�

Plant Geography of ChileAn Essay on Postmodern Biogeography

Den Naturwissenschaftlichen Fakultätender Friedrich-Alexander-Universität Erlangen-Nürnberg

zurErlangung des Doktorgrades

vorgelegt vonAndrés Moreira-Muñozaus Los Angeles, Chile

�

Gedruckt mit Unterstützung des Deutschen Akademischen Austauschdienstes (DAAD)

�

Als Dissertation genehmigt von den Naturwissenschaftlichen Fakultäten der Universität Erlangen-Nürnberg

Tag der mündlichen Prüfung: Donnerstag �4. Juni �007

Vorsitzender derPromotioskommision: Prof. Dr. Eberhard Bänsch

Erstberichterstatter: Prof. Dr. Michael Richter

Zweitberichterstatter: Prof. Dr. Tod Stuessy

4

�

A Paola, Silene, Coyán y Sayén

�

Contents Zusammenfassung / Summary...............................................................................................8

� Introduction .......................................................................................................................... ��

�.� Inventing plant geography ............................................................................................ ���.� Chorology and Arealkunde ........................................................................................... ���.� Naming the plant world ................................................................................................ �9�.4 From the map to the tree: cladistics and biogeography ................................................ �4�.� The fragmented map of modern biogeography ............................................................. �7�.� Postmodern biogeography: de(re)constructing the map ............................................... �9

� Chile, a Remote Corner on Earth ......................................................................................... �7

�.� Romancing the South: the discovery of a virgin world ................................................ �72.2 Classification of the Chilean plants: a modern (but not definitive) synthesis ............... 4�2.3 Geographical classification of the Chilean flora ........................................................... 47�.4 Excursus: vegetation maps and Vegetationsbilder ........................................................ �02.5 Geographic ranges in the latitudinal profile .................................................................. ��

� Geographic Relationships of the Chilean Flora ................................................................... 7�

3.1 Pantropical floristic element ......................................................................................... 7�3.2 Australasiatic floristic element ...................................................................................... 7�3.3 Neotropical (American) floristic element ..................................................................... 7�3.4 Antitropical floristic element ......................................................................................... 773.5 South-temperate floristic element ................................................................................. 803.6 Endemic floristic element ............................................................................................. 8�3.7 Cosmopolitan floristic element ..................................................................................... 8�

4 Biogeographic analysis ........................................................................................................ 89

4.�. To be or not to be disjunct ............................................................................................ 904.2 The austral v/s the neotropical floristic realm ............................................................... 9�4.� Analysis of Endemism .................................................................................................. 9�4.4 The disintegration of the endemic and the south-temperate elements ........................ �0�4.5 Plant geography of the Chilean Pacific islands ........................................................... �0�

� Palaeogeography: insights into the Evolution of the Chilean Flora .................................. ��7

�.� Continental movements: fragmenting the Earth’s surface .......................................... ��7�.� Oceanic transgressions over the continental surface .................................................. ����.� The Andean uplift........................................................................................................ ��8�.4 The last �0 000 years: surviving the Ice Age .............................................................. ��95.5 Alternative palaeogeographies: against scientific consensus ...................................... ���

7

6 Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography ................ ��9

�.� Molecular dating, the reigning paradigm .................................................................... ��9�.�. Phylogeny of Chilean plants ...................................................................................... �4��.� Vicariance v/s dispersal in the Chilean Flora .............................................................. �4��.4 Sloppy biogeography v/s harsh geology? ................................................................... �49�.� Species and speciation ................................................................................................ ����.� Re-inventing an origin for the land plants .................................................................. ��7

7 Back to Postmodern Biogeography ................................................................................... ��7

7.� Biogeography as a social science ................................................................................ ��77.� Biogeography toward a science of qualities ............................................................... �78

8 Conclusions: toward a biogeographic synthesis of the Chilean Flora ............................... �8�

Footnotes ............................................................................................................................... �8�

References ............................................................................................................................. �88

Appendices ............................................................................................................................ ���

Appendix A: List of Chilean genera, geographic distribution, floristic elements ............. ���Appendix B: Genera shared by several Chilean regions .................................................. �47Appendix C: Matrix for PAE: Distribution of Endemic Genera ....................................... ���Appendix D: Native Genera in the Chilean Pacific Islands ............................................. ���

Agradecimientos / Danksagung / Acknowledgements ......................................................... ���

Lebenslauf ............................................................................................................................. ��7

8

Zusammenfassung

Die historische Biogeographie hat sich in den letzten Jahrzehnten sowohl theoretisch als auch methodisch stark weiterentwickelt. Konzepte wie Dispersion, Vikarianz oder Panbiogeographie zeigen konstante Synergien im Zuge der Erneuerung von Theorie und Praxis. Die historische Biogeographie muss sich dabei, wie alle Naturwissenschaften, der großen Herausforderung stellen, sich der Moderne und der Postmoderne anzupassen. Die vorliegende Arbeit analysiert die wichtigsten phytogeographischen Gegebenheiten in Chile unter besonderer Berücksichtigung der geographischen Verbreitung von Gattungen und ihrer globalen Beziehungen. Die Auswirkungen dieser Verhältnisse auf die Entwicklung der chilenischen Flora werden dabei besonders unter verschiedenen paläogeographischen Szenarien diskutiert. Außerdem wird die Situation im Kontext der modernen und postmodernen Wissenschaftstheorie betrachtet. Das erlaubt eine Kontext-orientierte Synthese der chilenischen Pflanzengeographie und stellt den Ausgangspunkt für weitere Forschungen in dieser Übergangsdisziplin zwischen Geographie und Biologie.

Schlagwörter: Pflanzengeographie, Panbiogeographie, Postmoderne, Floristische Elemente, Chile.

Summary

Historical biogeography has seen a rapid theoretical and methodological developed in the last decades. Concepts such as dispersal, vicariance, or panbiogeography show a constant synergy in the renewal of the theory and the praxis. The present thesis examines the most important phytogeographical issues in Chile with special consideration of the distribution and geographical relationships of the genera. The consequences of these relations on the evolution of the Chilean Flora are discussed thereby, particularly under different possible palaeogeographical scenarios. In addition the conditions in the context of the modern and postmodern science theory are considered. This permits a context-oriented synthesis of the Chilean plant geography, that set the starting point for further research in this crucial field between geography and biology.

Keywords: Plant Geography, panbiogeography, postmodern science, floristic elements, Chile.

9

� Introduction

�0

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

��

1 Introduction

“Between geographers and historical biogeographers there has been relatively little communication and, like Stoddart [�98�], I consider this a problem. By far the majority of those publishing work with biogeographical themes are biologists. Most biology-trained biogeographers appear to have little or no familiarity with the theoretical, philosophical, and methodological literature of geography; this, at least, seems to be the only conclusion that can be drawn from the almost total absence of referral to such in their papers” (Smith �989).

This is an essay on empirical biogeography. An intend to distinguish, compare, and integrate the geo and the bio.

The first and so far the only Plant Geography of Chile was written �00 years ago by Karl Reiche (�907). The German botanist was appointed at the Museo Nacional de Historia Natural in �89� by Federico Philippi, the son and scientific successor of the great naturalist Rudolf A. Philippi, who arrived in Chile �8�� highly recommended by Alexander von Humboldt. Those were still times of discovery: every plant, every animal, even the rocks needed to be first found, then kept, catalogued, compared, described, illustrated. Those were good times for being a naturalist.

Thanks to the botanical knowledge accumulated in 3 centuries of discovery and classification (see chapter �), plus his incredible capacity, Reiche could publish his Flora de Chile in five volumes (�89�-�9��) and his Plant Geography, this latter requested for the series Die Vegetation der Erde by the German botanists Adolf Engler and Oscar Drude (Reiche �907).

The Flora de Chile is under a new revision in a current effort lead by the Universidad de Concepción, in a plan that comprises six volumes, from which the second has been almost completed (Marticorena & Rodríguez �99�-�00�). It seems to be also the time for a renewal of Reiche’s Plant Geography. Not few things have changed in �00 years: plants have been renamed and reclassified; taxonomy and systematics have suffered deep changes; biology, geography, and biogeography have undergone paradigmatic vicissitudes.

In such a composite discipline like biogeography, today any intend to integrate the different views that shape it, must confront not only the differences inherent to the diverse disciplines involved, but also the more general conflicts that affect today any scientific endeavour. For the one side there have been sincere intends to integrate phytogeography and zoogeography in one corpus of integrative and synthetical biogeography (e.g. Croizat �9�8, �9��). On the other side, the biogeographic arena is getting more and more fragmented due to a plethora of methods (Ebach et al. �00�), and the ultimate synthesis is getting more and more elusive (Lomolino & Heaney �004). Some speak about the crisis of biogeography (e.g. Riddle �00�), but this crisis is not restricted to biogeography and seems to be more general, as the crisis of reductionistic modern science

��

in its failure to account for the real world problems, as challenged by postmodern theoretical scientists.

To which extend biogeography assumes and reflects the conflicts, assumptions and challenges inherent to modern and postmodern science will be discussed upon the analysis of Chilean plant geography. Aspects of systematics, cladistics, evolutionary theory, palaeogeographic scenarios and their relationship to biogeography will be discussed in the thesis.

The �st chapter sets the theoretical basis for the endeavour of doing plant geography in the ��th century, in a constantly changing world (sensu Ebach & Tangney �007). The �nd chapter has to do with the discovery of the Chilean plant world for modern science, as well as the taxonomic and plant geographic classifications of the flora. Any biogeographic intend needs a solid classification basis, and therefore an update of the Chilean flora following a modern classification system has been made. The �rd chapter deals with the geographic relationships of the Chilean flora, further analyzed in chapter four. Chapter five deals with the possible origins of the Chilean flora as it can be learned from palaeogeography. Chapter six takes up again some of the problems and conflicts announced in the �st chapter, in relation to the Chilean flora. Chapter seven intends to summarize the problems expressed in the former chapter, and the final chapter exhibits some conclusions about the Chilean plant geography.

��

1.1 Inventing plant geography

“Biogeography, however, is just one part of our attempt to understand the Universe, and the classification of this attempt at understanding into multiple disciplines (biogeography, ecology, geology, cosmology, chemistry, etc.) is an artificial one designed for our convenience. Indeed the philosopher Midgley [�989, �00�] has strongly argued that we have impeded our understanding of the world by overemphasizing these unnatural divisions. Biogeography does not need a rigorous definition beyond a general statement that it is about the distribution of organisms in space and time” (Wilkinson �00�).

