 51 4 Biogeographic Regionalization ...... 129 52 4.1 The Chilean Plants in the Global Concert ...... 130 53 4.2 The Austral v/s the Neotropical Floristic Realm ...... 136 54 4.2.1 Floristic Elements in the Latitudinal Proﬁle ...... 137 55 4.2.2 Similarity Along the Latitudinal Gradient ...... 139 56 4.3 Regions and Provinces ...... 142 57 4.3.1 Endemism as the Base for Regionalization ...... 144 58 References ...... 147 59 60 Part III Islands Biogeography 61

62 5 Paciﬁc Offshore Chile ...... 153

63 5.1 Rapa Nui ...... 153

64 5.2 Islas Desventuradas ...... 157

65 5.3 Juan Fernández Archipelago ...... 159

66 5.3.1 The Unique Plant World of Juan Fernández ...... 159

67 5.3.2 Floristic Similarity of Juan Fernández ...... 161

68 5.3.3 Origins of the Fernandezian Flora ...... 167

69 5.3.4 Conservation of Juan Fernández Plants ...... 172

70 References ...... 177 71 6 Islands on the Continent: Conservation 72 Biogeography in Changing Ecosystems ...... 181 73 6.1 Fragmentation v/s Conservation on Chilean 74 Landscapes ...... 181 75 6.2 Global Change Biogeography: A Science of 76 Uncertainties ...and Possibilities ...... 185 77 References ...... 190 78 79 Part IV Case Studies on Selected Families 80

81 7 Cactaceae, a Weird Family and Postmodern Evolution ...... 197

82 7.1 Cacti Classiﬁcation ...... 198

83 7.2 Chilean Representatives and Their Distribution ...... 198

84 7.2.1 Cacti Distribution in Chile ...... 200

85 7.2.2 Areas of Endemism ...... 202

86 7.3 Notes on Cacti Conservation ...... 209

87 7.4 Biogeographic Insights: Spatial and Temporal Origins ..... 214

88 7.5 Cactus Postmodern Evolution ...... 216

89 References ...... 217

90 UNCORRECTED PROOF Contents xv

91 8 Asteraceae, Chilean Richest Family ...... 221 92 8.1 Classiﬁcation of Chilean Asteraceae ...... 221 93 8.2 Floristic Elements of Chilean Asteraceae ...... 223 94 8.3 Biogeographic regionalization of the Chilean Asteraceae .... 229 95 8.4 Asteraceae Evolutionary Biogeography ...... 234 96 8.4.1 Origin and Dispersal Routes ...... 238 97 8.4.2 Dispersal Capacities ...... 238 98 8.5 Conservation v/s Invasions ...... 239 99 8.5.1 Invading Biogeography ...... 242 100 References ...... 244 101 9 Nothofagus, Key Genus in Plant Geography ...... 249 102 9.1 Taxonomy and Phylogeny ...... 249 103 9.2 Diversity and Distribution ...... 254 104 9.3 Speciation v/s Extinction ...... 256 105 9.4 Vicariance v/s Dispersal and Centres of Origin ...... 260 106 9.5 Nothofagus and Associated Taxa ...... 262 107 9.6 Synthesis and Outlook ...... 263 108 References ...... 264 109 110 Part V Where to from Here? Projections of Chilean Plant Geography 111

112 10 All the Possible Worlds of Biogeography ...... 269

113 10.1 The Fragmented Map of Modern Biogeography ...... 269

114 10.2 Postmodern Biogeography: Deconstructing the Map ...... 270

115 10.3 Sloppy Biogeography v/s Harsh Geology? ...... 273

116 10.4 Just Some Possible Worlds ...... 275

117 10.4.1 Connections Over Land Bridges ...... 276

118 10.4.2 And What About a Closer Paciﬁc Basin? ...... 276

119 10.4.3 Three Models of Gondwana Fragmentation +

120 One Dispersal ...... 278

121 10.5 The “New Biogeography” ...... 281

122 10.6 Coda: The Geographical Nature of Biogeography ...... 283

123 References ...... 285 124 11 Epilogue: The Juan Fernández Islands 125 and the Long-Distance Dispersal of Utopia ...... 291 126 References ...... 293 127

128 Index ...... 329

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135 UNCORRECTED PROOF 01 Abbreviations 02

12 13 ACC Antarctic circumpolar current 14 cfr. Refer to 15 Chap. Chapter 16 col. sect. Colour section 17 ENSO El Niño Southern Oscillation 18 Fig. Figure 19 GIS Geographic information systems 20 K/T boundary Cretaceous/Cenozoic boundary 21 m asl Metres above sea level 22 mya Million years ago 23 Sect. Section 24 SGO National Herbarium Santiago, Chile 25 SNASPE National public protected areas system 26 yr BP Years before present 27

45 UNCORRECTED PROOF

xvii 01 About the Author 02

14 Andrés Moreira-Muñoz was born in Los Angeles (Chile), studied at the German

15 School in Santiago and graduated as Professional Geographer at the Pontiﬁcia

16 Universidad Católica de Chile. Botanical interest was inherited from his grand-

17 father and mother, both renowned botanists at the Museo Nacional de Historia

18 Natural in Santiago. He obtained his doctoral degree in Geography from the

19 University Erlangen-Nürnberg, Germany, under the direction of the plant geogra-

20 pher Prof. Michael Richter.

21 He currently occupies a position as assistant professor at the Instituto de

22 Geografía, Pontiﬁcia Universidad Católica de Chile, and develops research projects

23 about the chorology of Chilean plants, conservation biogeography and ﬁeld-based

24 education.

45 UNCORRECTED PROOF

xix 01 Part I 02 03 Geobotanical Scenario 04

45 UNCORRECTED PROOF 01 Chapter 1 02 The Extravagant Physical Geography of Chile 03

13 Abstract Current Chilean vascular ﬂora and its biogeographical patterns are 14 strongly related to the geographical features of the territory, past and present. Main 15 characteristics of the physical geography of Chile are described, with emphasis on 16 the geologic and climatic changes that affected the biome conﬁguration since the 17 Devonian onwards. Approaching the present time, the effects of the Pleistocene 18 glaciations in the distribution of several communities are discussed. 19

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

22 due to its impressive geographical contrasts: it contains the driest desert on the

23 planet, formidable inland ice ﬁelds, active volcanoes, fjords, geysers, a vast coastline

24 and the major highs of the Andes.

25 Chile stretches for 4,337 km along the south-western margin of South America from the Altiplano highs at 17◦35S to Tierra del Fuego, the Islands Diego Ramírez 26 ◦ 27 and Cape Horn at 56 S (Figs. 1.1 and 1.2 (col. sect.)). The country’s boundary to

28 the west is the wide Paciﬁc Ocean. The national territory includes several groups

29 of Paciﬁc oceanic islands, principally Rapa Nui (Easter Island), the Juan Fernández

30 archipelago, and the Desventuradas Islands (Fig. 1.1) (Chap. 5). Besides this the 2 31 nation has a geopolitical claim on a portion of 1,250,000 km in Antarctica. Though

32 geopolitical interests are beyond the scope of this book, and despite the modest

33 presence of extant vascular plants in Antarctica (only Deschampsia antartica and

34 Colobanthus quitensis), the Continent of Ice is of high interest regarding the origin

35 of the Chilean plant world (Sect. 1.2, Box 9.1). The eastern margin of mainland Chile is the Andes cordillera, which reaches to a 36 ◦ 37 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 38 ◦ 39 Ojos del Salado volcano at 27 06 S, reaching 6,893 m asl. Contrary to the long latitudinal extent, in width Chile rarely extends more than 200 km, reaching a maximum 40 ◦ ◦ 41 of 360 km at Mejillones (23 S) and a minimum of 90 km at Illapel (31 37 S). The

42 difference in altitude from the coast to the high Andes creates a series of bioclimatic

43 variations in the altitudinal proﬁle (Fig. 1.6). These variations, coupled with the cli-

44 matic latitudinal gradient, create a variety of geographic conditions that dramatically

45 UNCORRECTED PROOF

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

46 90” W 72” W 47 48 17” S 49

52 Islas Desventuradas 53

56 Juan Fernández

57 archipelago 110” W 58 Isla Salas y Gómez 59

60 Rapa Nui (Isla de Pascua)

69 Islas Diego 70 Ramírez 53” W 71

75 Oceanic Chile 76 Continental / Antarctic Chile 77

80 0 1,000 2,000 km 81

82 83 90° S 84

85 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 86 map provided by ESRI Labs 87

90 UNCORRECTED PROOF 1 The Extravagant Physical Geography of Chile 5

91 a b 92

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128 Fig. 1.2 Physical geography of Chile: a Valle de la Luna, Atacama desert, 23◦S; b Cerro Las 129 Vizcachas, Cordillera de la Costa, 33◦S; c rocky coast at Concón, Valparaíso (32◦50S); d Laguna ◦ 130 del Inca, Portillo, Andean pass to Argentina (32 50 S); e Glaciar Los Perros, Torres del Paine, ◦ ◦ 131 Campos de Hielo Sur (51 S); f southern fjords and Cordillera de Darwin (55 S) (photo credits: a,

132 b, d–f A. Moreira-Muñoz; c S. Elórtegui Francioli)

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135 UNCORRECTED PROOF 6 1 The Extravagant Physical Geography of Chile

136 affect the Chilean vegetation from the arid North to the humid temperate rainforests 137 in the South (Sect. 1.3).

138

139 140 1.1 Tectonics and Physiography 141 142 The main character of Chilean landscapes is driven by tectonic forcing: the geo- 143 logical evolution of Chile is related to the east-directed subduction of the Nazca 144 Plate beneath the South American Plate (Pankhurst and Hervé 2007) (Fig. 1.3). 145 The Chile Rise is an active spreading centre that marks the boundary between the 146 Nazca Plate and the Antarctic Plate at the so called Chile Triple Junction (Fig. 1.3). 147 The Nazca Plate is being subducted at a rate of ~65 mm/year (to the North of the 148 Triple Junction), while the Antarctic Plate is being subducted at a slower rate of 149 ~18 mm/year (Barrientos 2007). According to Ranero et al. (2006), the amount of 150 sediments to the trench is variable in space and time: north of 28◦S, due to aridity, 151 there is a relatively small amount of erosion and sediment supplied to the trench; in 152 the mid-latitude, the well developed river drainage system supplies much material 153

154

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156 Cocos Plate 157

158

159 a e s Galápagos 160 Pacific Plate Per 161 u -Chile Trench 162 cific Ri a Nazca Plate 163 t P 164 as b E Desventuradas 165 Rapa Nui Pitcairn Is. 166

167 Juan Fernández 168

169 Chile Ri c 170 South American 171 se Plate 172

173 Chile Triple d 174 Junction 175 Scotia Plate 176 Antarctic Plate

177

178 Fig. 1.3 Tectonic main features and volcanic zones of South America: a northern volcanic zone;

179 b central volcanic zone; c southern volcanic zone; d austral volcanic zone (adapted from Orme

180 (2007), by permission ofUNCORRECTED Oxford University Press; see also Stern et al. (2007 PROOF)) 1.1 Tectonics and Physiography 7

◦ 181 to the trench; south of ~ 40 S glacial-interglacial periods might have controlled the 182 amount of sediment supplied to the trench (Ranero et al. 2006). 183 A prominent feature of the Nazca Plate is the Juan Ferrnández hot spot chain, ◦ 184 a series of disconnected seamounts that disappear into the trench at 33 S (Ranero 185 et al. 2006) (Fig. 1.3). Subduction is accompanied by intense magmatic and seismic 186 activity (Orme 2007). Great earthquakes occur somewhere along the western South 187 American margin every few years, and “no recorded human generation in Chile has 188 escaped the damaging consequences of large earthquakes” (Barrientos 2007, p 263). 189 Indeed, while writing these lines, on the 27th of February 2010, an earthquake with 190 a magnitude of 8.8 followed by a tsunami affected Central-south Chile, resulting in 191 hundreds of deaths and thousands homeless. 192 Together with earthquakes, the active volcanism along the length of the country 193 is also a good reminder of the active tectonic processes acting below the surface 194 (Box 1.1).