Plant geography, phytogeography, geobotany, chorology, Arealkunde. These concepts have been used for two centuries in distinct and similar ways since Alexander von Humboldt’s Essai sur la géographie des plantes (Humboldt 1805). The limits between these fields have never been clear and it seems to be a waste of time delimiting them strictly. Augustin Pyramus de Candolle (�8�0) called the subject just Géographie botanique and his son Alphonse de Candolle (�8��) added the adjective [Géographie botanique] raisonné. Later Schröter wrote about genetische Pflanzengeographie oder Epiontologie (Schröter �9��), while Meusel preferred Vergleichende Arealkunde or Geobotanik (Meusel �94�), and Walter added the adjective floristisch-historische [Geobotanik] or reduced it to Arealkunde (Walter �9�4, Walter & Straka �970). Schmithüsen proposed that geobotanists emphasize the plant and therefore should call their study plant geography = Pflanzengeographie; geographers should have their emphasize in the physiognomy of plant communities, i.e. the vegetation, and therefore should call it vegetation geography = Vegetationsgeographie (Schmithüsen �9�8). This proposal has not been followed by geographers like Richter (�997) in his Pflanzengeographie, in the search for an integrative concept in which patterns and processes are combined. Some botanists wrote about botanical geography (e.g. Thiselton-Dyer �909), others about geographic botany (e.g. Raup �94�)�. For the present thesis I took the original concept of A.P. de Candolle, simply traduced as plant geography. In this form I can aside honour Stanley A. Cain’s great piece Foundations of plant geography (Cain �944). The present work is tied to the tradition of German chorology/chorography and Arealkunde, this latter an already extinct discipline.

1.2 Chorology and Arealkunde

„The Germans have a convenient term fort the science of area, Arealkunde. The term chorology is already in international usage, but it is less definite and more inclusive than Arealkunde. Two possible translations of Arealkunde are areology (which has an entirely different usage in astronomy)� and spatiology; but perhaps the best term for the science of area is areography, that portion of geography which deals especially with area” (Cain �944, �47).

�4

1.2.1 Origin of chorology

The term chorology was introduced by Ernst Haeckel in the natural sciences (Friis �998), from the Greek word Chora (χώρα) = the residence, the distribution district.

„Unter Chorologie verstehen wir die gesammte Wissenschaft von der räumlichen Verbreitung der Organismen, von ihrer geographischen und topographischen Ausdehnung über die Erdoberfläche. […] Im weitesten Sinne gehört mithin die gesammte „Geographie und Topographie der Thiere und Pflanzen“ hierher, sowie die Statistik der Organismen, welche diese Verbreitungs-Verhältnisse mathematisch darstellt“ (Haeckel �8��, pp. �8�-�87).

Haeckel recognizes A. von Humboldt and F. Schouw as the founders of plant geography, but both did not used the term chorology. The term geographische Florenkunde was more in use by the botanists at the end of the �9th century (Drude �890), later replaced by genetische Pflanzengeographie (Irmscher �9��, �9�9). Suessenguth (�9�8) and Vester (�940), this latter in his extensive thesis about the area types of the angiosperm families, wrote about Arealgeographie. At that time O. Schwarz used again the term chorology, as Phytochorologie (Schwarz �9�8).

Chorology became one of the most important terms in German geography when geographers tried to move from a descriptive to an analytical science. This challenge at the beginning of the �0th Century touched not only geography but all sciences. Geography suffered the transformation from a chorographic into a chorologic science. Geographers involved in this task were Peschel, Marthe, Richthofen and above all Hettner (�9�7), although some accuse him, as having not yet abandoned a descriptive science (see Holt-Jensen �999). The substance of a chorologic science was the analysis of the relations between specific places or regions. This was the emphasis of regional geography, an approach that prevails till today in many geographical institutes. Only in the �0’s geography saw a real change, mainly due to the work of William Bunge (�9��). He favoured a geography based on spatial analysis, which stressed the geometrical arrangement and patterns of geographic phenomena. Then the term chorology lost its meaning in geography.

1.2.2 Vergleichende Arealkunde

At the end of the �9th century there was under the botanists an imperative need to map the growing botanical knowledge. This not only concerned the pure representation, as it was so good expressed by R. von Wettstein and his geographisch-morphologische Methode. The analysis of distribution of the taxa could be the key for the solution of phylogenetic and systematic problems (Wettstein 1898) (figure 1.1)�. Die Pflanzenareale was a series that intended to compile the distributional knowledge available at that time in a central collection (Hannig & Winkler 1926-1940) (figure 1.2). „Only the diagrammatical representation on area maps can offer the necessary descriptiveness of distribution conditions of extant and fossil plants and supply thus the indispensable basis for the floristic analysis, the plant-geographical arrangement and the history of the development of the floristic regions, as well as for certain phylogenetic problems of

��

theoretical systematics“ (Hannig & Winkler �9��-�940). The discipline was called Vergleichende Arealkunde = comparative Arealkunde, because „although many area maps will actually show interesting distribution conditions, others will only obtain its value in the relationship with other areas” (Hannig & Winkler 1926-1940). The series achieved only five volumes, but anyhow presented the maps for �4 families, ��8 genera and �4�8 species.

1.2.3 Bloom time and fall

The first one who explicit synonimized Arealkunde with the older term chorology was Hermann Meusel in his fundamental work Vergleichende Arealkunde (Meusel �94�). Afterwards came Walter’s Arealkunde (Walter �9�4, �. ed. by Walter & Straka �970). Both Meusel and Walter understood Arealkunde as floristisch-historische Geobotanik. The authors propose the discipline as an integrative one. They recognized as a basic topic the cartographic representation but by far not the only one. In addition, the object of study were the nature of the areas (e.g. size and form), the relationship with systematics, the development of the floras in the course of geologic history, and the floristic relationships (floristic elements, area types). Meusel and his co-workers of the

Figure �.� Distribution map of morphologic similar Gentiana species (section Endotricha), applying the geographic-morphologic method (von Wettstein �898)

��

arealkundliche school at Halle/Saale brought Arealkunde/Chorologie to its highest point, with the work Vergleichende Chorologie der Zentraleuropäischen Flora, published in three volumes (Meusel et al. �9��, �978; Meusel & Jäger �99�). From the �0’s to 70’s we can mention also the large mapping work of NW Europe from Swedish E. Hultén, and his Circumpolar plants (Hultén �9�0, �9�4, �97�). Overseas, the work of C. van Steenis and M. van Balgooy, Pacific Plant Areas, appeared in five volumes between 1963-1993. With a more regional relevance it is worth of mention the geobotany at Erlangen (e.g. Gauckler �9�0, Milbradt �97�, Welß & Lindacher �994).

The earlier maps were an auxiliary tool for solving more general problems about the relationships

Figure �.� Donat‘s distribution maps in Hannig & Winkler (�9��-�940)

�7

in and between floras, but slowly the initial sense went lost and more emphasis was given to the maps. First these maps represented areas or outlines, but after the Atlas of the British Flora from Perring & Walters (�9��), geobotanists are more concerned with dot/grid methods „The influence of this first national atlas following the dot/grid method led in Europe to a true renaissance of floristic mapping and consequently to an indeterminable number of atlases at the national, regional and local scale” (Haeupler 2005). Examples are the floristic atlas of Germany (Haeupler & Schönfelder �988), of Bavaria (Schönfelder & Bresinsky �990), or the Flora of the Regnitz area (Gatterer & Nezadal �00�). In the meantime chorology broke down and lost its chorological character, maintaining as principal goal only the cartographic representation, i.e. as a Chorographie in the sense of pre-Hettner geography. Its comparative nature, the question about the origin of floras, about the relations between floras, went thereby lost. There are today only few users of chorology in its original sense, like Huxley et al. (�998) as successors of F. White and its extensive work on Africa (White �97�, �99�); or Cope (�000) as the last representative of the chorological work with grasses at Kew. Already in the preface of his work Meusel (�94�) realized “…During the boom that affected plant-geographical research in the last decades, chorology stayed remarkably behind vegetation science (phytosociology). This may be partially connected with the fact that the methods of modern ecology are better suited for vegetation analysis than for floristic problems”. Afterwards the vegetation science developed much more strongly, by integration of concepts of the ecology (e.g Clements), the Zürich-Montpellier school (Braun-Blanquet) and phytosociology (Tüxen). This was also recognized by Rothmaler: “…Working directions like the ecology detached themselves as independent branches of science, so that the original (floristic) plant geography went adrift. This however stands in close connection with taxonomy; a complete separation of both spheres of activity is not possible” (Rothmaler �9��). Floristic historical plant geography has been therefore taken over by other disciplines, partially by palaeobotany, historical biogeography or systematic botany.

1.2.4 Arealkunde v/s areography

As cited at the beginning of this section, Cain (�944) suggests that Arealkunde could be translated as areography. Areography developed in fact as an important subdiscipline of biogeography and harbours a whole chapter in the newest edition of the mainstream textbook Biogeography (Lomolino et al. �00�). To areography belong the analysis of geographic ranges (e.g. Gaston �00�) and ecogeographic rules, like Bergmann’s rule, Allen’s rule, and perhaps the most famous Rapoport’s rule, about latitudinal richness gradients. The Argentine ecologist E. Rapoport can certainly be recognized as the re-finder of areography (Rapoport 1982), although he most surely borrowed the term (not the meaning) from Cain (�944). He knew the term chorology, however decided himself for areography, because “the term chorology has been used by some authors as a synonym for biogeography”. Rapoport explained the new discipline as metabiology, and transformed the qualities of areas into quantifiable characteristics to allow their analysis via statistical methods. Therefore the work is one of the foundation-stones of the rapidly growing discipline macroecology (Brown �99�, Blackburn & Gaston �00�). Indeed Rapoport’s rule is still

�8

constantly examined and discussed in ecological and biogeographical journals �0 years after its proposal. Rapoport’ s areography attaches great importance to form, size and development of the distribution ranges; however it seems that it has lost the connection to historical biogeography, palaeobiogeography and systematics.

1.2.5 Post mortem

Parallel to the rapidly arising areography, chorology and Arealkunde broke down, and are not even mentioned in English textbooks (e.g. Cox & Moore �00�, Lomolino et al. �00�). Arealkunde is still referred to in German textbooks, but always in relation to the old good days of Meusel and Walter (e.g. in Richter �997, Schroeder �998, Frey & Lösch �004). There is however no advancement of the discipline; Arealkunde could not adapt to the modern times (and did never get a proper translation into English), and virtually went extinct.

Contrary to Arealkunde, “chorology is still in use, in quite a variety of different ways, but many modern studies do appear to be within that remit” (Williams �007, p. �9). “Chorology is the study of the mechanisms of distribution related to taxon origins… and authors like de Queiroz [2005] essentially embraces what is chorology” (Williams 2007). This definition is only partially correct: chorology certainly has to do with mechanisms of distribution, as an intend to explain static patterns of distribution (which would be the study object of a chorography). But chorology do not necessary has to do only with the mechanisms (vicariance and/or dispersal) that embrace unrelated histories (as in de Queiroz 2005). Chorology has it strengthens in its comparative nature …“although many area maps will actually show interesting distribution conditions, others will only obtain its value in the relationship with other areas” (Hannig & Winkler �9��-�940). In this sense chorology is in the line of Parenti and Ebach’s comparative biogeography, and is closer attached to systematics (Rothmaler �9��, Stuessy et al. �00�, Parenti & Ebach in prep.). Chorology and Arealkunde have trespassed their essence to current more integrative approaches, like Croizat’s panbiogeography (Croizat �9�8). For the description and cartography of distributions, the most suited concept is still the old-fashion term Chorography.

�9

1.3 Naming the plant worldThe plant geographical task is intrinsically related to the systematic task, specially in its classificatory aim: taxonomy4. A brief summary of the history of taxonomy can be found in Stuessy (�00�) and Cullen & Walters (�00�), as resumed from earlier studies, e.g. Croizat (�94�), Lawrence (�9��), Davis & Heywood (1963), Griffiths (1974), Morton (1981), Walters (1986), and Stevens (1994). Integrating Stuessy’s and Cullen & Walters’ views we can identify seven phases in the history of plant taxonomy (table �.�.)

Table �.� Phases in the history of plant taxonomy From Stuessy (�00�) and Cullen & Walters (�00�).

Phase Period Issues

� �00 BC - �4�0 Pioneer Greek Theophrastus wrote several manuscripts dealing with plants, translated as Enquiry into Plants (Hort �9��, quoted by Stuessy �00�).

� c. �4�0-���0 Age of the Herbalists, practical knowledge against diseases

� ���0 - �7�� Classification per se, searching for revealing ‘God’s system of classification used at the Creation’, i.e. as artificial systems, (e.g. Caesalpino, Bauhin, Ray de Tourneffort, culminating in the system of Linnaeus’ Species Plantarum of �7��.

4 �770-�880 The search for natural systems (e.g. Jussie’s Genera Plantarum, de Candolles’ Prodromus).

� �880-�9�0 Phylogenetic systematics, mainly enforced by Darwin’s Origin of Species (�8�9). The hierarchical structure was no longer viewed as the work of the Creator but rather as a result of the Earthly process of organic evolution. Classifications were organized along phylogenetic principles or ideas as to what might be primitive, what derived, and which groups might have evolved from other (Stuessy �00�) (e.g. systems of Engler & Prantl, Hutchinson, Takhtajan, Cronquist).

� �9�0-�980 Phenetic studies avoiding phylogenetic speculation and returning to a ‘more natural’ approach, sometimes using numerical methods

7 �980 onwards Resurgence of phylogenetics mainly based on cladistics and molecular information.

Cullen & Walters (�00�) emphasize that from the pre-Linnean period to the present three aspects have been contending for predominance:

a) the analytic, i.e. accurate and clear methods of identification and reference;b) the synthetic, i.e. grouping of units on the base of similarities and differences in a hierarchical

system, in a system that allows inferences (predictions) about the characteristics and properties of the units;

c) the phylogenetic, which seeks to demonstrate evolutionary relationships.

“At different periods, different aspects have been in the ascendant … Most of the practical taxonomic results produced have been in the form of Floras, monographs and revisions, which –on the whole- are concerned only with the first two aspects above. Running

�0

alongside this whole process is the need for a stable nomenclatural system to facilitate communication. … Gilmour [1936/1989] puts this very elegantly: ‘It [the taxonomic process] is a tool by the aid of which the human mind can deal effectively with the almost infinite variety of the universe. It is not something inherent in the universe, but it is, as it were, a conceptual order imposed on it by man for his own purpose’. Thus an element of convenience is built in to the idea of these higher groups from the beginning”. (Cullen & Walters �00�, p. 47).

Also Stuessy permits himself a sceptical thought: “This need to classify may relate to our use of language and its logical structure, it may reflect our general insecurity about the life-experience and our desire to control it better or it could possibly reflect how life itself is really organized” (Stuessy �00�, p. �4). This question lies on the basics of systematics and have been largely debated, mostly by philosophers of science (e.g. Endersby �00�, Hull �00�). The early words of the German philosopher E. Cassirer are still appropriate to discuss the subject. Cassirer speaks about “the naïve view of the world”. According to his view “systematics consists of grouping things according to their similarities. Our concepts of groups or classes are supposed to arise by our distinguishing the similarities of things from their differences; as we ascend in abstracting thought to ever more inclusive classes, the similarities which we recognize become ever fewer, until I suppose they eventually disappear. From an ontological viewpoint this kind of thinking must be judged naïve, because it involves presuming that the similarities (common attributes) of things have a simple hierarchical distribution. But there is in nature no unique hierarchy of similarities, as can be seen from the following example taken from Popper [�9�9, p. 4��]” (Cassirer �9��, as quoted by Griffiths 1974, p. 88). Popper’s example is reproduced here as figure 1.3.

In the practice this dilemma has been tried to be solved by taxonomists in augmenting the characters and states� under evaluation, or in providing a priori a phylogenetic framework to organize the similarities. As shown in table �.�, phylogenetic systematics arose only after the general acceptance of Darwin’s Origin of Species, and the consequent changes in biological theory.

Woodger (1952) explained this change in biological theory: “In the Linnean system of classification

Figure 1.3 Similarities between simple geometrical fi-gures. Some figures are similar with respect to shading or its absence; others are similar with respect to shape; and others similar with respect to size (after Popper 1959, as quoted by Griffiths 1974, P. 88)

��

of animals and plants a species was a set or class, in fact it originally meant a smallest named class in the system. But a class or set is an abstract entitiy and thus has neither beginning nor end in time. We cannot, therefore, speak of an origin of species if we are conceiving species in the Linnean manner. The doctrine of evolution is not something that can be grafted, so to speak, onto the Linnean system of classification. The species of Darwin and the species of Linnaeus are not at all the same thing – the former are concrete entities with a beginning in time and the latter are abstract and timeless” (Woodger 1965, p. 19, quoted by Griffiths 1974). In this sense, Stuessy (�00�) recognizes three different schools of systematics that developed over the past 40 years: phenetics (end �9�0s into the �9�0s), cladistics (�970s-�990s) and phyletics (�000 onwards).

In the following the emphasis will be put on some aspects of phenetics and cladistics (figure �.4), being the latter the current dominating paradigm in systematics since the translation of Willi Hennig’s Grundzüge einer Theorie der phylogenetischen Systematik into English (Hennig �9�0, �9��).

1.3.1 Phenetics

According to Stuessy (2006) “Phenetics was the first attempt to place biological classification on a more explicit footing. To do so, it openly rejected evolutionary interpretations, at least in the process itself, being too complex and mired in circular reasoning (e.g Sokal & Sneath �9��)”. These latter authors already knew Hennig’s work, but they were not impressed by the various rules Hennig set out for inferring phylogeny, nor by Hennig’s methods of reciprocal illumination. They found them too liable to produce error, like reasoning in a circle, “the piling of hypotheses upon hypotheses … Their goal was to develop clear, logical, straight-line methods of classification. First a systematist must do A, then B, and only then C – no circles, no spirals, no back-tracking.” (Hull �00�, p. ��0). Phenetics treated characters and states as particles of information to be assessed mathematically for purposes of grouping and ranking. All characters and states were treated as equal (usually) and overall similarity was emphasized. It stressed the importance of the basic data matrix of all life – the matrix of characters and states that must be used in classification. It is no doubt that this approach was possible thanks to the advances in computation science, without which it would have been impossible to evaluate large amounts of data (Stuessy �00�).

Figure �.4. David Maddison‘s graphi-cal representation of the debate bet-ween cladistics and phenetics during the 80‘s. [http://david.bembidion.org/]

��

1.3.2 Cladistics

Cladistics viewed the quantitative aspects of phenetics favourably, but advocated the reinstatement of evolutionary dimensions into classification (e.g. Hennig 1966, Wiley 1981). This approach focused on determining branching patterns of evolution (representing phylogeny), which could be determined mathematically. Particular characters and states were selected that were believed to have maximum phylogenetic value. Cladistics then restores evolutionary interpretations back into classification, but it also maintained quantitative comparisons (Stuessy 2006).

The methods proposed by Hennig (�9�0, �9��) have been coupled with special computer programs during the last decades, and became the reigning paradigm in systematics. The methodology rests, according to Cullen & Walters (�00�), in three a priori axioms: a) monophyly: principle that any classificatory group should be monophyletic, in contrast to polyphyletic or paraphyletic (box �.�);b) the principle that assumes that the evolution of a group proceeds by the minimum number of possible steps;c) every evolutionary event is a bifurcation of a lineage into two. This implies that each group has one, and only one, other group to which it is most closely related: the sister- group.

polyphyletic (Greek for „of many races“) a group which its mem-bers have in common evolved separately in different places in the phylogenetic tree. Equivalent-ly, a polyphyletic taxon does not contain the most recent common ancestor of all its members.

paraphyletic (Greek para = near and phyle = race) a group that contains its most recent common ancestor, but does not contain all the descendants of that ancestor.

monophyletic: a group (Greek: „of one race“) that onsists of a common ancestor and all its descendants.

clade -- A monophyletic taxon; a group of organisms which includes the most recent common ancestor of all of its members and all of the descendants of that most recent common ancestor. From the Greek word „klados“, meaning branch or twig.

cladogenesis -- The development of a new clade; the splitting of a single lineage into two distinct lineages; speciation.

cladogram -- A diagram, resulting from a cladistic analysis, which depicts a hypothetical branching sequence of lineages leading to the taxa under consideration. The points of branching within a cladogram are called nodes. All taxa occur at the endpoints of the cladogram.

sources: UCMP glossary: http://www.ucmp.berkeley.edu/glossary/gloss�phylo.htmlUnderstanding evolution: http://evolution.berkeley.edu/evolibrary/article/phylogenetics_0�

Box �.� Basics of cladistics

��

Cladistics arose in the search for a system that rests upon scientific objectivity and less on the subjective view of the taxonomist “to create a rigorous, objective methodology for reconstructing phylogenies upon which classification can be based” (Endersby 2001).