195

196

197 198 Box 1.1 Living Under the Volcano 199 200 Chilean active and inactive volcanoes comprise ca.10% of the circum-Paciﬁc 201 “ring of ﬁre” (Pankhurst and Hervé 2007). These are mostly andesitic stra- 202 tovolcanoes that occupy almost the entire length of the country, especially 203 at the “South Volcanic Zone”, that encompass most of the South American 204 active volcanoes (Stern et al. 2007) (Figs. 1.3 and 1.4 (col. sect.)). More than 205 150 potentially active volcanoes have been detected, and 62 of them erupted in 206 historical times (González-Ferrán 1994). One of the most recent is the erup- ◦ 207 tion of Volcán Chaitén (43 S) on May 2008, which was responsible for the 208 obligate abandonment of the homonymous town. The ash column reached a 209 height of 15 km and spread wide upon the Atlantic (Fig. 1.5). Apart from its 210 consequences and risks for human occupation, volcanism has been a constant 211 source of disturbance in the Chilean ecosystems, especially in the southern 212 temperate forests (Milleron et al. 2008). 213

214 215 Taking account of its tectonic and morphostructural features, Chile can be classi- 216 ﬁed in a broad sense in ﬁve macrozones (Fig. 1.6) (Charrier et al. 2007; Stern et al. 217 2007): 218

219 ◦ 220 (a) The Coastal Cordillera occupies the western part of the proﬁle from 18 Sto ◦ 221 Chiloé Island (~ 42 S). It comprises the coastal batholith that consists predom- 222 inately of Late Palaeozoic and Mesozoic igneous rocks, with paired belts ◦ 223 of Palaeozoic metamorphic rocks cropping out south of Pichilemu (34 23 S) 224 (Pankhurst and Hervé 2007). Very impressive is the high riffs (“acantilado”) ◦ 225 that stretches fromUNCORRECTED 0 to 800 m asl at Iquique (20 S). PROOF 8 1 The Extravagant Physical Geography of Chile

226 a c 227

228

229

230

231

232

233

234 235 b 236

237

238

239

240

241

242 d 243

244

245

246

247

248

249

250 251 Fig. 1.4 Chilean volcanoes: a Parinacota volcano, 18◦10S; b steam expulsion of Volcán Lascar ◦ ◦ 252 (23 20 S), on December 1996; c Volcán Chaitén (42 50 S), false colour Aster satellite image:

253 plume of ash and steam advancing ca. 70 km to the north-east on January 2009; d lava ﬁelds ◦ 254 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)) 255

256

257

258 (b) The Central Depression is a tectonic downwarp with a Mesozoic to Quaternary 259 sedimentary fill of volcanic, glacial and fluvial origin. This main agricultural ◦ ◦ 260 and urbanized region ranges from 18 S to Copiapó (27 S), and again from ◦ ◦ ◦ ◦ 261 Santiago (33 S) to Chiloé (42 S). It is absent between 27 and 33 S, in the 262 so called zone of transverse river valleys or “Norte Chico” (Weischet 1970; 263 Charrier et al. 2007). This zone corresponds also to the “flat slab” zone, a 264 zone free of recent volcanic activity, associated to the subduction of the Juan 265 Fernández Ridge (Fig. 1.3). 266 (c) The main Andean Cordillera is a chain of mountains that dates back to the 267 Miocene, whose emergence continues today (see Box 1.5). It can be subdivided ◦ 268 in three segments: Forearc Precordillera and Western Cordillera, between 18 ◦ ◦ ◦ 269 and 27 S; High Andean Range, between 27 and 33 S (flat-slab subduction

270 UNCORRECTED PROOF 1.1 Tectonics and Physiography 9

271

272 (a)

273

274

275

276

277

278

279

280

281

282

283

284

285

286

287

288

289

290

291 292 (b) (c) 293

294

295

296

297

298

299

300

301 302 Fig. 1.5 Examples of volcanic activity in historical times: a ash expulsion by Volcán Antuco on 303 the 1st March 1839, as represented in Claudio Gay’s Atlas (Chap. 2); b eruption of Volcán Carrán

304 in 1955 (from Illies 1959); c Volcán Chaitén eruption photographed on May 26, 2008 (photo by J.N. Marso, courtesy of the USGS) 305

306

307

308 ◦ ◦ 309 segment); and Principal Cordillera, between 33 and ca. 42 S (Charrier 310 et al. 2007). 311 (d) Patagonian Cordillera: the Andes’ continuation right down into Tierra del Fuego 312 at the southern tip of Chile, with a continuous reduction in height (Pankhurst 313 and Hervé 2007). The origin of this low portion of the Andes has been related 314 to an allochtonous Palaeozoic terrane (see Box 1.2). The west-southern margin

315 UNCORRECTED PROOF 10 1 The Extravagant Physical Geography of Chile

Profile 20° S 316 4,000 317 3,000 318 2,000 20° 1,000 319 0 0 20 40 60 80 100 120 140 160 km 320 a Profile 30°S 321 b Ojos del 4,000 322 Salado 3,000 323 c 2,000 1,000 324 30° 0 0 20 40 60 80 100 120 140 km 325 Monte Aconcagua Profile 33°S 5,000 326 4,000 3,000 327 Cordillera de La Costa (La Campana) a 2,000 328 1,000 0 329 40° 0 20 40 60 80 100 120 140 km Profile 37.5° S 330 2,500 331 2,000 Cordillera de Nahuelbuta 1,500 332 1,000 500 333 0 e 0 20 40 60 80 100 120 140 160 180 200 220 km 334 Profile 40°S 1,600 335 50° 1,200

336 800 Cordillera Pelada

400 337 0 338 f 0 20 40 60 80 100 120 140 160 km Profile 50°S 339 2,000 Campo de Hielo Sur 340 1,500 Fjords and channels zone 1,000 341 500 342 0 0 20 40 60 80 100 120 140 160 km 343

344 Fig. 1.6 Physiography of continental Chile, on the base of SRTM (Shuttle Radar Topography Mission) data (http://www2.jpl.nasa.gov/srtm/) (for national political limits see Fig. 1.1). 345 Altitudinal proﬁles have been produced with ArcGIS 9 based on Aster GDM data (http://asterweb. 346 jpl.nasa.gov/gdem.asp). Note variations in the vertical scale, not homogeneous 347

348

349 ◦ 350 of the land (42 to the South) is modeled by recent glaciations that carved the 351 coastal areas into fjords and archipelagos comprising thousands of little islands 352 (Pankhurst and Hervé 2007). It has been calculated that the coastal extension 353 of Chile including these islands and southern archipelagos reaches 83,850 km! 354 (IGM 2005). 355 (e) The Andean foreland of the southern Patagonian Cordillera or Magallanes basin 356 consists of Upper Jurassic to Early Cenozoic sedimentary deposits (Charrier 357 et al. 2007; Fosdick 2007). 358

359

360 UNCORRECTED PROOF 1.2 Past Climate and Vegetation 11

361 362 Box 1.2 Patagonian Vicissitudes 363 364 The remarkable landscape and ﬂora of Patagonia motivated early naturalists 365 like the Perito Francisco P. Moreno to propose an independent origin of this 366 microcontinent from the rest of South America (Moreno 1882, as quoted by 367 Ramos 2008). The characteristic landscape and rocks led Moreno to remark 368 strong afﬁnities to other southern landmasses like Antarctica, Australia, and 369 New Zealand, suggesting that Patagonia was the rest of a sunken continent. 370 This view was retained even during the time of continental drift discussion 371 (e.g. Windhausen 1931). Current geologic and palaeomagnetic data suggests 372 that indeed, Patagonia has seen successive periods of breaking and drifting 373 during the whole Palaeozoic (Rapalini 2005; Ramos 2008). The TimeTrek 374 model (see also Pankhurst et al. 2006) shows an amalgamation of Patagonia 375 to Antarctic Peninsula during Late Carboniferous (300 mya), and a grad- 376 ual separation from Antarctica into the Cretaceous (120 mya) (Fig. 1.8). 377 Biotic exchange between South America and Antarctic Peninsula may have 378 been favoured (and then prevented) more than just one time, following rather 379 exchange cycles (Fig. 1.7). 380

381

382

383

384

385

386

387

388

389 a b c 390 391 Fig. 1.7 Positions of Patagonia: a in the Late Carboniferous (300 mya) aggregated to the 392 Antarctic Peninsula; b in the Early Cretaceous (120 mya), separated from Antarctica; c in

393 the Eocene (50 mya), again close to the Antarctic Peninsula. Modeled with TimeTrek v 4.2.5, Cambridge Paleomap Services 394

395

396

397

398 1.2 Past Climate and Vegetation

399

400 Tectonic and geomorphologic processes, coupled with the oceanic-atmospheric 401 system, have had enormous effects on the botanical evolution and its physiognom- 402 ical expression (i.e. the vegetation). The main aspects of the palaeogeographical 403 evolution of the territory will be resumed hereafter.

404

405 UNCORRECTED PROOF 12 1 The Extravagant Physical Geography of Chile

406 COOL WARM COOL WARM COOL WARM COOL WARM COOL 30°C 407

408

409 20°C 410

411

412 10°C P Ordovici Cret C Devoni Neogene C Q Permi J a S ri Tr a ua u l

413 a a il m r r eogene ass a u tern ass b ceo b ri 414 onif. ri a a a ic a a ic n a n ry us n n 415 n 145 2 199 3 4 65 251 299 44 542 416

416 2.5 3 59 88 3 417 418 Fig. 1.8 Global climate change since the Cambrian onwards. Adapted from Frakes et al. (1992) 419 and Scotese et al. (1999). Dates have been updated with the 2004 Geologic Time Scale (Gradstein 420 et al. 2004)

421

422

423 Palaeobotanical studies of Chile date back to Engelhardt (1891), Ochsenius

424 (1891), Dusén (1907), Berry (1922a, b), Fuenzalida (1938, 1966) among others.

425 More recent advances are centered in the Cenozoic (e.g. Cecioni 1968; Nishida

426 1984; Troncoso and Romero 1998; Hinojosa 2005). Constant improvement of the

427 methods applied to the study of “climatically sensitive” sediments (e.g. coals, salt

428 deposits, evaporites), together with studies in diversity patterns in global vegetation

429 through time, are beneﬁting our understanding of the evolution of plant biomes in

430 space and time (Willis and McElwain 2002).

431 The ﬂoristic and vegetational history of southern South America is strong related

432 to the tectonic and climatic history of the Gondwana continent (McLoughlin 2001). AQ1 433 “During the 500 million years that Gondwana and its fragments existed, the ‘Earth’

434 global climate system has shifted from ‘Ice House’ conditions to ‘Hot House’ condi-

435 tions four times” (Scotese et al. 1999) (Fig. 1.8). These global climatic ﬂuctuations

436 have constantly affected the biotic evolution and biogeography: ﬂoristic regions

437 can be tracked back even to the mid-late Silurian, the time when according to

438 most palaeobotanical evidence, the vascular plants have conquered the land surface

439 (Willis and McElwain 2002; Raymond et al. 2006)(Box2.3).