“In sum, the path leading from a desire among systematists to make classification more scientific to the present state of affairs has been long and tortuous. Species began as a significant level of organization in the taxonomic hierarchy. Increasingly, it is thought of as just one level out of many. Taxonomists began by thinking that the Linnaean hierarchy is the ideal way of representing phylogenetic trees, but as they attempted to make the relationship between hierarchical classifications and phylogeny clearer and more objective, they were forced to realize that representing even the most straightforward phylogenetic relationship (e.g., sister groups) is very difficult, if not impossible. One solution to this problem is to abandon the Linnaean hierarchy“ (Hull �00�, p. ���).

The abandonment of the Linnean hierarchy have turned in fact in one of the most controversial issues in the history of systematics, due to the arise of a new unranked system based solely on terminal taxa, the Phylocode (e.g. de Queiroz & Cantino 2001, de Queiroz 2006). But on the other hand, authors like Grant (�998, �00�) still maintain that cladistic and taxonomic systems are different schools, equally effective, each in its own way.

1.3.3 Is a rose really just a rose?

“A rose is still a rose, but everything else in botany is turned on its head” �

The rise of cladistics came to a peak in �998, when the new angiosperm classification (APG �998) appeared on the newspaper, on TV and in several popular magazines, achieving popular attention for several weeks. The press talked about a botanical revolution, a breakthrough, about genetic revelations. Appearing in the media is not normal for scientific botany, and specially for taxonomy to get front page newspaper headlines is unprecedented. “Was this an hazardous minute of fame?, is there a real revolution?, or was this coverage promoted by the proper scientific group? The popular interest was partly because of the claimed accuracy of the new system, and because this accuracy was based on DNA sequencing via computers – two sexy topics for the popular press” (Endersby �00�).

Endersby (2001) first noted the unprecedented system of publication, that can be seen as a modern manifesto: the publication as a group, not as an individual (APG �998). This effort of collaboration, between 29 botanists with their own interests and personal conflicting views, has but many readings:

a) the collective work allowed more comprehensive results that its members would have achieved individually;

b) publication under the group name is also a rhetorical device that draws attention to its objectivity;

�4

c) creating a group identity excludes practitioners of alternative forms of taxonomy.In this way, emphasizing the novelty of the work and its claim to be objective, the group gained a better position for its approach and for an ongoing funding. But to what extent the result represents a real botanical revolution is a matter of discussion, and interviews done by Endersby (�00�) for his paper put a question on the supposed objectivity:

“there is, however, still an element of subjectivity, or tradition, involved, viz. with respect to which groups should be named” (Bremer �999, quoted by Endersby �00�).

“…the orders are almost purely and simply conveniences, and in a number of cases there is no real reason whatsoever to prefer one delimitation of a group over another... (Stevens �999, quoted by Endersby �00�).

“…we didn’t want it to be cited as ‘somebody et al’, because it did represent the work of so many different people in different places… we wished to stress the fact that this wasn’t a system of classification done by an expert - that is somebody like a Cronquist, or a Takhtajan, or Thorne ... We wanted to explicitly say that botanists had gone beyond the need for authorities in this way... (Chase �999, quoted by Endersby �00�).

But APG’s effort just changed the authority of somebody toward the authority of the group, the consensus, valid only for insiders and not for outsiders of alternative taxonomy. However, where the supposedly objectivity stayed is a question that still remains (see further discussion in section �.�.�).

1.4 From the map to the tree: cladistics and biogeography

„Ich setzte also im Erwachsenenalter die Spielereinen meiner Kindheit mit Karten fort: ich verband Städte gleicher Größe durch gerade Linien, einmal, um festzustellen, ob im Eisenbahn- oder Straßennetz gewisse Regeln erkennbar seien, ob es regelhafte Verkehrsnetze gäbe, zum anderen, um die Abstände zwischen gleich großen Städten zu messen. Dabei füllten sich meine Karten mit Dreiecken, oft gleichseitigen Dreiecken – die Abstände gleich großer Städte untereinander waren also annährend gleich --, die sich zu Sechsecken zusammenschlossen. Ich stellte weiter fest, daß in Süddeutschland die kleinen Landstädte sehr oft sehr genau einen Abstand von �� km voneinander haben. Mein Ziel war abgesteckt: Gesetze zu finden, nach denen Anzahl, Größe und Verteilung der Städte bestimmt sind...“ (Christaller �9�8, p. 9�) 7

The citation is not just anecdotic but exemplifies the importance of the cartographic work for the comprehension of spatial phenomena, even as such an intuitive play done by a child. In this case the child grew and turned to be one of the most recognized geographer of the �0th century, as creator of the central places theory.

Distribution maps were and are still central in the development of biogeography since the first attempts in the �9th century by Lamarck & de Candolle (Ebach & Goujet �00�) and F. Schouw

��

(Mennema �98�) (see sections �.� and �.�). But modern biogeography is changing the maps for the branching trees, and has more to do with the topology of cladograms and area cladograms than with distribution maps. This trend can be followed back to the work of Gareth Nelson and Norman Platnick, by an approach they called vicariance biogeography, latter named cladistic biogeography (Humphries & Parenti �98�).

Box �.� Basics of cladistic biogeography

In cladistic or vicariance biogeography distributions of monophyletic groups of taxa over areas are explained by the reconstruction of area cladograms. These area cladograms are hypotheses of historical relationships between areas and are derived from phylogenetic and distributional information of the monophyletic groups concerned. A first-order explanantion for correspondence between phylogenetic relationships of taxa and historical relationships among areas is vicari-ance.

When formation of barriers or splitting up of areas triggered speciation (i.e. vicariance), all spe-cies are endemic to their own area. In such simple cases, derivation of an area cladogram is trivial. Replacement of taxa in the taxon-cladogram by their areas of distribution results in area clado-grams with an own and unique terminal node for each area. However, real data are mostly the result of other processes such as extinction of species in part of their range or dispersal of species over the formed barriers.

As a result of these processes, taxa of a monophyletic groups can become widespread or sympatric in their distribution. In order to obtain area cladograms with an own and unique terminal node for each area, additional steps are necessary. In vicariance biogeography these additional steps are implemented in three assumptions about the cause of widespread and sympatric taxa.

Source: Marco G.P. Van Veller home page: http://home.hccnet.nl/m.van.veller/ See also Hum-

AA A

A

B B

B

C

C C

CD

D D

��

1.4.1 Vicariance biogeography v/s dispersalism

Gareth Nelson and Norman Platnick developed this approach based on the work of León Croizat (Nelson & Platnick �98�), but vicariance as an explanation of disjunct distributions is already found in the early works of J.D. Hooker (Turrill �9��), and in Ratzel’s biogeography (Ratzel 1901). But more specifically, as Nelson and Platnick recognized, their views had grown out of the work of Hennig, Croizat and Karl R. Popper. “Although they [Nelson & Platnik] admitted that Croizat found Hennig’s principles of phylogenetic systematics antithetical to his own principles of panbiogeography, and that neither Hennig nor Croizat had ever mentioned Popper (and vice versa), nevertheless Nelson and Platnick considered their own work as synthesizing and extending the views of these three authorities” (Hull �00�, p. ��4). They extendedly discussed the science of branching diagrams - cladistics in its most general sense (box �.�). “It was primarily through the championing of Hennig’s principles of biogeography, first by Lars Brundin of the Swedish Museum of Natural History and then by a young American ichthyologist, Gareth Nelson, that Hennig finally gained some recognition among English-speaking biologists” (Hull 2001, p. ���).

Nelson and Platnick’s approach was further extended by other authors that found in this approach a real changing paradigm from the Darwinian dispersalist biogeography. Dispersalism long dominated the biogeographic scene since the publication of The Origin of Species, and was further developed by the New York School of biogeography, in the figures of W.D. Matthew (1871–1930), K.P. Schmidt (�890–�9�7), G.G. Simpson (�90�–�984), P.J. Darlington, Jr (�904–�98�) and G.S. Myers (�90�–�98�), (reviewed by Nelson & Ladiges �00�).

Dispersalist biogeography is based on the assumption that taxa originated in relatively small areas (centres of origin) and therefore tries to reconstruct the routes that organisms covered to colonize known past or present ranges. A good example of this view is Simpson’s Splendid Isolation: The Curious History of South American Mammals (Simpson �980). In fact, Croizat called dispersalism “the science of the curious, the mysterious, the improbable” (Croizat �9�8).

Many authors saw the problems in the search for the centre of origin and migration routes, like botanist Stanley Cain (�90�-�99�), who analized Adams’ �� criteria for the analysis of the centres of origin (table 1.2), and asserted that “… [centres of origin] have beeen largely accepted without question, despite the lack of substantiating data in some cases, and have been variously and somewhat loosely employed” (Cain �944, p. �8�).

After almost ��0 years of dispersalism, vicariance biogeography was taken by biogeographers as the new paradigm, and most of the literature in the 80’s and 90’s was engaged with the analysis of area cladograms.

But dispersalism was only taken a rest, and returned like the Fenix, with the new methods of molecular dating, as well expressed in the apologia of de Queiroz (2005), and recent papers of

�7

emphatic advocates (e.g. Renner �00�, McGlone �00�) (see the discussion in chapter �). An interesting point expressed by Grehan (�007) regarding current biogeographic studies is in direct relationship to Christaller’s quotation at the beginning of this section: “To discern the geographic context of evolution, Darwinian biogeographers look to historical theories of ecology, systematics, molecular clocks, and geology – anything but distributions themselves – as the empirical data of biogeography… One only has to see how often distribution maps are left out of the picture, even for papers focusing on biogeographic theory and method” (Grehan �007, pp. 8�-84).

Table �.� Adams’ �� criteria for centres of origin, as revised by Cain (�944, pp. �8�-���).

�. Location of greatest differentiation of a type �. Location of dominance or greatest abundance of individuals�. Location of synthetic or closely related forms 4. Location of maximum size of individuals �. Location of greatest productiveness �. Continuity and convergence of lines of dispersal7. Location of least dependence upon a restricted habitat 8. Continuity and directness of individual variations radiating from the centre along highways of dispersal9. Direction indicated by geographical affinities �0. Direction indicated by the annual migration routes of birds ��. Direction indicated by seasonal appearance ��. Increase in the number of dominant genes toward the centres of origin

��. Centre indicated by the concentricity of progressive equiformal areas

1.5 The fragmented map of modern biogeographyWilli Hennig, the great theorist behind the current cladistics paradigm, explained the criterion of veracity of his phylogenetic approach, with the reconstruction of a fragmented map:

“Suppose a geographer has obtained fragments of a topographic map of an unknown land. He will make every effort to reconstruct the map from the fragments. How can he succeed if the original map is unknown to him? He was not present when the map was torn up. The geographer must try to assign each fragment to its original place in the totally of all the recovered fragments. He will proceed by trying to find, for a portion of a river present on one fragment of the map, the adjoining piece of the same river on another fragment. If he directs his attention to a single geographic element in his map fragments, such as rivers, he is likely to go wrong. Thus the three sections of a river, a, a’, and a’’ (figure 1.5 a) could seem to be upper, middle, and lower parts of the same river. His error becomes obvious if he considers other elements (“characters”) of his map fragments. They remain isolated; pieces of roads and railroad lines do nor join up (figure 1.5 a). But if all geographic elements are satisfactorily fitted together (figure 1.5 b) the geographer will be convinced that the fragments have been assembled correctly, even though he did not know the original condition of the map” (Hennig �9��, p. ��0-���) 8.

�8

In section 1.3 cladistics was briefly introduced, therefore Hennig’s metaphor serves here more as an excuse to discuss the conflicting view in the current fragmented biogeographical scene. As stated by Riddle (�00� and references therein), modern biogeography seems to suffer a protracted identity crisis, since there is an evident lack of fully integrative approaches to determine “the roles of earth history and ecology in the geography of diversification” (Riddle 2005). The plethora of methods that are in common use (see a revision by Crisci et al. �00� and Morrone �00�) leave no opportunity for this integration. Nelson & Ladiges (�00�) characterized current biogeography as a “mess of methods”.

Critics come and go between the exponents of different approaches: Ebach & Humphries (�00�) and Ebach et al. (�00�) expose their arguments for a deconstruction of phlylogenetic biogeography at historical, theoretical and methodological levels. “The new theoretical framework within historical biogeography claimed by Van Veller et al. [�00�] is no more than a repeat of the pre-tectonic ideology that looks for centres of origin and direct lineages” (Ebach et al. �00�). Van Veller et al. argue “because ontological views are not subject to direct empirical testing, cladistic and phylogenetic biogeography may both be considered valid research programmes, each of which has its own particular focus of attention” (Van Veller et al. �00�). Furthermore, Donoghue & Moore (�00�) propose that cladistic biogeography has failed to become a truly productive research programm because its scope has been fatally over-simplified by being restricted to assessing the topological congruence of phylogenetic trees as depicted by general area cladograms. But Parenti & Ebach (in prep.) reject Donoghue & Moore’s integrative biogeography program because in their view it has been used to reinforce equivocal divisions, such as that between phylogenetically and ecologically focused methods.

“In our view, biogeography is one field with one aim – to explain the distribution of taxa today as elements of form, time and space regardless of whether the data is morphological or molecular and whether taxa are represented as species, sub-species or populations…The dismissal of Earth process from historical and ecological biogeography may have a sociological, political and perhaps an emotional raison d’etre. Whatever the reason, the rationale is unclear and alienating to non-biological disciplines. There is no reason to doubt the role of geological process, namely tectonics as the primary cause for geographical and environmental composition (soils, water)

Figure �.� Hennig‘s fragmented map as test of the criterion of veracity (Hennig �9��)

a b

�9

changes. Intrinsic changes cannot occur without an external stimulus. Evolution is not a constant battle between the eaters and those to be eaten, but rather changes in organisms that are living and surviving on a dynamic Earth” (Ebach & Humphries �00�).

After the period of discovery that marked the origin of modern nature sciences (with A. von Humboldt) “historical and ecological biogeography and their many subdivisions evolved, diverged, and eventually flourished (or languished) as increasingly more distinct disciplines” (Lomolino & Heaney 2004). This diversification and growth of distinctive scientific disciplines established a presumed need to specialize that resulted in more and more splitering.

Lomolino & Heaney (�004) interpret such a splitering rather optimistic: “the greatest strides we can make in unlocking the mysteries and complexities of nature in this fundamentally interdisciplinary science are those from new synthesis and bold collaborations among scientists across the many descendant disciplines, long divergent but now reticulating within a strong spatial context – the new biogeography.” They continue: “From the vicariance hypotheses of Josef Dalton Hooker in the mid �800s to the most recent methods of analyzing reticulating phylogenies and phylogeographies, geographic variation over time and space is the key. How life forms vary across kingdoms, from the unicellular organisms to the greatest beasts, and from ancient to current (and to the future), how all this varies across geographic gradients -this is the realm of the new biogeography” (Lomolino & Heaney �004).

Lomolino & Heaney (�004) continue with their optimism: “the revitalization will continue in earnest, largely through the efforts of broad-thinking scientists who no longer shy away from but embrace the complexity of nature, and who foster collaborations and conceptual reticulations in modern biogeography”.

1.6 Postmodern biogeography: de(re)constructing the map

„Vivimos tiempos en que constantemente el postmodernismo confronta el discurso hegemónico de la ciencia para denunciar sus excesos y exponer sus límites“ (Morrone �00�, p. 87) 9

Lomolino & Heaney (�004) recognized that after the specialization and splitering of biogeography “…the grand view, the ultimate synthesis across space and time, became murky and more elusive”. Lomolino & Heaney (�004) expect that the great synthesis will come from “encourage creative development and applications of the comparative approach, deconstructing and reassembling more comprehensive explanations for the diversity and distribution of biotas”. But this is in conflict with the specialization, and some authors claim that biogeography is currently “relegated to the interesting pay-off at the end of systematic papers” (Upchurch �00�).

�0

Furthermore, in proposing to deconstruct current approaches and promoting creative applications, Lomolino & Heaney are going beyond from modern reductionistic science and enter the complex and more unpredictable region of postmodern science. In the search of the grand view, the ultimate synthesis across space and time we come unavoidably to León Croizat’s analytical and synthetical biogeography: Panbiogeography (Croizat �9�0). In fact, Croizat’s catch-phrase the earth and the biota evolve together has become one of the most popular slogans in the history of biogeography. Croizat expressed this explicitly in dozens of articles and several books like Space, time, form, the biological synthesis (Coizat �9��) (see Heads & Craw �984 for a complete bibliography). Croizat’s synthetical view is what biogeographers have been searching since the beginning of the science as a modern task. Croizat, while opposing the dominant dispersalism of his time, is one of the major responsible of the revitalization, quite possible a scientific revolution, that biogeography is today experiencing (Lomolino & Heaney �004). The deconstruction suggested by Lomolino & Heaney (�004) is in tune with the revision that is today occurring in many sciences, specially the social sciences. The critical view of postmodernism left nothing untouched and our world view is changing more rapidly as many like to recognize it.

1.6.1 Objectivity and postmodern science

The philosopher Frederick Ferré wrote a challenging essay for the mainstream book Principles of Conservation Biology (Meffe & Carroll �997) about the limitations of reductionist modern science in responding to the real world problems. Only postmodern science would “nurture the human hunger for quality: for beauty, balance, creative advance” (Ferré �997). The reaction of Cotterill (�999) for such a naïve view was drastic: “postmodernism has little, if anything, to offer these nor any other science – especially where conservation biology is tasked with informing and changing policy… this uncritical sanction in a mainstream textbook on conservation biology kindles suspicions that a deeper problem afflicts the discipline’s scientific integrity” (Cotterill �999). Ferré’s essay got the sanction, and disappeared from the �d edition of the book (Groom et al. �00�). Cotterill and Attwell further developed his critique in a large article about Postmodernism and African conservation science, defending the traditional protectionist conservation in Africa, against community-based approaches. They conclude that “postmodernist thinking has had a significant negative impact on conservation science in Africa, largely by marginalising the central issue of human population pressure” (Attwell & Cotterill �000). Although this is not the forum for a philosophical rebuttal, reducing the problem of conservation to a single factor like population pressure is the typical reductionist view of modern science.

Attwell & Cotterill (�000) are very conscientious of the concerns of postmodernism. “In particular, it is the reductionist approach of science that finds such disfavour in postmodernist intellectual circles. The postmodernist view questions the objectivity of observation and ultimately the truth of scientific knowledge. Postmodernists may give equal credibility to explanations of the world, no matter whether the knowledge base has been arrived at via the scientific method or whether it has arisen through folklore or anecdote. The notion that science has no more claim to truth than, say, tribal mythology, is an extreme form of what is termed cultural relativism. It has been aptly

��

described as fashionable salon philosophy by Dawkins [�99�], who questions the entire credibility of the postmodernist programme [Dawkins �998]. In its extreme manifestations, postmodernism is the antithesis of science [Wilson �998]” (Attwell & Cotterill �000, p. ��0).

Remarkably, Attwell & Cotterill (�000) recognise that “this movement, by which knowledge can be used irrespectively of its scientific validity, is encapsulated in what is now called New Age thinking. Conversely, the traditional approach to the philosophy of science, to which we subscribe, is to view scientific knowledge as objective“ (Attwell & Cotterill �000, p. ��0).

1.6.2 Postmodernism as the villain?

From the same tribune that holds Attwell & Cotterill’s (�000) claims, the journal Biodiversity and Conservation, we can learn: “One strand in postmodern thought is concerned to re-appraise the status of scientific knowledge. One element in this re-appraisal is to challenge the alleged value-free status of scientific knowledge. The claim to be value-free involves two distinct theses. First, it is claimed that the realm which the natural sciences investigate is a factual realm. If values attach to nature, it is not the task of science to explore them: science investigates the factual character of the world, aims to discover facts about it. The second claim or ideal of value freedom involves, not the subject matter to be investigated, but the scientific investigator. The scientist does not import any values into the enquiry. He or she is a purely rational, value-neutral, investigator. Any attachments to, or interpretations or evaluations of, the subject matter which the scientist as an individual might hold are set aside when the scientist is engaged in his or her professional investigations” (Howarth �99�, p. 787).

Howarth (�99�) concluded “This may not be a criticism of science. It may be more a criticism of those other institutions in our culture which listen only to the voice of science and relegate questions about significance and value, and their answers, to the realm of the merely subjective preference and opinion. Merleau-Ponty [�9��] invokes us to listen to another voice: The whole universe of science is built upon the world as directly experienced, and if we want to subject science itself to rigorous scrutiny and arrive at a precise assessment of its meaning and scope, we must begin by reawakening the basic experience of the world of which science is the second-order expression. Science has not and never will have, by its nature, the same significance qua form of being as the world which we perceive, for the simple reason that it is a rationale or explanation of that world” (Merleau-Ponty �9��, as quoted by Howarth �99�, p. 797).