440

441

442 1.2.1 The Palaeozoic (542–251 mya)

443

444 Several orogenic events affected the western margin of Gondwana from the Late 445 Proterozoic to the Palaeozoic (Ramos and Aleman 2000; Pankhurst et al. 2006). 446 The Famatinian orogeny in the Ordovician (~490–450 mya) is characterized by the 447 amalgamation of several allochtonous terranes, like Cuyania and Chilenia, imply- 448 ing that North America had collided with West Gondwana by that time (Astini et al. 449 1995). Mejillonia and Patagonia terranes amalgamated in the Early Permian, as the 450 last convergence episodesUNCORRECTED (Ramos 2009)(Box 1.2). The development PROOF of preAndean 1.2 Past Climate and Vegetation 13

451 foreland basins during the Palaeozoic, set the stage for the initiation of the Andes 452 long before the event that culminated in massive Cenozoic uplift (Orme 2007). 453 During the Late Palaeozoic, Gondwana became amalgamated to the supercontinent 454 of Laurussia to form the vast single landmass called Pangaea. 455 From the Early Devonian to the Late Carboniferous (400–300 mya), global vege- 456 tation evolved from one dominated by small, weedy plants, only several decimetres 457 in height, to fully forested ecosystems with trees reaching sizes of 35 m (Willis 458 and McElwain 2002). During the Middle to Late Devonian (390–360 mya) warm, 459 humid climates with high levels of atmospheric CO2 prevailed worldwide, favouring 460 the appearance of earliest arborescent forms of plants (see Box 2.3). 461 By the Late Carboniferous (330–299 mya) the southern flora consisted 462 mainly of likely pteridosperms, lycopsids, Cordaites and Ginkgophytes (Vega and 463 Archangelsky 1997). Diversity was rather low, and the southern flora was uni- ◦ ◦ 464 formly developed across Gondwana between 30 S and 60 S (Anderson et al. 465 1999; DiMichele et al. 2001). However, Cúneo (1989) suggests that floristic 466 differentiation was also apparent on the west coast of South America. The pres- 467 ence of Lepidodendron and Sigillaria (lycopod trees) has been reported from 468 the Carboniferous deposits of Chile (Charrier 1988). Late Carbonifeous ended 469 in a widespread glaciation, one of the most severe in Earth’s history. The 470 Permo-Carboniferous glaciation (310–290 mya) lasted for around 30 million years 471 (Beerling 2002); Gondwanan continents were locked in deep glaciation (Fig. 1.9).

472

473

474

475

476

477

478

479

480

481

482

483

484

485

486

487

488 Subtropical desert 489

490 Cool temperate

491

492 Ice sheet

493

494 Fig. 1.9 Late Carboniferous biomes (adapted from Willis and McElwain (2002) on a TimeTrek

495 4.2.5 model, CambridgeUNCORRECTED Paleomap Services) PROOF 14 1 The Extravagant Physical Geography of Chile

496 The Permian (299–251 mya) was characterized by major global climate changes, 497 from glaciated (icehouse) to completely ice-free (hothouse) stages (Fig. 1.8). “With 498 the onset of glaciation in the Permian, the flora changed dramatically with the 499 appearance of Glossopteris and the disappearance of most of the Late Carboniferous 500 elements” (DiMichele et al. 2001, p 467). By the Middle Permian, one of the most 501 striking vegetation changes was the relatively increased proportion of seed plants 502 together with a reduction of the swamp-dwelling lycopsids and sphenopsids (Wnuk 503 1996, McAllister Rees et al. 2002). Glossopteris, a gymnosperm genus with many 504 species, turned to be the characteristic plant of Gondwana (DiMichele et al. 2001). 505 Indeed, Glossopteris dominant presence across Gondwana is one of the keys that 506 supported the continental drift theory of Alfred Wegener. Botrychiopsis, another 507 typical species from west Gondwana, went extinct when the environmental condi- 508 tions typical of a greenhouse stage were created by the end of the Permian (Jasper 509 et al. 2003). 510 The Permian flora of Gondwana was significantly more diversified than the one 511 of the Late Carboniferous (Cúneo 1989), and the floristic provinciality changed dur- ◦ ◦ 512 ing the course of the Permian. The belt located between 60 and 45 Sinwestern 513 Gondwana was called the “Southern temperate semiarid belt of middle latitudes”, 514 characterized by Glossopteris and moderately thermophilic vegetation with abun- 515 dant tree-ferns and lycopods (McLoughlin 2001; Chumakov and Zharkov 2003) 516 (Fig. 1.10).

517

518

519

520

521

522

523

524

525

526

527

528

529

530

531

532 Subtropical desert

533 Mid latitude desert 534 535 Cold temperate 536

537 Cool temperate

538

539 Fig. 1.10 Middle Permian biomes (adapted from Willis and McElwain (2002) on a TimeTrek

540 4.2.5 model, CambridgeUNCORRECTED Paleomap Services) PROOF 1.2 Past Climate and Vegetation 15

541 1.2.2 The Mesozoic (251–65.5 mya)

542

543 The transition from the Palaeozoic to the Mesozoic is characterized by a dramatic 544 event: the Permian-Triassic extinction event, which apparently saw the destruction 545 of 90% of marine life on Earth due to extensive volcanism, under other causes 546 (Benton and Twitchett 2003). The impacts on the terrestrial ecosystem were not 547 so drastic, or paradoxically even favorable for some plants (Looy et al. 2001). 548 The Triassic (251–199.6 mya) climate was relatively warm compared to today, 549 and continentality and aridity were more extended due to the permanence of the 550 single continent Pangaea. The Triassic flora remained broadly similar to that of the 551 Permian, dominated by gymnosperms (seed ferns, cycads, and ginkgos). During 552 the Triassic, Glossopteris-dominated communities were replaced by Dicroidium 553 (a seed fern) dominated floras across the Southern Hemisphere (McLoughlin 2001). 554 Also, the major radiation of conifers, e.g. the Araucariaceae began in the Triassic 555 (see Sect. 2.2). Other important components of the southern flora were ginkgo- 556 phytes, putative gnetales, bennettitales, and cycadales, plus many lycophytes and 557 osmundacean, gleicheniacean, dicksoniacean, dipteridacean and marattiacean ferns 558 (McLoughlin 2001, p 286; Artabe et al. 2003) (see Sect. 2.2). 559 The Jurassic (199.6–145.5 mya), better known for the diversification of charis- 560 matic faunal groups like the dinosaurs, is also considered one of the most important 561 periods in plant evolution. By the Early Jurassic, both composition and distribu- 562 tion of southern hemisphere vegetation had changed dramatically. Glossopteris and 563 Dicroidium no longer dominated the southern flora. Instead they were replaced by 564 cycads, bennetittes, ginkgos, and conifers, and for the first time global floras con- 565 tained a significant portion of forms that are recognizable in our present floras. The 566 floral assemblage for Cerro La Brea, Mendoza, Argentina (Early Jurassic) shows the 567 presence of 14 taxa belonging to the Equisetaceae, Asterothecaceae, Marattiaceae, 568 Osmundaceae, Dipteridaceae, and several conifers (Artabe et al. 2005). 569 While Gondwana drafted towards the equator, five distinct biomes settled dur- 570 ing the Early Jurassic (McAllister Rees et al. 2000) (Fig. 1.11). Southern South 571 America must have been occupied by a “winterwet biome” with a climate simi- 572 lar to that of today’s Mediterranean-type one. The relatively increased proportion 573 of plants with small leaves and other xerophytic features clearly indicates seasonal 574 water deficits (Willis and McElwain 2002). In the Middle Jurassic, main compo- 575 nents of this biome, like Cycadales, Bennettitales, conifers, ferns, and Sphenopsids, 576 reached northernmost Chile, i.e. current arid Atacama (Fuenzalida 1966; Herbst and 577 Troncoso 1996). 578 Quattrocchio et al. (2007) listed more than a hundred species from the Jurassic of 579 the Neuquén basin, Argentina. Clearly dominant groups were the Cheirolepidiaceae, 580 Araucariaceae and Podocarpaceae, together with Cyatheaceae, Osmundaceae, 581 Marattiaceae, Dipteridaceae, Lycopodiaceae, Schizaeaceae, Anthocerotaceae, 582 Ricciaceae, Cycadales/Bennettitales, Caytoniaceae and Gnetales. The authors 583 further propose an environmental model in which the Araucariaceae and 584 Podocarpaceae occupied mostly high-altitude places, while ferns, cycads and 585 Cheirolepidiaceae mayUNCORRECTED have been restricted to more low-lying PROOF and humid places 16 1 The Extravagant Physical Geography of Chile

586 (Fig. 1.12). Let us keep in mind that there was still not such thing like an elevated 587 Andes (Box 1.5)

588

589

590 Box 1.3 Gondwana Breaks-Up

591

592 Most authors recognize three major separation events of Gondwana that

593 affected the evolution of the South American ﬂora: the separation between

594 W and E Gondwana during the Jurassic (180–150 mya); the separation

595 America/Africa between 119 and 105 mya, and the split between Antarctica

596 and southern South America (32–28 mya) (Table 1.1). These ages serve as

597 reference; but there is no real consensus on the time of fragmentation of the

598 different components. The crucial separation of Australia from Antarctica

599 and South America from Antarctica and the development of the Drake

600 Passage is still a controversial issue: “South America may have separated AQ2 601 from Antarctica as early as the Late Jurassic (Smith et al. 1994), or as late AQ3 602 as the Palaeocene (Hay et al. 1999) or Eocene (Brundin 1988)” (Orme 2007, AQ4 603 p 10) (see Box 9.1). The TimeTrek model shows indeed a much earlier sepa-

604 ration of South America and Antarctica at around 130 mya (Early Cretaceous)

605 (Fig. 1.8).

606

607

608

609

610

611

612

613

614

615

616

617

618

619

620

621

622

623 Subtropical desert 624 625 Winterwet 626

627 Warm temperate

628

629 Fig. 1.11 Early Jurassic biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5

630 model, Cambridge PaleomapUNCORRECTED Services) PROOF 1.2 Past Climate and Vegetation 17

631

632 Cycadales Podocarpaceae

633 Cheirolepidiaceae Araucariaceae v 634 Lycopodiaceae and ferns v v v v 635 v v v v v v v 636 v v v v Igneous rocks v v v v 637 v v v v v v v v v Marine deposits v v v Continental v 638 v v v v v v deposits v v v v v v v v v v v v v v v v v 639 640 Fig. 1.12 Palaeoenvironmental reconstruction of middle Jurassic ﬂora from Neuquén, Argentina 641 (adapted from Quattrocchio et al. 2007)

642

643

644

645

646 Southern Floras during Early Cretaceous did not differ much from the 647 Late Jurassic ones (Fig. 1.13). Most famous is the middle Cretaceous, 648 known as the period of expansion and radiation of the angiosperms (see 649 also Box 2.4). Angiosperms evolving during this time include a number of 650 families that constitute a significant part of the present-day global flora (e.g. 651 Betulaceae, Gunneraceae, Fagaceae/Nothofagaceae). For the early Late Cretaceous 652 (Cenomanian to Coniacian), Troncoso and Romero (1998) reported a Neotropical 653 flora showing a notable change compared to the previous ones. They reported 654 the definitive replacement of the dominance of gymnosperms by angiosperms, 655 including representatives of extant families, such as the Lauraceae, Sterculiaceae, 656 Bignoniaceae, and Monimiaceae; and from extant genera like Laurelia, Peumus, 657 and Schinopsis (this last genus is currently not present in Chile). 658 By the Late Cretaceous, (Campanian-Maastrichtian) Troncoso and Romero 659 (1998) reported a Neotropical flora with marginal presence of Nothofagus 660 (Campanian first appearance of Nothofagus in Antarctica; Maastrichtian first 661 appearance of Nothofagus in the fossil record from Central Chile and Tierra del 662 Fuego) (see also Chap. 9). In spite of its marginal presence, it is the peak of northern ◦ 663 expansion of Nothofagus in South America, reaching 30 S (Torres and Rallo 1981) 664 (Fig. 1.14). This expansion of Nothofagus is challenging since the Late Cretaceous 665 is considered a rather greenhouse world. It is but possible that transient small ice- 666 caps existed during this mostly warm period. It has been proposed that relatively 667 large and short-term global sea level variations may have been connected with small 668 and ephemeral ice sheets in Antarctica, probably related to short intervals of peak 669 Milankovitch forcing (Gallagher et al. 2008). 670 Southern South America, already isolated from the rest of western Gondwana, 671 was occupied mainly by a “subtropical desert” and a “warm temperate” biome 672 (Fig. 1.14), the latter being characterized by Araucariaceae, Nothofagaceae, 673 Proteaceae, and Winteraceae (Willis and McElwain 2002). “The presence of tropi- 674 cal elements in the austral margin of South America gives support to the expansion 675 of a warm climateUNCORRECTED towards high latitudes during the mid Cretaceous” PROOF (Barreda 18 1 The Extravagant Physical Geography of Chile