1.6.3 Deconstructing biogeography

Postmodernism has long permeated the social sciences, but the biological/physical sciences are much more immune to these concerns�0. Chorley expressed it with an excellent aphorism applied to the theory of geomorphology:

��

“Whenever anyone mentions theory to a geomorphologist, he instinctively reaches for his soil auger” (Chorley �978)

But even in such hard disciplines like geomorphology, the philosophical analysis to strengthen its intellectual foundation is on its course (e.g. Chorley �978, Osterkamp & Hupp �99�, Rhoads & Thorn �99�).

Perhaps the first explicit proposal for a deconstruction of biogeography, is the one of M. Heads on a guest editorial of the Journal of Biogeography, with the title ‘Mesozoic tectonics and the deconstruction of biogeography’ (Heads 1990). Heads briefly proposed an alternative view for the evolution of the Australasian biota, arguing that “approaches to biogeography have been based all to often on consideration of particular lineages, emphasizing purely theoretical ancestor-descendant relationships and have maintained a blind spot towards the general effect of phases of modernization on a landscape and its biota”. Heads did not stop there and took the challenge of join “recent developments in biogeographic theory integrate the findings of Croizat, Derrida and a Continental philosophical tradition often scored by the English-speaking world” (Craw & Sermonti �988, quoted by Heads �990). Later in the decade he published with R. Craw and J. Grehan the book Panbiogeography, that resumes most of the early ideas from Croizat coupled with new methods and opportunities of development (Craw et al. �999). More recently Heads (�00�a) wrote an extended revision on the history of panbiogeography.

Closing this chapter, two examples of recent evaluation of some long standing theories in ecological biogeography are worth of mention. The long standing dynamic Equilibrium Theory of Island Biogeography from MacArthur & Wilson (�9�7) has been recently questioned: “Functional areography provides convincing arguments for a postmodern deconstruction of major principles of the dynamic Equilibrium Theory of Island Biogeography” (Walter �004). Brown & Lomolino (�000) also found that “a dynamic equilibrium between contemporary rates of immigration and extinction, is clearly contradicted by phylogenetic and fossil evidence of a long, pervasive legacy of history on the diversity and composition of most insular lineages”. On another front, Marquet et al. (�004) proposed a novel approach in the study of species richness. The authors call for a “deconstruction of biodiversity patterns”, under the basic assumption that assessing diversity as richness does not adequately characterize the way in which species differ from each other, and which cause them to respond in different ways to changes in the environment. They claim that “deconstruction should be consciously performed as a methodological strategy”.

Under this critical scenario for the further development of biogeography, any attempt to do a Plant Geography needs nowadays to treat carefully every evidence. The task is not easy since many complementary but intrinsically different disciplines come to play: systematics, cladistics, geology, palaeogeography, every one with its own paradigms, assumptions, advantages and limitations.

��

“For history is like a nymph glimpsed bathing between leaves: the more you shift perspective, the more is revealed. If you want to see her whole you have to dodge and slip between many different viewpoints. This technique, which ought to commend itself to historians, might also restore to subjectivists a prospect of objectivity. Even the most dedicated subjectivist should be able to imagine what it would like: objectivity would be the result of compiling or combining all possible subjective points of view. Every time we take notice of each other, therefore, we get a little closer to truth. Those who refuse to acknowledge anyone’s existence save their own must approach truth by imagining a variety of perspectives. To see things from no point of view is not even theoretically possible. If we try to see from every point of view, we shall never attain our goal, but it is at least meaningful to speak of seeing from every point of view, whereas it is literal nonsense to speak of seeing from none.” (Fernández-Armesto �997, Truth, p ��8).

Following Fernández-Armesto’s idea, along the thesis the plant geography of Chile will be analized from different point of views: from the traditional views to the alternative ones. The effort has to be seen still as a work in progress, since:

“...Toda división fitogeográfica deberá, entre tanto, considerarse como provisional o aproximada. Podrán expresarse opiniones, pero discutir divisiones, límites o nomenclatura, o tratar de imponer ideas en forma dogmática, será solo gastar tiempo y papel sin mayor provecho para la ciencia.” (Cabrera �9��, as quoted by Ribichich �00�)��

�4

��

� Chile, a Remote Corner on Earth

��

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

�7

2 Chile, a Remote Corner on Earth

“Durante la Conquista, cuando los españoles recorrían América buscando esos metales [plata y oro] y llevándose todo lo que encontraban al paso, Chile se consideraba el culo del mundo, porque comparado con las riquezas del resto del continente tenía muy poco que ofrecer” (Isabel Allende, Hija de la Fortuna, �999)

The knowledge of the Chilean flora (to modern western science) has been build step by step since the first Spanish conquerors put their footprint on this land, but it is still far from complete or accurate. Many white areas on the floristic map remain to be found (depending of course on the scale of the map). Unknown territories… unknown plants?

2.1 Romancing the South: the discovery of a virgin worldThe discovery of the Chilean plant world can be tracked back to the circum-navigation of Hernando de Magallanes and Sebastián Elcano. A detailed relate of early collectors, botanists, and naturalists that contributed to the knowledge of the Chilean flora has been done by Reiche (�907) and Marticorena (�99�) (there is also a complete bibliography in Marticorena �99�, and an online summary by Muñoz-Schick & Moreira-Muñoz �00�). Some highlights in the discovery of the Chilean plant world have been briefly resumed in box 2.1.

Box �.� The discovery of the Chilean plant world

The discovery of the Strait of Magellan by Hernando de Magal-lanes and his crew (�� october ���0) set the start point for the European exploration of South America. Magallanes gave the name ‘Cabo de Las Vírgenes’ to the eastern Cape that allowed them to get into the Strait. During the first days of exploration, the observations are about a native wood to make fire which smoke smell well. They refer undoubtedly to the wood of ‘ca-nelo’, Drimys winteri. The sailors early noted the properties of canelo’s bark and herbs like Cardamine and Apium against scur-vy. The first notes from sailors refer of course the medicinal, nutritional and wooden properties of the new plants found.

Hipólito Ruiz and José Pavón did the first specific botanical excursion to Chile. With the purpose of extending the collections of the Real Botanical Garden of Madrid, king Carlos III of Spain sent several naturalist expeditions to different countries; Peru and Chile corresponded to Ruiz and Pavón. They made collections in some points between Talcahuano and Santiago between �78� and �78�. The work they published later in � volumes has more than

�00 illustrations. Here as example‚ the ‚copihue‘, Lapageria rosea.

�8

The English naturalist Joseph Dalton Hooker (�8�7-�9��) ac-companied the expedition to the South Pole of John Clark Ross (1839 - 1843). He was the first who collected intensively in the southern lands (Kerguelen, Tasmania, New Zealand, Falkland, Tierra del Fuego) and who noticed the close floristic relation-ships between these territories. His Botany of the Antarctic Vo-yage was published in several volumes between �844-�8�0.

Box �.� The discovery of the Chilean plant world (continuation)

In �8�7 the German naturalist Eduard Poep-pig arrived in Chile. He explored the country for two years and it was the first time that a foreign naturalist stayed in Chile for a long period. He also visited other countries of South America, and the work of � volumes and �00 illustrations (written with S. Endli-cher), was accompanied for the first time by coloured drawings. The night from 9. to �0. January 1827 Poeppig crossed for the first time the equator aboard the ship ‘Gulnare’.

“Leicht war die Stunde, in der wir aus der nördlichen Halbkugel schie-den, einer der feierlichten und bedeutsamsten eines langen Reiselebens … Fast will es dem Reisenden dünken als trete er mir dem Eintritte in eine neue Welt auch in ein neues Leben…“. (after Morawetz & Rösser

�998). The illustration corresponds to Puya alpestris.

The Jesuit Juan Ignacio Molina (1737-1829) is considered the first Chilean naturalist. After the expulsion of Chile of the Jesuit Compa-ny (�7�8), Molina published in Italy his Compendio della storia ge-ografica, naturale, e civile del Regno del Chile (Molina �77�). This and coming works (Molina �78�, �8�0) were for a long time the main source of knowledge on natural sciences of Chile. The illust-ration represents araucaria, the Chilean palm (Jubaea) and ‚culén‘ (Otholobium glandulosum).

�9

After its Independence from the Spa-nish dominion in �8�0, the young Re-public of Chile had to better know its natural resources. The French profes-sor of natural sciences, Claude Gay (�800-�87�), already living in Chile, was contracted by the government to do the first intensive scientific explo-ration of the territory. He intensively traveled between Copiapó and Chiloé (�8�0 - �84�) (Muñoz Pizarro �944).

The great Charles Darwin (�809-�88�), same as J.D. Hooker, was very young when he traveled around the globe aboard the Beagle between �8��-�8��. His impressions abouth the southern biota and geology were fundamental for the later development of the theory of evolution. He traveled in Chile between �8�4-�8�� and collected almost �.�00 plant specimens, that were later studied by J.D. Hooker. After climbing the Cerro La Campana, he wrote on his diary: „We spent the day on the summit, and I never enjoyed one more thoroughly. Chile, boun-ded by the Andes and the Pacific, was seen as in a map. The pleasure from the scenery, in itself beautiful, was heightened by the many reflections which arose

from the mere view of the grand range, with its lesser parallel ones, and of the broad valley of Quillota directly intersecting the latter. Who can avoid admiring the wonderful force which has upheaved these mountains, and even more so the countless ages which it must have required, to have broken through, removed, and levelled whole masses of them?” (Darwin �8�9).

Afterwards he published his masterpiece Historia Física y Política de Chile, which consists of �8 volumes (8 of botany) and � illustra-tion atlases (one cultural and one physical) (Gay �84�-�8�4). This was a work without precedents in America to that date. The mis-sion that the Government of Chile had trusted him also included the formation of a Cabinet of Natural Sciences, that was the base of the Museo Nacional de Historia Natural. Before him, the flora of Chile was known as compound of around �00 species and in his work �7�7 new species are described. Illustration of Desfontainia spinosa (source: www.memoriachilena.cl)

Box �.� The discovery of the Chilean plant world (continuation)

Rudolf Amandus Philippi (�808-�904) had to leave Germany due to political problems and arrived in Chile in �8��. He is recognized as the main naturalist in the history of Chilean science (Castro et al. �00�), having done studies in botany, zoology, geology, paleotology, etnology, etc. He gave a big impulse to the Museo Nacional de Historia Natural.

R.A. Philippi

F. Philippi

R.A. Philippi got soon the collaboration of his son Federico Philippi (�8�8-�9�0), exploring vast regi-ons in Chile and the Norte Grande, territories just annexed to Chile after the Pacific war. There were a great amount of collaborators who provided ma-terial to the Philippi. With the impulse and the bo-tanical explorations of father and son, the amount of �7�7 species described to that date rises to more that 7497 species. Of the �7�0 species described by them, 90% of the collections are conserved at the National Museum as ‘type exemplars’. This is an exception under Latin American countries, where most of the early collections and specially the type specimens ended in Europe or the U.S.A.