676 Table 1.1 Three stages in the break-up of Gondwana (as resumed by McLoughlin 2001)

677 Major separation Palaeoreconstructions on a TimeTrek 678 events Period and causes v. 4.5.2 model 679

680 (W Gondwana / During Middle to Late Jurassic E Gondwana) (180–150 mya): breakup 681 associated with development 682 of a series of deep seated 683 mantle plumes beneath the

684 extensive Gondwanan

685 continental crust in S Africa (c 182 mya) and the 686 Transantarctic mountains 687 (c 176 mya) (Storey 1995) 688

689

690

691 Africa–S Early Cretaceous (119–105

692 America mya): opening of the South separation Atlantic Ocean, due to the 693 emplacement of 694 Plume-related 695 Parana-Etendecka 696 continental ﬂood basalts in

697 Brazil and Namibia (137–127 mya). Final 698 break-up of Africa and S 699 America was completed only 700 at 80 mya 701

702 West Early Oligocene (ca 30 mya):

703 Antarctica-S beginning at ~35–30.5 mya America as a subsidence in the Powell 704 Basin followed by seaﬂoor 705 spreading. Opening of the 706 Drake Passage between the 707 southern tip of South

708 America and the northern end of Antarctic Peninsula 709 allowed deep water 710 circulation and the 711 installation of the Antarctic 712 Circumpolar Current (ACC)

713 between 41 and 24 mya (see Box 9.1) 714

715

716 and Archangelsky 2006). Troncoso and Romero (1998) also reported the pres- 717 ence of Neotropical palaeoﬂoras in the mid- and Late Cretaceous from Magallanes 718 and Tierra del Fuego. Microfossils assigned to the Arecaceae (Palmae) have been 719 reported since the Maastrichtian (Hesse and Zetter 2005). 720 UNCORRECTED PROOF 1.2 Past Climate and Vegetation 19

721

722

723

724

725

726

727

728

729

730

731

732

733

734

735

736

737

738

739 Fig. 1.13 Illustration of the biotic assemblage from the limit Jurassic/Cretaceous (145.5 mya) 740 of the Southern Cone. Theropod dinosaur on a swamp surrounded by ginkgos, araucarias, and 741 arborescent ferns (original illustration by Sergio Elórtegui Francioli) 742

743

744

745

746

747

748

749

750

751

752 Tropical summerwet 753

754 Subtropical desert

755 Mid latitude desert 756

757 Warm temperate 758

759 Cool temperate

760

761

762 Fig. 1.14 Late Cretaceous biomes; arrow shows northernmost expansion of Nothofagus (see text) 763 (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5 model, Cambridge Paleomap 764 Services) 765 UNCORRECTED PROOF 20 1 The Extravagant Physical Geography of Chile

766

767

768 Box 1.4 Floral Extinction at the K/T Boundary ?

769 A permanent question is whether massive extinction events that mostly 770 affected the terrestrial fauna affected as well the global flora (McElwain and 771 Punyasena 2007). It seems that at the K/T boundary, at least several groups 772 suffered similar luck than dinosaurs, plesiosaurs, and ammonoids. For exam- 773 ple, the seed-ferns, a group that dominated the vegetation formations in many 774 parts of the world from the Triassic to the Cretaceous, are considered to have 775 disappeared at the end of the Cretaceous. Neverthless, exceptions are the rule, 776 and there is a seed-fern fossil recent discovered in Tasmania that has been 777 dated from the Early Eocene (McLoughlin et al. 2008). 778 Recent findings on the Lefipán Formation in NW Chubut province dated 779 as Maastrichtian, supports the catastrophic character of the K/T boundary 780 (Cúneo et al. 2007). The discovery of a highly diversified assemblage of 781 dicot leaves with probably more than 70 species, as well as several mono- 782 cots, podocarp conifers, and ferns, suggests that the latest Cretaceous floras 783 were probably more diverse than those known from Patagonia during the 784 Palaeocene. This means that the K/T event indeed affected the terrestrial 785 ecosystems of southern latitudes. The recovery of floral diversity must have 786 taken most of the Palaeocene until the recovering of plant richness by the early 787 Eocene (Cúneo et al. 2007). 788

789

790

791

792

793 1.2.3 The Cenozoic (65.5 mya Onwards)

794

795 The deep-sea oxygen isotope record permits a detailed reconstruction of the 796 Cenozoic global climate, that has suffered a number of episodes of global warm- 797 ing and cooling, and ice-sheet growth and decay (Zachos et al. 2001) (Fig. 1.15). 798 The most pronounced warming occurred from the Mid-Palaeocene (59 mya) to the 799 Early Eocene (52 mya), showing a peak in the so called Early Eocene Climatic 800 Optimum (52–50 mya) (Fig. 1.15). This period was one of the warmest periods in ◦ 801 the Earth’s history: temperature estimates of between 9 and 12 C higher than present 802 have been proposed (Zachos et al. 2001). This optimum was followed by a trend 803 toward cooler conditions in the Late Eocene. According to Zachos et al. (2001), ice- 804 sheets appeared in the Early Oligocene, and persisted until a warming phase that 805 reduced the extent of Antarctic ice in the Late Oligocene Warming (Fig. 1.15). From 806 this point (26–27 mya) until the middle Miocene (15 mya), the global ice volume 807 remained low with the exception of several brief periods of glaciation. This warm 808 phase peaked in the Middle Miocene Climatic Optimum (17–15 mya), and was fol- 809 lowed by a gradual cooling and reestablishment of a major ice-sheet on Antarctica 810 towards the Plio/PleistoceneUNCORRECTED (Zachos et al. 2001) (Fig. 1.15). PROOF 1.2 Past Climate and Vegetation 21

811 Early Eocene Climatic Optimum 812 y a 813 12

ent d Late Oligocene 814 s Warming 815 8 Late Palaeocene Middle Miocene 816 Climate Optimum

ove pre ove Thermal Maximum

817 ab 4 West Antarctic 818 Ice-sheet 819 re (°C)

u Glaciation

t 0 820 Glaciation East Antarctic a

821 Ice-sheet Palaeoc. Eocene Oligocene Miocene Plio/Pleist. 822 K Temper 823 70 60 50 40 30 20 10 0 Age (mya) 824 825 Fig. 1.15 Global climatic ﬂuctuations during the Cenozoic, based on global deep-sea oxygen and 826 carbon isotope records (adapted from Zachos et al. 2001)

827

828

829 In the Early Palaeocene (~65–55 mya) the global position of South America 830 had moved close to the present-day position (Fig. 1.14). Nevertheless, the cold cir- 831 cumpolar ocean current had not yet developed, and Paciﬁc Ocean currents carried 832 heated tropical waters to high latitudes. As a consequence, a permanent ice cover at 833 the poles was absent, and the prevailing low relief of the continents, coupled with 834 high seas, resulted in rain-bearing winds penetrating far into the interior of all the 835 main landmasses (Willis and McElwain 2002). 836 South America was mainly occupied by “tropical everwet”, “subtropical desert” 837 and “warm temperate” biomes. The warm temperate biome was composed of ever- ◦ 838 green and deciduous dicots (e.g. Nothofagus), and podocarps. South of 70 S, and 839 widespread in Antarctica, a “warm cool temperate biome” was established, com- 840 posed mainly by Araucaria, Podocarpus, Dacrydium, evergreen Nothofagus, and to 841 a minor extent members of the Loranthaceae, Myrtaceae, Casuarinaceae, Ericaceae, 842 Liliaceae, and Cunoniaceae (Truswell 1990). 843 Troncoso and Romero (1998) emphasized the neotropical character of 844 the Palaeocene palaeoﬂoras of Central and Southern Chile. Zonal vegeta- 845 tion was composed mainly of rainforests with palms, mangroves, and in the 846 higher parts, azonal vegetation composed of Gymnosperms (Cheirolepidaceae, 847 Araucariaceae, Podocarpaceae, Zamiaceae) and Nothofagus, accompanied by 848 Myrtaceae, Proteaceae and Lauraceae. Fossil Boraginaceae related to extant Cordia 849 have been described by Brea and Zucol (2006) from the Late Palaeocene of Chubut, 850 Argentina. A rich assemblage of micro- and megafossils has been described by ◦ 851 Troncoso et al. (2002) from the Ligorio Márquez Formation in Aisén (47 S). Of the 852 twenty leaf species reported, fourteen are from the Lauraceae; the rest corresponding 853 to the Melastomataceae, Myrtaceae, Sapindaceae, and others. Furthermore, seven 854 Pteridophyta, two conifers, and four angiosperms are represented by palynological 855 species. In spite of thisUNCORRECTED predominantly tropical character, the presence PROOF of temperate 22 1 The Extravagant Physical Geography of Chile

856 taxa like Nothofagus and Podocarpaceae confirms the warm temperate tendency at ◦ 857 47 S (Okuda et al. 2006). 858 Recently Iglesias et al. (2007) reported a greater species richness than was 859 previously known from Palaeocene Patagonia, including more than 43 species of 860 angiosperm leaves. At the end of the Palaeogene, representatives of most of the 861 angiosperm modern classes and many orders were already present in southern South 862 America (Gandolfo and Zamaloa 2003; Prámparo et al. 2007). 863 Eocene (55.8–33.9 mya) floras of Southern South America show subtropical 864 to fully tropical forests, with zones of seasonal dryness in Chile (Romero 1986). 865 The three extant South American tribes of the Proteaceae were already present 866 in the early Eocene, forming the Australia-Antarctica-South America connection 867 (González et al. 2007). Late Eocene fossil leaves, flowers and fruits assigned to the 868 Escalloniaceae have also been reported as being involved in this austral connection 869 (Troncoso and San Martín 1999). 870 Remarkable is the presence of Eucalyptus macrofossils in the Patagonian Early 871 Eocene (Gandolfo et al. 2007), since the genus shows an extant distribution 872 in Australasia, mainly Australia and Tasmania (not New Zealand). The South 873 American macrofossils reported by Gandolfo et al. (2007) are to date the most 874 ancient register for the genus. 875 The Laguna del Hunco palaeoflora in NW Chubut, Argentina, shows the most 876 complete example of Early Eocene vegetation in South America. This palaeoflora 877 is one of the world’s most diverse Cenozoic assemblages of angiosperms (Wilf 878 et al. 2005, 2007). This assemblage comprises tropical elements restricted today 879 to temperate and tropical Australasia (e.g. Dacrycarpus, Papuacereus, Eucalyptus); 880 tropical elements (e.g. Roupala, Bixa, Escallonia), and the disjunct element South 881 America/Australasia (e.g. Eucryphia, Orites, Lomatia) (see Fig. 3.5 (col. sect.)). 882 Fossil plants at Laguna del Hunco are extremely abundant, diverse (>150 leaf 883 species), and well-preserved. During the early Eocene the area was a subtropi- 884 cal rainforest with land connections both to Australasia via Antarctica and to the 885 Neotropics (Fig. 1.16). 886 Wilf et al. (2007) suggest that the Laguna del Hunco plant lineages retreated 887 to geographically disparate rainforest refugia following post-Eocene cooling and 888 drying in Patagonia. Only few lineages adapted and persisted in temperate South 889 America. 890 The continuous decrease in temperature during the Eocene allowed a new dis- 891 placement of Nothofagus towards South-Central Chile. Therefore this time-span is 892 characterized by a mixed tropical-subantarctic palaeoflora (Troncoso and Romero 893 1998). In spite of the prevalence of mixed palaeofloras during the Eocene, results ◦ 894 obtained by Gayó et al. (2005) at Bahía Cocholgüe (36,5 S) suggest that tropical 895 floras persisted in central Chile during the Early Eocene and formed a belt between ◦ ◦ 896 25 S and 37 S. This persistence of tropical floras (composed mainly by Lauraceae 897 and Myrtaceae) might be related to the influence of the Early Eocene Climatic 898 Optimum (Fig. 1.15) and to a shrinking tropical belt (Gayó et al. 2005). 899 The transition from the Eocene to the Oligocene (33.9–23.03 mya) was a period 900 of significant globalUNCORRECTED climatic cooling and increased aridity, major changesPROOF in oceanic circulation, and the initiation of ice on Antarctica (Zachos et al. 2001; Convey 1.2 Past Climate and Vegetation 23

901

902

903

904

905

906

907

908

909

910

911

912

913 Tropical summerwet 914

915 Subtropical desert

916 Warm temperate 917 918 Cool temperate 919

920 Fig. 1.16 Early Eocene biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5 921 model, Cambridge Paleomap Services) 922

923

924 et al. 2008) (Fig. 1.15). Major reorganization and redistribution of global vege- 925 tation followed these climatic trends, with a reduction of tropical forests and the 926 expansion of temperate vegetation toward the equator (Willis and McElwain 2002). 927 A Subantarctic palaeoﬂora expanded its distribution range across southern South 928 America, occupying an area that became to extend from the island of Tierra del 929 Fuego to the south of Central Chile (Romero 1993) (Fig. 1.17). 930 This implicates the massive retreat of tropical and subtropical compo- 931 nents from the Sapindaceae and Lauraceae, the generic replacement of gen- 932 era in the Rhamnaceae, Myrtaceae, Bignoniaceae, Flacourtiaceae/Salicaceae; 933 and the regional extinction of several families like Moraceae, Annonaceae, 934 Dilleniaceae, Malpighiaceae, Vochysiaceae, Tiliaceae, Sterculiaceae, Sapotaceae, 935 and Styracaceae (Troncoso and Romero 1998). Permanent ice sheets persisted on 936 Antarctica until the Late Oligocene (26–27 mya), when a warming trend reduced 937 the extent of Antarctic ice (Zachos et al. 2001). 938 From the “Late Oligocene Warming” (26–27 mya) (Fig. 1.15) until the Middle 939 Miocene (~15 mya), the global ice volume remained low and water showed slightly 940 higher temperatures, intermingled with brief periods of glaciation (Zachos et al. 941 2001). This was followed by a gradual cooling and reestablishment of a major 942 ice-sheet on Antarctica by 10 mya (Fig. 1.18). The continental interior became 943 increasingly arid/cold and large areas of shorelines were exposed due to a falling sea 944 level. Outside the core a depauperate “cold temperate” biome survived, having lost 945 its main forests componentsUNCORRECTED and with some herbs and C3 grasses PROOF remaining. During 24 1 The Extravagant Physical Geography of Chile

946

947

948

949

950

951

952

953 Tropical everwet 954 Subtropical summerwet 955

956 Woody savannah 957

958 Warm/cool temperate

959 Cool/cold temperate 960

961 Cold temperate 962 Ice sheet 963

964 Fig. 1.17 Early Oligocene biomes; arrows show mixture of tropical and austral ﬂoras (adapted 965 from Willis and McElwain (2002) on a TimeTrek 4.2.5 model, Cambridge Paleomap Services) 966

967

968

969

970

971

972

973

974

975

976

977 Tropical everwet

978 Subtropical summerwet 979 (woody savannah) 980 Warm temperate 981 Cool temperate 982

983 Cold temperate 984

985 Ice sheet

986 987 Fig. 1.18 Miocene biomes (adapted from Willis and McElwain (2002) on a TimeTrek 4.2.5 model, 988 Cambridge Paleomap Services)

989

990 UNCORRECTED PROOF 1.2 Past Climate and Vegetation 25

991 the Late Miocene most of Western South America was occupied by a “cool temper- 992 ate” biome. The “winterwet” and “subtropical summerwet” biomes were restricted 993 to a reduced proportion of today’s Atacama Desert (Fig. 1.18). 994 The Miocene (23.03–5.33 mya) is characterized by a development of mod- 995 ern angiosperm families like Asteraceae, Poaceae, Malvaceae, Fabaceae and 996 Cyperaceae, related to more open communities replacing tropical forests in southern 997 South America (Barreda et al. 2007; Palazzesi and Barreda 2007). Tropical forests 998 were still abundant during the Early Miocene of Patagonia; the vegetation increas- 999 ingly acquired a more complex aspect, due to the wider distribution of grasses and 1000 shrubs. Barreda et al. (2007) list 60 angiosperm families present in the fossil record 1001 during the Miocene in Argentina. More diversified families are the Malvaceae (33 1002 taxa), Fabaceae (32), and Asteraceae (25). 1003 During the Early and Middle Miocene, the subantarctic flora reached the south- 1004 ern part of Central Chile (Troncoso and Romero 1998). In the middle Miocene of 1005 Central Chile a change from the previous subantarctic palaeoflora into a mixed 1006 palaeoflora with a predominance of neotropical taxa and the retreat of subantarc- 1007 tic taxa occurred (Hinojosa 2005). The subsequent subtropical palaeoflora that 1008 occupied central Chile during the lower to mid Miocene, 20–15 mya, developed 1009 under a warmer and more humid palaeoclimate, with an incipient Andean rain 1010 shadow effect, is the nearest ancestor of the sclerophyllous modern vegetation of 1011 central Chile (Hinojosa et al. 2006). 1012 In northern Chile, the hyperarid climate became established at the Oligocene/ 1013 Miocene boundary (ca. 25 mya) (Dunai et al. 2005; Nalpas et al. 2008), and was 1014 followed by more humid (semiarid periods interrupted by short arid events up to the 1015 earliest Late Pliocene (Hartley and Chong 2002, Box 3.1)). 1016 Global deep-sea oxygen and carbon isotope records indicate additional cooling 1017 and small-scale icesheet expansion on west-Antarctica during the Late Miocene 1018 until the Early Pliocene (6 mya). The early Pliocene is marked by a subtle warm- 1019 ing trend between 3.3 and 3 mya. Afterwards cooling again increased (Zachos et al. 1020 2001).

1021

1022

1023 Box 1.5 Slow or Rapid Andean Uplift?

1024

1025 The Late Miocene has been proposed as the initial phase of the Andes uplift.

1026 Gregory-Wodzicki (2000), on the base of palaeobotanical data, proposed a

1027 surface uplift in the order of 2,300–3,400 m asl since the late Miocene at

1028 uplift rates of 0.2–0.3 mm/year. More recently Ghosh et al. (2006) obtained

1029 results that indicate a surprisingly rapid uplift of the Bolivian Altiplano at AQ5 1030 an average rate of 1.03 +-0.12 mm per year between ~10.3 and ~6.7 mya (i.e.

1031 from 0 to 4,000 m asl since the Middle/Late Miocene). These results challenge

1032 the known forces responsible for the uplift and are in conﬂict with geological

1033 evidence (e.g. Hartley 2003, proposed a proto-Central Andean mountain range

1034 placed between 15 and 9 mya). Geomorphological evidence, i.e. lahar deposits

1035 UNCORRECTED PROOF 26 1 The Extravagant Physical Geography of Chile

1036 ◦ ◦ 1037 in the Coastal Cordillera of central Chile (33 40 –34 15 S) still supports an 1038 Oligocene–Miocene uplift of the Andes (Encinas et al. 2006). New ﬁndings by 1039 Garzione et al. (2008) and Hoke and Garzione (2008) based on isotope data, 1040 suggest that the Andes elevation remained relatively stable for long periods 1041 (tens of millions of years), separated by rapid (1 to 4 million years) changes 1042 of 1.5 km or more. 1043

1044

1045 Most families already present in the mid to late Miocene continue to be present

1046 during the Pliocene (5.33–1.81 mya) (e.g. Arecaceae, Lauraceae, Myrtaceae,

1047 Anacardiaceae, Asteraceae, Chenopodiaceae/Amaranthaceae). Several families like

1048 the Fabaceae increasingly diversiﬁed (Barreda et al. 2007).

1049 Southern South America was dominated by grasslands, steppes, and shrublands,

1050 with rainforests restricted to the moist temperate forests of south-western Patagonia

1051 (Dowsett et al. 1999; Haywood et al. 2002) (Fig. 1.19). Barreda et al. (2007) ◦ 1052 recognize a neotropical palaeo-ﬂoristic province from 32 S to the north and a proto-

1053 espinal/steppe province to the south, together with a Nothofagacean province at the

1054 southwest.

1055 The Cerro Centinela palaeoﬂora in Central Chile contains representatives of

1056 more than 20 modern families of different afﬁnities: tropical genera not found in

1057 Chile any longer (e.g. Nectandra Ocotea, Miconia), subtropical genera (Schinus,

1058

1059

1060

1061

1062

1063

1064

1065 Tropical everwet 1066

1067 Subtropical summerwet 1068

1069 Woody savannah

1070 Desert (hyperarid / arid) 1071

1072 Cool temperate 1073

1074 Cold temperate

1075 Ice sheet 1076

1077 Andes 1078

1079 Fig. 1.19 Pliocene biomes (adapted from Dowsett et al. (1999) and Haywood et al. (2002), on a

1080 TimeTrek 4.2.5 model, CambridgeUNCORRECTED Paleomap Services) PROOF 1.2 Past Climate and Vegetation 27

1081 Schinopsis, Acacia) and temperate or austral genera (Araucaria sección Eutacta, 1082 Nothofagus) (Troncoso and Encinas 2006). Appealing is the presence of the 1083 fern genus Dicksonia, found today mainly in Australasia (Malesia, New Guinea, 1084 Australia, New Caledonia) and in Juan Fernández (Chap. 5). 1085 The end of the Cenozoic, traditionally treated as the “Quaternary”, has been 1086 divided into two epochs: the Pleistocene (1.8 mya to 11,500 year BP) and the 1087 Holocene (11,500 year BP to the present). The Pleistocene is vastly known as the 1088 most recent epoch of glaciations. The last Pleistocene glaciation cycle is known 1089 from southern Chile as the Llanquihue glaciation, which is correlated with the 1090 Wisconsin/Weichselian glaciations in the northern hemisphere, according to global 1091 cooling data (Andersen et al. 1995;Lowelletal.1995; Moreno et al. 2001). 1092 In Chile glaciations affected to some extent all the ecosystems ranging from 1093 the arid north to the humid south. Several proxy-data used for Quaternaty palae- 1094 oreconstructions are speciﬁc for each environment: rodent middens in the north 1095 (Betancourt and Saavedra 2002; Maldonado et al. 2005), tree rings in central Chile 1096 (Barichivich et al. 2009), and sediment cores containing fossil pollen in the for- 1097 merly glaciated south (Heusser 2003; Moreno 2004)(Box 1.6). The pollen analysis 1098 is complemented with the study of macrofossils (leaves), beetles (e.g. Ashworth 1099 et al. 1991), and more recently, chironomid stratigraphies (Massaferro et al. 2009).

1100

1101

1102 Box 1.6 Six Steps for Palaeoenvironmental Reconstruction

1103

1104 Hereafter the main steps for getting pollen sample cores are brieﬂy exposed

1105 (arbitrary extracted from Heusser 2003, Chap. 10):

1106

1107 1. Select a suitable site for sampling, on the base of aerial photographs and

1108 topographic maps, referring to the glacial borders and little sedimentary

1109 bogs or mires. Take account of accessibility for heavy coring equipment.

1110 2. Get a piston sampler equipped with core tubes 5 cm in diameter and 1 m

1111 in length, with 1.5 m long extension rods.

1112 3. With the piston sampler managed by three to four people, get the samples.

1113 You may need to build a wooden platform on the bog and a chain host to

1114 lift the sampler to the surface.

1115 4. Extrude increments onto clear plastic, examine and describe them (color,

1116 texture, layers) and wrap them in aluminum foil. Take multiple cores at

1117 each coring location to ensure overlap at core breaks.

1118 5. Once in the laboratory, identify the pollen grains under a microscope, at

1119 every < 5 cm interval in every sample core.

1120 6. Voilá! You are ready to begin your own palaeoenvironmental

1121 reconstruction.

1122

1123 Note: Calvin and Linda Heusser, together with an international research team,

1124 worked for more than 40 years in southern Chile. They could get thousands

1125 UNCORRECTED PROOF 28 1 The Extravagant Physical Geography of Chile

1126

1127 of samples from 50 coring sites to reconstruct the glacial history and discern 1128 the palaeoecological factors responsible for vegetation changes over 50,000 1129 years.