40

Francisco Fuentes (�879-�9�4) was incharged of the National Herbarium in �9��, as Karl Reiche leaved the country and accepted an appointment in Mexico. Fuentes made the first study on plants of the Easter Island (Fuentes �9��) and on Monocots that had not been treated in Reiche‘s Flora.

Gualterio Looser (�898-�98�) showed a wide interest in the natural sciences, having been appointed as chief of anthropology at the National Museum. Since a journey to Juan Fernández in �9�� he demonstrated more interest in botany, publishing till �97� abouth �40 works mostly dedicated to the ferns. He became the national authority in this theme. He also made big efforts in the translation

into spanish of main texts from Reiche (plant geography) and Skottsberg.

Box �.� The discovery of the Chilean plant world (continuation)

Karl Reiche (�8�0-�9�9), a German professor of natural sciences, was appointed as encharged of botany in �90� by Federico Philippi, when this last took the direction of the Museo Nacional de Historia Natural. Reiche published the second Flora of Chile (after Gay) in � volumes (Reiche 1896-1911). He also wrote the first plant geography of Chile (Gründzuge der Pflanzenverbreitung in Chile, �907, translated to Spa-nish by Gualterio Looser as Geografía Botánica de Chile).

Federico Johow (�8�9-�9��) arrived in Chile �889 to teach natural sci-ences at the Instituto Pedagógico, Instituto Nacional and Universidad de Chile. He is the author of the milestone Estudios sobre la flora de las Islas de Juan Fernández, still an obligate reference for the islands (Johow �89�). He should have been responsably for the new Flora de Chile with Karl Reiche, but due to problems with the National Museum he was removed

from this effort.

Ivan Murray Johnston (�898-�9�0), US American prominent botanist, stu-died the coastal desert flora and vegetation. His papers „The coastal flora of the departments of Chañaral and Taltal“, and „The flora of the Nitrate Coast“ (�9�9 a,b), are considered classical studies of the northern Chilean coast, where vegetation is so highly related to sporadic precipitacion due to El Niño events (Dillon et al. �00�) and the fog‘s water content (Muñoz-Schick et al. �00�).

4�

Mutisia decurrens, from Sinop-sis de la Flora Chilena (�9��)

Carl Skottsberg (�880-�9��), the Swedish botanical eminence, pre-sident of the 7th International Botanical Congress in �9�0, is one of the most impressive botanists that explored the Chilean territory. He came first with the Swedish expedition to the South Pole bet-ween �90� and �90� and afterwars explored several times mainly Patagonia and the Pacific islands. He published about 128 articles and several books about the Chilean botany and plant geography. His Vegetationsverhältnisse längs der Cordillera de los Andes süd-lich von 41 Grad... (�9��) and his Natural History of Juan Fernan-dez and Easter Island (�9�0-�9��) are till today the most complete works that treat these territories.

Carlos Muñoz Pizarro (�9��-�97�), was a student of Francisco Fuentes. After a Guggenheim scho-larship in the USA, under the direction from I.M. Johnston, he was encharged of the Botanical Section at the Museo Nacional in �94�. He initiated a program of organisation of the collections. Around �0,700 exemplars were revised, catalogued and organized. In �9��, with a grant from the Rockefeller Foundation, Muñoz Pizarro explored, together with his wife Ruth Schick, the principal European herbaria, carrying out the photographic registry of the type collections of Chilean plants. The acquired knowledge allowed him to publish several important books: Sinopsis de la Flora Chi-lena (1959, 2d ed. 1966), the principal synthetical work of the Chilean flora still in use; Flores Sil-vestres de Chile (�9��); Chile: Plantas en extinción (�97�).

Box �.� The discovery of the Chilean plant world (continuation)

4�

2.2 Classification of the Chilean plants: a modern (but not definitive) synthesis Keeping in mind the discussion in section 1.2, there are several classification systems in which we can organize the Chilean Flora. Taxonomy is mainly a modern European task, transferred to the rest of the world since Linnaeus invented the binomial classification. There are also traces of an aboriginal taxonomy, and Villagrán (�00�) suggests to recognize this millenary task as a ciencia indígena. But other languages dominate the sciences, and most of the aboriginal knowledge went lost con la cruz y la espada.The system used in the present thesis for organizing and analysing the Chilean families and genera is the one proposed by Pryer et al. (2004) for the vascular plants (figure 2.1) and specifically the APG system in its 2d version (APG II 2003) for the angiosperms (figure 2.2), with further updates, as continuously tracked by Stevens (�00� onwards).

2.2.1 Floristic composition: major families and genera

Reiche’s (�907) statistics were composed of �4� families and 7�� genera for the Chilean Flora (table �.�), as he critically revised previous Philippi’s account of 8�� genera, as preparing the new Flora de Chile (Reiche �89�-�9��). Cullen & Walters (�00�) recognize a trend of increase in the number of families in different classifications in time, and in fact the number of genera and families has grown since Reiche’s revision. Muñoz Pizarro (�9��) in his Sinopsis de la Flora Chilena gave a number of �8� families and 9�0 native genera��, and Marticorena’s (�990) checklist considered �7� families an 8�7 genera (table �.�).

Leptosporangiate ferns x x x x x

Horsetails x x x x x

Marattioid ferns x x x x x x

Ophioglossoid ferns x x x x x x

Wisk ferns x x x x x

Gnetophytes x x x

Conifers x x x

Ginkgo x x x

Cycads x x x

Angiosperms x x x

Quillworts x x x x

Spikemosses x x x x

Clubmosses x x x x

Lyco

phyte

s

Trach

eoph

ytes

Euphy

lloph

ytes

Sperm

atoph

ytes

Monilo

phyte

s = fe

rns

„Pter

idoph

ytes“

„Fern

s and

...“

„...fer

n allie

s“

„Eup

orang

iate f

erns“

Trac

heop

hyte

s

Lyco

phyt

es

Eup

hyllo

phyt

es

Spe

rmat

ophy

tes

Mon

iloph

ytes

= fe

rns

Monophyletic Paraphyletic

Figure �.� Cladogram showing relationships among the major lineages of vascular plants (from Pryer et al. �004)

4�

Table �.� Historical statistics for the Chilean native vascular plants

Orders Families GeneraF. Philippi �88� - - 8��Reiche �907 - �4� 7��Muñoz Pizarro �9�� 9� �8� 9�0*

Marticorena �990 - �7� 8�7* including many naturalized

Recently, incorporating molecular data, some families have been disintegrated, like the Scrophulariaceae; most of the Chilean genera are today classified under the Plantaginaceae (Albach et al. �00�). Several genera should join the new proposed families Gratiolaceae or Linderniaceae (Rahmanzadeh et al. �00�). On the other side, also former Scrophulariaceae have been erected to

NymphaeaceaeAustrobaileyales

Canellales 1/1

Laurales 4/6Magnoliales

Piperales 3/3

Chloranthaceae

AcoralesAlismatales 7/15Asparagales 10/38Dioscoreales 1/2Liliales 4/7PandanalesArecales 1/2

CommelinalesPoales 7/102

Zingiberales

Ceratophyllales 1/1

Ranunculales 4/10Proteales 1/4

Berberidopsidales 2/2Gunnerales 1/1

Caryophyllales 12/69Santalales 3/9Saxifragales 4/10

CrossosomatalesGeraniales 4/9Myrtales 3/17Celastrales 1/1

Cucurbitales 2/2

Fabales 2/25

Fagales 2/2

Malpighiales 10/20Oxalidales 3/6

Rosales 3/21

Brassicales 4/28Malvales 2/13Sapindales 3/9

Cornales 2/8Ericales 6/17GarryalesGentianales 3/16Lamiales 15/58Solanales 2/29

Apiales 3/ 26Aquifoliales

Asterales 5/137Dipsacales 1/3

Amborellaceae

aste

rids

rosi

dscore

eud

icot

s

eudi

cots

mon

ocot

sangi

ospe

rms

euasterids II

euasterids I+ unplaced Boraginaceae /13

+ unplaced Desfontainiaceae /1+ unplaced Escalloniaceae /2

+ unplaced Icacinaceae /1

+ Vitales 1/1

eurosids II

eurosids I

magnoliids

commelinids

+ Zygophyllales 2/7

Figure 2.2 Cladogram showing relationships between angiosperm orders (slightly modified from APG 2003). Num-bers represent native families/genera present in Chile. The absence of number means no representation in Chile.

44

family status, i.e. Calceolariaceae (Olmstead �00�). Chenopodiaceae are now being submerged into Amaranthaceae; Empetraceae and Epacridaceae into Ericaceae (Kron �99�, Kron & Luteyn �00�).

With gains and losses, the statistics of the Chilean flora stayed more or less stable during the last 15 years. Anyhow, under this dynamic scenario, a synthesis of the flora is a work in progress (Marticorena �990). After a revision of the most recent literature and a couple hundreds of monographs, the following account for the Chilean flora can be given, updated to 31 January 2007 (table 2.2, appendix A). The extant Chilean vascular native flora is composed by:

59 orders, 179 families, 813 genera, and about 4.333 species

This statistics include the offshore oceanic flora; Pacific genera not represented in the continent, and other endemic genera (+ one endemic family) are shown in table �.�.