1130

1131

1132

1133

1134

1135

1136

1137

1138

1139

1140

1141

1142

1143

1144

1145

1146

1147 ◦ 1148 Glaciation effects were especially drastic from 42 (Chiloé) southward, were 1149 glaciers and ice lobes virtually devastated the temperate forests at the Last Glacial 1150 Maximum (LGM) between 29,400 and 14,450 year BP (Fig. 1.20). Vivid remnants 1151 of this widespread glaciation are the Campo de Hielo Patagónico Norte and Campo 1152 de Hielo Patagónico Sur, together with Cordillera de Darwin in southernmost 1153 Patagonia (Fig. 1.20). 1154 At the LGM, periglacial effects like solifluction and glaciofluvial activity also 1155 should have affected the Andes, the longitudinal depression, and the coastal ◦ 1156 Cordillera between 39 and 43 , affecting principally the Valdivian and evergreen 1157 northpatagonian forests (Heusser 2003). 1158 Glacial conditions forced forest formations to migrate equatorward and tree- 1159 lines to lower in altitude (Villagrán et al. 1998; Heusser 2003). Vegetation close 1160 to the glaciated areas was structurally open, forming a steppe-tundra and turning 1161 to parkland and open woodland towards north-central Chile. In the northern part ◦ 1162 of the Central Depression (Tagua Tagua, 34.5 S), at ~14,500 year BP, Lateglacial 1163 warmth and dryness induced the retreat of Nothofagus-Prumnopitys woodland first 1164 by a spread of grassland and ultimately by herb-shrub communities composed by 1165 xeric Amaranthaceae and Asteraceae (Heusser 1997). The presence of Nothofagus 1166 dombeyi type pollen until ~10,000 year BP in the Central Depression exemplifies the 1167 downward altitudinal migration of taxa: this species is today restricted to the Andes 1168 at this latitude, which is also its northern distribution limit (see Sect. 9.1, Fig. 9.7). 1169 Similar situation was suffered by conifers in the south: the current disjunct range of 1170 several species in bothUNCORRECTED cordilleras is a relict of a formerly wider distributionPROOF (before 1.2 Past Climate and Vegetation 29

1171 68° 1172 76°

1173

1174

1175 –30°

1176

1177

1178 LGM ice extension 1179

1180 Campos de Hielo

1181

1182

1183

1184 Distribution 1185 PastNew –40° 1186 Drapetes 1187

1188 Huperzia

1189

1190

1191

1192 1193 1 1194

1195

1196 2 –50° 1197

1198

1199

1200 0 250 500 km 1201 3 1202

1203

1204 1205 Fig. 1.20 Maximal extension of the last cycle of the Llanquihue glaciation (after Denton et al. 1206 1999; Heusser 2003). Remnants of the Pleistocene glaciations: (1) Campo de Hielo Patagónico 1207 Norte, (2) Campo de Hielo Patagónico Sur, (3) Cordillera de Darwin. Also shown is the past and

1208 current distribution of Huperzia fuegiana and Drapetes muscosus (adapted from Heusser (2003) and Moore (1983), and collections of the National Herbarium SGO) 1209

1210

1211

1212

1213

1214

1215 UNCORRECTED PROOF 30 1 The Extravagant Physical Geography of Chile

1216 the colder period at 30,000–14,000 year BP), as shown by the (fossil) presence of 1217 Fitzroya and Pilgerodendron in the Central Depression (Villagrán et al. 2004). 1218 Termination of the last glaciation was (differentiated locally) more or less at 1219 15,000 year BP. Subantarctic species at low altitude in Los Lagos-Chiloé region, 1220 Like Lepidothamnus fonkii (Podocarpaceae), Astelia pulima (Asteliaceae) and 1221 Donatia fascicularis (Stylidiaceae), migrated to higher altitudes. Other species like 1222 Huperzia fuegiana (Lycopodiaceae) and Drapetes muscosus (Thymelaeaceae) were 1223 pushed to the south and are today restricted to southernmost Patagonia or Fuegia 1224 (Fig. 1.20). 1225 The impacts of the Quaternary glaciations were not restricted to the southern 1226 Andes at all, and affected also the vegetation in the Central Chilean Andes (Villagrán 1227 et al. 1998). The changes on vegetation associated to the last glaciations cycle, as 1228 emphasized by these authors are:

1229

1230 (1) Existence of disjuct populations of conifers (Fitzroya, Araucaria, Prumnopitys, 1231 ◦ 1232 Austrocedrus) on the Costal cordillera between 37 and 42 S, dissociated from

1233 the main distribution on the Andes (see maps in Sect. 2.2). These are remnants

1234 of a widespread distribution of these species in the central depression when the

1235 climate was cooler. Same for the Magellanic moorlands located on the summits of the Cordillera de la Costa. 1236 ◦ 1237 (2) Current concentration of endemic and monotypic species between 36 and 40 S

1238 has been interpreted as the function of refugia for laurifolious forests during the

1239 Pleistocene.

1240 (3) High level of endemism at species level showed by the extant high Andean ﬂora

1241 of central Chile is the consequence of repeated cycles of isolation associated to

1242 the adavance of Andean glaciers during the Pleistocene. (4) The current presence of Andean scrub communities in the Coastal cordillera 1243 ◦ 1244 between 32 and 34 , could be the expression of a range expansion of these

1245 communities experimented during the last glaciaton.

1246

1247 The Andes of Central Chile were locally glaciated on the tops of the mountains, 1248 and it is presumed that the Andean ﬂora descended altitudinally and also migrated 1249 northwards during the Pleistocene (Simpson 1971; Villagrán et al. 1998). 1250 Especially the altitudinal migrations are hypothesised to have affected commu- 1251 nities as a whole, but there is also possible that there was a recombination and 1252 redistribution of the taxa forming today these communities (Patricio Moreno pers. 1253 comm.). Whether the Quaternary glaciations affected only the distribution ranges 1254 or had deeper evolutionary implications is still a matter of discussion (Box 1.7). 1255 As example, as the glacial tongues advanced down from the Andes into the cen- 1256 tral depression, valleys like the Río Maule (36◦S) and Río Biobío (37◦ to 38◦S) 1257 could have acted as barriers, interrupting gene ﬂow between plant populations 1258 and communities, as resulting from the study of Hypochaeris acaulis populations 1259 (Tremetsberger et al. 2003). 1260 UNCORRECTED PROOF 1.3 Current Climate and Vegetation 31

1261 1262 Box 1.7 How to Survive a Glaciation? The Refugial Debate 1263 1264 The Campos de Hielo Norte and Sur are considered the biggest inland ice- 1265 caps after Greenland, current reminders of the maximal extension of the 1266 Pleistocene glaciations. Figure 1.20 suggests almost complete depletion of the 1267 southern biota at the LGM. But the relative rapid reoccupation of deglaciated 1268 areas under warmer conditions by the rainforest taxa suggests the continued 1269 permanence of exemplars somewhere not so far from the glacial lobes. 1270 Debate continues on possible impacts from the ice ages and possible loca- 1271 tions of refugial sites in the Southern Andes (Knapp and Mallet 2003). As 1272 2/3 of the actual area of the southern forests was depleted, the traditional 1273 view proposes that taxa mostly survived the glaciations in the foreland of the 1274 glaciers and on several nunataks. This view has been recently challenged by 1275 Fickert et al. (2007), who suggest, based on research on six active glaciers 1276 (e.g. Monte Tronador in southern Chile), that the size of possible refugia 1277 would be considerably enlarged if debris-covered glaciers are considered. 1278 The nunatak theory just offers a too small area for a survival of viable plant 1279 populations (Fickert et al. 2007). Debris-covered glaciers should be added to 1280 the recent systematization of three main types of glacial refugia proposed by 1281 Holderegger and Thiel-Egenter (2009): i.e. nunatak, peripheral and lowland 1282 refugia. Concrete results provided by Premoli et al. (2000) suggest that the 1283 populations of Fitzroya cupressoides survived the Last Glacial Maximum in 1284 multiple refugia rather than in only one refugium, such as an ice-free area of 1285 coastal Chile (Single Refugium hypothesis). Multiple refugia in the eastern 1286 side of the Cordillera are also hypothesized for the survival of Austrocedrus 1287 chilensis (Pastorino and Gallo 2002). 1288 Pleistocene and Holocene changes have disrupted species ranges, extir- 1289 pated local populations, and changed selective pressures (Premoli et al. 2000), 1290 but it is doubtful that they affected speciation processes. Some authors have 1291 emphasized the role of the last glaciations in speciation, but others call this a 1292 “failed paradigm” (Klicka and Zink 1997). It is possible that molecular stud- 1293 ies of populations of Quaternary species help deﬁne the relict characteristic of 1294 these species (Willis and Niklas 2004) (see Sect. 9.6). 1295

1296

1297

1298

1299 1.3 Current Climate and Vegetation

1300

1301 A few crucial features of the present climate and vegetation will be outlined here, as 1302 a detailed description of these aspects is beyond the scope of this book. The reader 1303 is redirected to the most updated references on these topics, especially Luebert and 1304 Pliscoff (2006), Veblen et al. (2007), Garreaud et al. (2009). A ﬁne synthesis of the 1305 southern Andean vegetationUNCORRECTED is also provided by Heusser (2003). PROOF 32 1 The Extravagant Physical Geography of Chile

1306 South America is situated within the influence of the Intertropical Convergence 1307 Zone (ITCZ) and related circulation systems (Orme 2007). This band is a major 1308 feature of the global circulation and the Chilean climate certainly depends upon 1309 this regional situation. The yearly N-S displacement of the Subtropical Anticyclone 1310 (South Pacific High) is one of the principal factors affecting the climatic latitudinal 1311 gradient in Chile (Garreaud and Muñoz 2004;Emcketal.2006) (Fig. 1.21). 1312 The displacement of the South Pacific High towards the south during the austral 1313 summer promotes Mediterranean-type climatic conditions in Central Chile (Luebert 1314 and Pliscoff 2006). During this displacement, tropical rainfall can reach the north- ◦ 1315 ern Altiplano, but this influence decreases at around 23 S (Vuille and Baumgartner 1316 1998) due to the influence of the Andes as a “climatic wall” (Garreaud and Aceituno 1317 2007). The displacement of the Anticyclone towards the North in the austral winter 1318 allows the entrance of the westerlies from the SW, promoting an intense and reg- ◦ 1319 ular rainfall period with a maximal influence at around 47 S (Luebert and Pliscoff 1320 2006). South of this latitude, the influence of the high pressures located around 1321 Antarctica generates a decrease in precipitation (Endlicher and Santana 1988). From 1322 Antarctica comes another important feature of the Chilean climate, i.e. the cool 1323 Humboldt Current derived from the Antarctic Circumpolar Current (Orme 2007). 1324 The Humboldt Current is characterised by a predominant northward flow of sur- 1325 face waters of subantarctic origin and by strong upwelling of cool nutrient-rich 1326 subsurface waters of equatorial origin (Thiel et al. 2007). The current produces a 1327 decrease in the coastal zone temperatures in North- and Central Chile, and aided 1328 by tropospheric subsidence creates a temperature inversion that increases the con- 1329 ditions of aridity in western South America from near the equator to beyond the

1330

1331 1332 January (southern summer) July (southern winter) 1333

1334

1335

1336

1337

1338

1339

1340

1341

1342

1343

1344

1345

1346 South Pacific and South Atlantic Highs Relative position of the ITCZ

1347 Polar front Surface dominant winds

1348

1349 Fig. 1.21 Atmospheric circulation over South America. Note the yearly N–S displacement of the

1350 Intertropical ConvergenceUNCORRECTED Zone (ITCZ) (adapted from Emck et al. 2006) PROOF 1.3 Current Climate and Vegetation 33

1351 Tropic of Capricorn (Trewartha 1961;Orme2007). Regarding precipitation, these 1352 factors generate a gradient of decreasing rainfall from the SW towards the NE, from 1353 ca. 5,000 mm in the SW to 0 mm in the Atacama (Fig. 1.22a). Only at the north- 1354 ern Altiplano this tendency is reverted by the tropical rains reaching northern Chile 1355 from the Amazonas basin. 1356 On the base of the early climatic classification done by W. Köppen 1357 (1930), Chilean bioclimatic classifications were progressively developed by Di 1358 Castri (1968), Quintanilla (1974), Di Castri and Hajek (1976), and Amigo and 1359 Ramírez (1998). Coupling the pioneer plant geographical works of Reiche (1907) 1360 and Pisano (1954), with the bioclimatic classifications and phytosociological infor- 1361 mation (e.g. Oberdorfer 1960), several attempts have been made for a vegetation 1362 classification of the country, like the ones from Schmithüsen (1956), Quintanilla 1363 (1983), and Gajardo (1994). 1364 A systematic revision of previous classification schemes integrated to the analy- 1365 sis of global climatic surfaces on a GIS-based platform allowed Luebert and Pliscoff 1366 (2006) to generate the most accurate bioclimatic and vegetation synthesis to date. 1367 The classification considers bioclimatic and floristic data on the base of the concep- 1368 tual framework promoted by Rivas-Martínez and Rivas-Sáenz (1996–2009). The 1369 application of the method for Chile resulted in the identification of 17 vegetation 1370 formations and 127 vegetational belts (Luebert and Pliscoff 2006). The highest rank 1371 of the bioclimatic classification is the “macrobioclimate”, and the five units that 1372 exist worldwide are found in Chile (Fig. 1.22b):

1373 ◦ 1374 (a) Tropical macrobioclimate: it extends from the border with Perú at 17 35 S ◦ ◦ 1375 towards the south till a diagonal limit at 23 S at the coast and at 31 S in the high 1376 Andes. The southern limit reflects on the one side the maximal influence of the 1377 polar fronts coming from the SW (Fig. 1.22b) and on the other side the maximal 1378 influence of the tropical moisture that reaches the Altiplano and the north- 1379 ern high Andes during the austral summer. Within this zone lies the Atacama 1380 Desert. 1381 (b) Mediterranean-type macrobioclimate: this macrozone is one of the most char- 1382 acteristic features of Chile, as one of only five regions at the global scale that 1383 share this type of climate, characterized mainly tdue to the marked seasonality in 1384 the thermal and precipitation regimes (Di Castri 1981). The limits of this mac- 1385 robiolimate in Chile have been controversial, but the proposal of Luebert and 1386 Pliscoff (2006) is concordant with the findings of Amigo and Ramírez (1998). It 1387 extends from the diagonal limit with the tropical bioclimate towards the South, ◦ ◦ 1388 till 37 S at coast and Andes, and till 39 S in the Central Depression. The 1389 Mediterranean-type macrobioclimate appears also in disjunct patches further ◦ 1390 South around 46–47 S, related to the western limit of the Patagonian steppe. 1391 (c) Temperate macrobioclimate: it occupies the major area in continental Chile, ◦ 1392 from the limit with the Mediterranean-type one at 37–39 Suptowestern 1393 Patagonia and the Magallanes region in the southern territories. It is thermally 1394 most homogeneous and precipitation can reach more than 5,000 mm on the 1395 southwestern fjordlandUNCORRECTED and island groups more exposed to the PROOF humid westerlies. 34 1 The Extravagant Physical Geography of Chile

1396

1397

1398

1399 20° 1400

1401

1402 a 2 1 1403

1404

1405

1406 3 1407

1408 30° 4 1409 5 1410 b 1411

1412 6 1 1413

1414 <10 1415 7 10–50 1416 40°

1417 50–200

1418 200–500 8a 1419 500–1000 1420 1000–3000 8b 1421 3000–5000 c 1422 > 5000 1423 12

1424 50° 1425 10

1426

1427 11 d 1428 9

1429 1430 e 1431 abc 1432 Fig. 1.22 a Annual precipitation, based on Schmithüsen (1956) updated with current available 1433 data; b ﬁve Chilean macro-bioclimates (adapted from Luebert and Pliscoff 2006); c distribu- 1434 tion of vegetation formations according Schmithüsen (1956): 1= Andean vegetation; 2= desert = = = 1435 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- 1436 forest; 8b= northpatagonic rainforest; 8c=subantarctic rainforest; 9= subantarctic moorlands; 1437 10=subantarctic deciduous forest, 11=east-patagonic steppe; 12=Campos de Hielo 1438

1439

1440 UNCORRECTED PROOF 1.3 Current Climate and Vegetation 35

1441 (d) Antiboreal macrobioclimate: it occupies a restricted portion of the southernmost 1442 continental extreme, affecting the Magellanic archipelagos and the southern 1443 part of Tierra del Fuego. Precipitation decreases notably toeads the east and 1444 temperature decreases towards the south. 1445 (e) Polar bioclimate: with increasing thermal cold conditions, it occupies the 1446 territory outside the American continent towards Antarctica.

1447

1448 Schmithüsen (1956) provided one of the most sinthetical and comprehensive 1449 accounts of the Chilean vegetation, valid till today (Fig. 1.22c). He also illustrated 1450 magistrally the latitudinal versus altitudinal distribution of the vegetation formations 1451 (Fig. 1.23). 1452 The principal vegetation formations and their main characteristics will be brieﬂy 1453 described here, based on Schmithüsen (1956) and Luebert and Pliscoff (2006) 1454 (Figs. 1.22c, 1.23 and Fig. 1.24 (col. sect.)).

1455 ◦ 1456 (a) The hyperarid desert formation or desert core extends from 18 S along the ◦ 1457 coast and interior zones towards the south till around 24 . Approaching 1458 the border with Peru, vegetation is restricted to the deep valleys of Azapa 1459 and Camarones, related to agriculture. At the heart of the Atacama, vegeta- 1460 tion is almost completely lacking; nevertheless, there are stands of natural 1461 and human induced forests of Prosopis tamarugo = Pampa del Tamarugal. 1462 Towards the Andes, scrub vegetation consists mainly of a low scrub (mator- 1463 ral) of Adesmia atacamensis, Cistanthe salsoloides, Atriplex imbricata, and 1464 Acantholippia deserticola. In a very thin belt between 2,000 and 2,800 m asl, 1465 big cacti of Browningia candelaris bear out the landscape (Fig. 7.1 (col. sect.)). 1466 (b) The sparse coastal shrub vegetation, also characteristic for coastal southern 1467 Peru, is called the “Lomas” formation, and consists of a rich assemblage of 1468 Eulychnia, Nolana, Heliotropium, Tetragonia, and Euphorbia species. This for- 1469 mation is highly dependent on fog and humidity to some extent related to the 1470 El Niño phenomenon (Box 7.1).

1471

1472 North Snow limit Upper vegetation limit South 1473 6000 5000 1474 4000

1475 3000 2000 1476 1000 1477 18 20 22 24 26 2830 32 34 36 38 40 42 44 46 48 50 52 54 56 1478

1479 Andean vegetation Desert scrub Valdivian forest North Patagonian rainforest 1480 Pre-Andean scrub Xerophytic scrub Austrocedrus chilensis Subantarctic rainforest and moorlands 1481 Sclerophyllous matorral Araucana araucana Browningia candelaris stands Patagonian steppe 1482 Deciduous forest Desertic core Fitzroya cupressoides 1483

1484 Fig. 1.23 Distribution of vegetation formations along the altitudinal proﬁle according

1485 Schmithüsen (1956) UNCORRECTED PROOF 36 1 The Extravagant Physical Geography of Chile

1486 a b 1487

1488

1489

1490

1491

1492

1493 1494 c d 1495

1496

1497

1498

1499

1500

1501 1502 f 1503

1504

1505

1506

1507

1508

1509

1510

1511 1512 e 1513

1514

1515

1516

1517

1518

1519

1520 Fig. 1.24 Chilean vegetation: a high-Andean vegetation, Azorella compacta at Parinacota 1521 (18◦30S); b high-Andean vegetation, vegas de Zapaleri, limit to Bolivia at 22◦50S; c Puya 1522 coerulea, characteristic of the xeric matorral at Cerro La Campana (33◦S); d slerophyllous for- ◦ ◦ 1523 est at Río Clarillo (33 47 S); e temperate forest at Navarino island (55 S); f conifer forest ◦ 1524 (Araucaria) with Nothofagus at El Cañi, Pucón (39 20 S). Photo credits: a Walter Welss; b–f

1525 A. Moreira-Muñoz

1526

1527

1528

1529

1530 UNCORRECTED PROOF 1.3 Current Climate and Vegetation 37

1531 The desert coastal scrub extends into the land’s interior between 24 and ◦ 1532 32 S, generating a transition zone from the desert towards the winter-wet 1533 Mediterranean climate of Central Chile. It encompasses an open scrub (mator- 1534 ral) composed by Adesmia spp., Bulnesia, Balbisia, and Heliotropium species. ◦ 1535 At 30 S, the vegetation gradually changes to a xerophytic scrub composed of 1536 Haplopappus spp., Porlieria chilensis, Flourensia thurifera, Colliguaja odor- 1537 ifera, Trichocereus cacti, and Puya species. This zone harbours one of the 1538 most interesting botanical phenomena, the “desierto florido” (Box 7.1), and a 1539 plant geographical icon = the fog forest of Fray Jorge, where the northernmost 1540 remnants of southern floristic elements are to be found (see Box 3.1). 1541 (c) The Andean vegetation is the formation occupying extreme, high environments, ◦ ◦ 1542 ranging from 17 30 S to ca. 40 S along the western Andean slope. This wide 1543 latitudinal extension encompasses a very different composition along the North- 1544 South profile, and in the altitude. The intermediate altitudinal belts show the 1545 structurally most developed vegetation: the lowest belts are affected by the 1546 aridity, the highest by low temperature. These intermediate belts are composed 1547 by tolares, dominated by Parastrephia species and pajonales (cushion grasses) 1548 of Festuca orthophylla. Stands of Polylepis tarapacana trees are to be found. 1549 Precordilleran belts are dominated by Fabiana ramulosa and Diplostephium 1550 meyenii. Above 4,700 m asl bofedales (high Andean wetlands) support the long 1551 tradition of Andean llama and alpaca pasture. Approaching the most arid part 1552 of the Atacama Desert there are shrubs of Fabiana and Baccharis, together with 1553 Atriplex and Acantholippia in the lower belts. Towards the south, sparse vege- 1554 tation is composed of Jarava frigida cushion grasses, several Adesmia species, 1555 with the addition of Mulinum, and Urbania species. The treeline changes con- 1556 stantly along the latitude gradient: in the north it is composed by queñoales = 1557 Polylepis tarapacana and P. rugulosa, in the central-north it is replaced by ◦ 1558 Adesmia shrubs with Ephedra between 31 and 34 S the treeline reappears 1559 by means of Kageneckia angustifolia accompanied by Guindilia trinervis. 1560 At this latitudinal range the Andean scrub is composed of Adesmia species, 1561 Tetraglochin alatum, Mulinum spinosum, and cushion Apiaceae like Azorella ◦ 1562 spp. and Laretia acaulis.From3260 to the South, at the lower limit of the 1563 Andean formation, appears the conifer Austrocedrus chilensis (Fig. 2.20). The ◦ 1564 Andean formation disappears at around 37 S, where it is replaced by deciduous 1565 forests. 1566 (d) Entering into the Mediterranean climate zone the vegetation changes to a sclero- 1567 phyllous high scrub or matorral esclerófilo. On favorable South oriented slopes 1568 this scrub shows characteristics of woodland, with trees reaching 20–25 meter 1569 in height. Typical species of the Central Chilean matorral are Peumus boldus, 1570 Cryptocarya alba, Quillaja saponaria, Maytenus boaria. In the quebradas, 1571 i.e. more humid stands, there appears a more hygrophyllous forest composed 1572 of Crinodendron patagua, Beilschmiedia miersii, Drimys winteri, and Persea 1573 lingue. In contrast, the most exposed and plain areas contain a woody savanna 1574 (espinal) mainly composed of Acacia caven and Prosopis chilensis.North

1575 UNCORRECTED PROOF 38 1 The Extravagant Physical Geography of Chile

1576 exposed slopes show a rich array of annual species and characteristic bromeli- 1577 ads, Puya chilensisP. berteroniana and P. coerulea, together with the cactus 1578 Trichocereus chiloensis. ◦ 1579 (e) Around 33 the coastal cordillera reaches far inside the continent, and above 1580 1,200 m asl, the sclerophyllous woodland leaves space for a deciduous forest 1581 composed of deciduous Nothofagus species. The northernmost populations at ◦ 1582 33 S seem to be remnants of an ancient distribution of the genus (Chap. 9). 1583 Deciduous forests dominate along the Andes and the coast towards the South, 1584 surrounding the Central Depression. The core of the deciduous forest between ◦ 1585 35 and 36 S is known as the maulino forest, a mesic forest type, dominated by 1586 the two broadleaved deciduous species Nothofagus alessandrii and N. glauca ◦ 1587 (San Martín and Donoso 1996). At around 38 S, this forest shows signs of 1588 the transition towards the temperate macrobiolimate, with the remarkable pres- 1589 ence of the resinous or conifer forests of Araucaria araucana at the coast 1590 (Nahuelbuta) and the Andes. Deciduous forests turn often into a krummholz 1591 of Nothofagus antarctica and N. pumilio composes the treeline along the Andes 1592 all the way to the Cape Horn. 1593 (f) Located well into the temperate macrobioclimate, and related to high precip- 1594 itation levels (>2,000 mm/year) is the broad-leaved (laurifolious) forest, also 1595 known as the Valdivian forest. It shows, same as the maulino forest, a “U” ◦ ◦ 1596 shape with a coastal and an Andean branch between 39 and 42 S. The his- 1597 tory of this forest has been vastly debated, and some of its components, like 1598 Aextoxicon punctatum, Laureliopsis philippiana, Dasyphyllum diacanthoides 1599 (Fig. 8.2 (col. sect.)), Luma apiculata, Laurelia sempervirens, Eucryphia cordi- 1600 folia, and Weinmannia trichosperma seem to be old remnants of Palaeogene 1601 floras (Sect. 1.2). ◦ ◦ 1602 (g) At around 41 S on the Andes and 41 30 on the coast, broad-leaved forests 1603 are replaced by an evergreen northpatagonian rainforest mainly composed of 1604 large trees pertaining to the Nothofagaceae: Nothofagus dombeyi, N. nitida, and 1605 N. betuloides. These rainforests are intermingled with the conifer forests of 1606 Podocarpus nubigenus, Fitzroya cupressoides and Pilgerodendrum uviferum. 1607 These evergreen forests dominate at the coast and interior, being replaced in 1608 altitude by the deciduous forest. 1609 (h) As the landscape gets more and more fragmented into fjords and little islands ◦ 1610 south of 47 , and the precipitation exceeds the 4,000 mm/year the vegeta- 1611 tion turns to a low physiognomy of moorlands, dominated by Astelia pumila, 1612 Donatia fascicularis, Oreobolus obtusangulus. Towards the East the moorlands 1613 get less humid and dominated by the moss Sphagnum magellanicum.Mostof 1614 the interior of Patagonia is covered by the two wide icefielkds Campo de Hielo 1615 Norte and Campo de Hielo Sur. To the South of this last icefield, the decid- 1616 uous forest of Nothofagus reappears, together with the subantarctic evergreen 1617 rainforest. In accordance with the marked precipitation gradient ranging from 1618 4,000 mm at the western side to 300 mm at the eastern side of the low Andes in 1619 southern Patagonia and Tierra del Fuego, a gramineous steppe of Festuca spp. 1620 dominates the landscape.UNCORRECTED PROOF References 39

1621 The scheme presented here, based on Schmithüsen (1956) and Luebert and Pliscoff 1622 (2006) corresponds to the potential vegetation, but all the formations and most of the 1623 vegetation belts that compose the Chilean vegetation are to a high degree affected by 1624 the long history of human occupation, from localized mining impacts in the north to 1625 extended forest substitutions in the south. The core of the deciduous forest at around ◦ ◦ 1626 38 Sto41 S has been transformed into agriculture, and in the Central Derpression 1627 only remnants of sclerophyllous forests remain in this mainly cultural landscape 1628 (Chap. 6).

1629

1630

1631 1632 References 1633 1634 Amigo J, Ramírez C (1998) A bioclimatic classiﬁcation of Chile: woodland communities in the 1635 temperate zone. Plant Ecol 136:9–26

1636 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 1637 and 42◦30s latitude: a short review of preliminary results. Quat Int 28:199–201 1638 Anderson JM, Anderson HM, Archangelsky S, Bamford M, Chandra S, Dettmann M, Hill R, 1639 McLoughlin S, Rösler O (1999) Patterns of Gondwana plant colonisation and diversiﬁcation. 1640 J Afr Earth Sci 28:145–167

1641 Artabe AE, Morel EM, Spalletti LA (2003) Characterization of the phytogeographic Triassic provinces from extratropical Gondwana. Ameghiniana 40:387–405 1642 Artabe AE, Ganuza DG, Spalletti LA, Zúñiga A, Morel EM (2005) Revisión de la paleoﬂora 1643 del cerro La Brea (Jurásico Temprano), provincia de Mendoza, Argentina. Ameghiniana 42: 1644 429–442 1645 Ashworth AC, Markraf V, Villagrán C (1991) Late Quaternary climatic history of the Chilean

1646 Channels based on fossil pollen and beetle analysis, with an analysis of the modern vegetation and pollen rain. J Quat Sci 6:279–291 1647 Astini RA, Benedetto JL, Vaccari NE (1995) The early Paleozoic evolution of the Argentine 1648 Precordillera as a Laurentian rifted, drifted, and collided terrane: a geodynamic model. Geol 1649 Soc Am Bull 107:253–273 1650 Barichivich J, Sauchyn DJ, Lara A (2009) Climate signals in high elevation tree-rings from

1651 the semiarid Andes of north-central Chile: responses to regional and large-scale variability. Palaeogeogr Palaeoclimatol Palaeoecol 281:320–333 1652 Barreda V, Archangelsky S (2006) The southernmost record of tropical pollen grains in the mid- 1653 Cretaceous of Patagonia, Argentina. Cretaceous Res 27:778–787 1654 Barreda VD, Anzótegui LA, Prieto AR, Aceñolaza PG, Bianchi MM, Borromei AM, Brea M et al. 1655 (2007) Diversiﬁcación y cambios de las angiospermas durante el Neógeno en Argentina. In: ◦ 1656 Archangelsky S, Sánchez T, Tonni EP (eds) Ameghiniana 50 aniversario, Publicación Especial 11:173–191 1657 Barrientos SE (2007) Earthquakes in Chile. In: Moreno T, Gibbons W, (eds) The geology of Chile. 1658 Geological Society, London, pp 263–287 1659 Beerling DJ (2002) Low atmospheric CO2 levels during the Permo-Carboniferous glaciation 1660 inferred from fossil lycopsids. Proc Natl Acad Sci 99:12567–12571

1661 Benton MJ, Twitchett RJ (2003) How to kill (almost) all life: the end-Permian extinction event. Trends Ecol Evol 18:358–365 1662 Berry EW (1922a) Contributions to the paleobotany of Peru, Bolivia and Chile. The John Hopkins 1663 Press, Baltimore. 1664 Berry EW (1922b) The ﬂora of Concepción-Arauco coal measures of Chile. Stud Geol (John 1665 Hopkins Univ) 4:73–142UNCORRECTED PROOF 40 1 The Extravagant Physical Geography of Chile

1666 Betancourt JL, Saavedra B (2002) Rodent middens, a new method for Quaternary research in arid

1667 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 1668 (Paleoceno superior), Puerto Visser, Chubut, Argentina. Ameghiniana 43:139–146 1669 Cecioni G, (ed) (1968) El Terciario de Chile: zona central. Editorial Andrés Bello, Santiago. 1670 Charrier R (1988) Condiciones Paleoclimáticas para el Carbonífero Superior y Pérmico Inferior en 1671 la mitad austral de América del Sur. Bol Mus Nac Hist Nat (Chile) 41:105–116

1672 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, 1673 pp 21–116 1674 Chumakov NM, Zharkov MA (2003) Climate of the Late Permian and Early Triassic: general 1675 inferences. Stratigr Geol Correl 11:361–375 1676 Convey P, Gibson JAE, Hillenbrand C-D, Hodgson DA, Pugh PJA, Smellie JL, Stevens MI (2008)

1677 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 1678 America and their relationship with other ﬂoral realms. Abstracts, 28th International Geological 1679 Congress, p 351 1680 Cúneo RN, Johnson K, Wilf P, Gandolfo MA, Iglesias A (2007) A preliminary report on the diver- 1681 sity of Latest Cretaceous ﬂoras from Northern Patagonia, Argentina. Abstract GSA Denver

1682 Annual Meeting Denton GH, Heusser CJ, Lowell TV, Moreno PI, Andersen BG, Heusser LE, Schlüchter C, 1683 Marchant DR (1999) Interhemispheric linkage of paleoclimate during the last glaciation. Geogr 1684 Ann 81A:107–153 1685 Di Castri F (1968) Esquisse écologique du Chili. In: Delamanre-Debouteville C, Rapoport E, (eds) 1686 Biologie del’ Amérique Australe, 4. Editions du Centre National de la Recherce Scientﬁque,

1687 Paris, pp 6–52 Di Castri F (1981) Mediterranean-type shrublands of the world. In: Di Castri F, Goodall DW, 1688 Specht RL, (eds) Mediterranean-type Shrublands. Ecosystems of the World vol 11. Elsevier, 1689 Amsterdam, pp 1–52 1690 Di Castri F, Hajek E (1976) Bioclimatología de Chile. Vicerrectoría Académica, Universidad 1691 Católica de Chile, Santiago.

1692 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 1693 Dowsett HJ, Barron JA, Poore RZ, Thompson RS, Cronin TM, Ishman SE, Willard DA (1999) 1694 Middle pliocene paleoenvironmental reconstruction: PRISM2. US Geological Survey Open 1695 File Report, 99–535. http://pubs.usgs . gov/of/1999/of99-535/ 1696 Dunai TJ, González López GA, Juez-Larré J (2005) Oligocene–Miocene age of aridity in

1697 the Atacama Desert revealed by exposure dating of erosion-sensitive landforms. Geology 33:321–324 1698 Dusén P (1907) Über die Tertiäre Flora der Magellansländer. In: Nordenskjold O (ed) 1699 Wissenschaftlichen Ergebnisse der Schwedischen Expedition nach der Magellansländern, 1700 1895–1897, vol I, pp 84–108 1701 Emck P, Moreira-Muñoz A, Richter M (2006) El clima y sus efectos en la vegetación. In: Moraes

1702 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 1703 Encinas A, Maksaev V, Pinto L, Le Roux JP, Munizaga F, Zentilli M (2006) Pliocene lahar deposits 1704 in the Coastal Cordillera of central Chile: implications for uplift, avalanche deposits, and 1705 porphyry copper systems in the Main Andean Cordillera. J S Am Earth Sci 20:369–381 1706 Endlicher W, Santana A (1988) El clima del sur de la Patagonia y sus aspectos ecológicos: un siglo

1707 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 1708 Fickert T, Friend D, Grüninger F, Molnia B, Richter M (2007) Did debris covered glaciers serve 1709 as Pleistocene refugia for plants? A new hypothesis derived from observations of recent plant 1710 growth on Glacier surfaces.UNCORRECTED Arct Antarct Alp Res 39:245–257 PROOF References 41

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1712 southern Chile: new constraints from apatite U-Th/He thermochronology. Abstracts 2007 GSA Denver Annual Meeting 1713 Frakes LA, Francis JE, Syktus JI (1992) Climate modes of the Phanerozoic. Cambridge University 1714 Press, Cambridge. 1715 Fuenzalida H (1938) Las capas de Los Molles. Bol Mus Nac Hist Nat 16:67–92 1716 Fuenzalida H (1966) Historia vegetacional de Chile. Estudios geográﬁcos . Facultad de Filosofía y

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