Table �.� Summary of Chilean plants (updated �� December �00�)

Orders Families Genera Native speciesferns and fern allies �8 �4 �0 ���gymnosperms � 4 9 ��monocots � �0 ��� 880dicots �� ��� �9� �,���TOTAL �9 �79 8�� 4,���

Table 2.3 Genera not found in continental Chile (* = family also absent from the continent)

Genus (family)Juan Fernández Islands ferns and fern allies Dicksonia (Dicksoniaceae)*

Thyrsopteris (Dicksoniaceae)*Arthropteris (Oleandraceae)*

gymnosperms -

monocots Juania (Arecaceae)Machaerina (Cyperaceae)Megalachne (Poaceae)Podophorus (Poaceae)

dicots Centaurodendron (Asteraceae)Dendroseris (Asteraceae)Robinsonia (Asteraceae)Yunquea (Asteraceae)Selkirkia (Boraginaceae)Haloragis (Haloragaceae)Cuminia (Lamiaceae)Lactoris (Lactoridaceae)*Zanthoxylum = Fagara (Rutaceae)Coprosma (Rubiaceae)x Margyracaena (Rosaceae)Santalum (Santalaceae)Boehmeria (Urticaceae)

4�

Desventuradas Islands ferns and fern allies -gymnosperms -monocots -dicots Lycapsus (Asteraceae)

Thamnoseris (Asteraceae)Nesocaryum (Boraginaceae)Sanctambrosia (Caryophyllaceae)

Isla de Pascua ferns and fern allies Davallia (Davalliaceae)*Microlepia (Dennstaedtiaceae)Diplazium (Dryopteridaceae)Dryopteris (Dryopteridaceae)Microsorum (Polypodiaceae)Psilotum (Psilotaceae)*Vittaria (Vittariaceae)*Doodia (Blechnaceae)

gymnosperms -monocots Kyllinga (Cyperaceae)

Pycreus (Cyperaceae)Axonopus (Poaceae)Stipa (Poaceae)

dicots Triumfetta (Malvaceae)Ipomoea (Convolvulaceae)

In the angiosperms, richest Chilean orders are the Lamiales (�� families, �8 genera), Caryophyllales (��/�9), Asparagales (�0/�8), Malpighiales (�0/�0), Poales (7/�0�), Alismatales (7/��), Ericales (6/17) and the Asterales (5/137) (figure 2.2).

The Asteraceae, usually the largest family in floras of arid or semi-arid regions (Goldblatt & Manning 2000), is also the most species- and generic-rich family in the Chilean flora (table 2.4). The second largest family is the Poaceae, what is also expected due to the global high richness of the family. Apiaceae, Brassicaceae, Fabaceae and Solanaceae follow in size, with Fabaceae behind Poaceae in species numbers.

Table 2.4 Ranking of the 20 largest families in the Chilean flora by size

FAM N° genera N° Chilean species Asteraceae ��� 8��Poaceae �7 �9�Apiaceae �4 9�Brassicaceae �� ���Fabaceae �� �7�Solanaceae �� ��4Caryophyllaceae �8 ��Cactaceae �� 9�Cyperaceae �� �4�Boraginaceae �� �07Rosaceae �� 4�Verbenaceae �� 8�Malvaceae �� 7�Alliaceae 9 �9Myrtaceae 9 ��Amaranthaceae 9 47Iridaceae 9 �8Campanulaceae 8 �7Lamiaceae 8 �9Plantaginaceae 8 4�

4�

The most species-rich genera are Senecio, Adesmia, Oxalis, Viola, Haplopappus, Poa, Carex, Solanum, Calceolaria, Berberis (table �.�). Some of these genera are cosmopolitan species-rich genera (e.g. Senecio, Oxalis, Viola), while others are strict neotropical genera (e.g Haploppapus, Calceolaria). Adesmia is a very interesting case, being the most species-rich genus restricted to the southern Andes of Chile/Argentina (with some representation in Perú and southern Amazonas) (a recent biogeographic analysis of Adesmia has been done by Mihoc et al. �00�).

Table 2.5 Ranking of the 20 largest genera in the Chilean flora (including Hypochaeris, Alstroemeria, Verbena each with �� spp.)

GEN N° Chilean species Senecio ��8Adesmia ���Oxalis ��7Viola 70Haplopappus �4Poa ��Carex ��Solanum ��Calceolaria �0Berberis 47Astragalus 4�Baccharis 4�Valeriana 4�Leucheria 4�Dioscorea 4�Cryptantha �9Chaetanthera �7Nolana ��Loasa ��Hypochaeris, Alstroemeria, Verbena ��

2.2.2 Endemic families

The Chilean flora comprises three monotypic or dytipic endemic families, Gomortegaceae, Francoaceae, and Lactoridaceae, this latter endemic to Juan Fernández (table �.�). Also there are four subendemic families restricted to Chile and adjacent territories in Argentina and Peru (table 2.7). The Chilean flora harbours 83 endemic genera, that will be further analysed in chapter 3.6.

Table 2.6. Endemic families of the Chilean flora

Family Genera N° speciesGomortegaceae Gomortega �Francoaceae Francoa

Tetilla ��

Lactoridaceae Lactoris � (Juan Fernández)

Table 2.7 Subendemic families of the Chilean flora (+adjacent areas in Argentina/Peru)

Family Genera N° species (Chile)Aextoxicaceae Aextoxicon � (�)Misodendraceae Misodendrum 8 (8)Philesiaceae Lapageria

Philesia � (�) � (�)

Malesherbiaceae Malesherbia �4 (�8)

47

Just as a brief comparison, Argentina harbours one endemic family (Halophytaceae) and 4� endemic genera (Zuloaga et al. 1999). The flora of Perú, composed of more than 17.000 species, and around �.400 native genera�4, has �� endemic genera but do not show any endemic family (Brako & Zarucchi 1993). The flora of Ecuador (2.110 native genera, and around 15.000 species), shows �� endemic genera and also none endemic family (Jørgensen & León-Yáñez �999).

2.3 Geographical classification of the Chilean flora Stuessy (2006) touches indirectly the task of biogeographical classification while listing the six principles of plant taxonomy. Principles 4 and �, at least, are more general, and applicable to biogeography:Pr. 4: Humans need hierarchical systems of information storage and retrieval to live and survive, including dealing with the living world.Pr. �: The assessed patterns of organismal relationship are used to construct hierarchical classifications of coordinate and subordinate groups that are information-rich and have high predictive efficacy; these are the taxonomic hypotheses that change with new information and new modes of analysis. While the hierarchical system of botanical regions has been otherwise criticised and compared with the hierarchical terminology of Roman military administration and control (Grehan �00�), others have shown that the biogeographical hierarchies are real (McLoughlin �99�, �994). Hereafter I present a summary of the different ways in which the Chilean flora has been classified at the global level (table �.8).

Table 2.8 The geographical classification of the Chilean flora

N° kingdoms

N° regions

G.R. Treviranus �80� 8 ‘Flor’ - Antarktische FlorAugust Pyramus de Candolle �8�0

- �0 le Chili and les terres Magellaniques

Schouw �8�� �� - South of 42° = Antarktisches Florenreich; 42° - 23° = Reich der Holzartigen Synanthereen (Compositae); North of 23° = Reich der Cactus und Piper

Alphonse de Candolle �8��

- 4� Region ��, Le Chili, and region ��, la Patagonie, la terre de Feu et les iles Malouines (Falkland).

Grisebach �87� - �4 Tropische Andean Flora = North to 23°; Chilenisches Übergangsgebiet = 23°-34°, Antarktisches Waldgebiet = 34°-��°

Engler �879-�88� 4 �� Südamerikanisches Florenreich = to 41°, Altoceanisches Florenreich = south from 41°

Drude �884 �4 �� South of 41° = Antarktisches Florenreich; North of 41° = Andines Florenreich

Drude �890 �4 �� Andines Florenreich and Antarktisches FlorenreichDiels �908 � 7 Antarktisches Florenreich and Neotropisches FlorenreichGood �947 (�974) � �7 Antarctic kingdom, Patagonian region, South of 4�°;

Neotropical kingdom, Andean region, North of 4�°.Mattick �9�4 � 4� South = Antarktisches Florenreich; North = Neotropisches

Florenreich

48

Takhtajan �9�� � �7 Antarctic kingdom, Patagonian region, South of 40°; Neotropical kingdom, Andean region, North of 40°.

Takhtajan �978 � �� Holantarctic kingdom, Chile-Patagonian region, South of ��° to Antarctic peninsula and Malvinas Islands (Falkland Is.)

Cox �00� � - Southamerican kingdomMorrone �00� � �� Austral kingdom, Andean region

2.3.1. The global classification

Gottfried R. Treviranus (�77�-�8�7), one of the naturalists who coined the term biology (Engels 2005) first intended a global floristic classification, organizing the world flora in 8 principal floras =Hauptfloren. This early classification included an Antarctic Flora (Antarktische Flor), which comprises Chile, Magallanes, Tierra del Fuego, and New Zealand (Treviranus �80�). Treviranus is to my knowledge the first biologist to recognise explicitly the floristic relationship between southernmost South America and Australasia, based on the early works of J.I. Molina (�740-�8�0), J. Banks (�74�-�8�0), and G. Forster (�7�4-�794)��. Treviranus noticed the floristic relations between New Zealand and Tierra del Fuego and also the existence of an antitropical floristic element, i.e. genera present in temperate areas from both hemispheres but absent in the tropics, such as Pinguicula, Salix, Fagus or Ribes. The relation appeared to him surprising since to that time the known flora of Tierra del Fuego was composed by less than 40 species! (Treviranus 1803, p. ���).

At this early stage biogeographic representation was inexistent, since the discipline was at its early stages of development. Only � years later Jean B. Lamarck and Augustin-Pyramus de Candolle published the “first biogeographical map” for the third edition of the Flore française (Ebach & Goujet �00�). A-P. de Candolle, in his Géographie botanique further classified the world in 20 floristic regions: Chile fitted into two regions, le Chili and les terres Magellaniques (de Candolle 1820). But de Candolle’s world classification still lacked a map. Three years later, Danish botanist Joakim Frederik Schouw (1789-1852) published the first ptytogeographical world map (Schouw 1823, Mennema 1985). Schouw proposed 25 floristic realms =Florenreiche. Southern South America was classified into two realms: the Reich der Holzartigen Synanthereen, and the Antarktisches Reich, from 40°S to the South. Applying the concept of endemism recently proposed by A-P. de Candolle, Schouw exposed explicitly the criteria for classifying and delimiting the floristic realms (Drude 1884, p. 13):

�. half of the known plants have to be native to the territory in question;�. ¼ of the genera had to be endemic or have their maximal distribution there;�. A plant realm had to have some endemic families.

De Candolle’s son Alphonse de Candolle (�8��) disputed the criteria Schouw used and proposed 4� botanical regions, but Chile maintained the division into two regions. But soon de Candolle the younger rejected such schemes of regions and turned to be the first critic of the task of

49

floristic classification: “Je tiens donc les divisions du globe par régions, proposées jusqu’à present, pour des systèmes artificiels.... Elles ont nui a la science”�� (A. de Candolle �8��, p. ��04-��0�, as quoted by Nelson �978). Later he states, with reference to his Geographie: “ouvrage du reste complètement différent de celui auquel mon père pensait, car les documents etaient devenus plus nombreus, et mes idées s’étaient singulièrement éloignees de celles qui régnaient dans la science depuis vingt ans “ �7(A. de Candolle Mémoires, p. �9�, as quoted by Nelson �978).

It seems to be that as botanical information became to be overwhelming, the task of phytogeographical classification was getting more and more difficult. One of the key naturalists in this growing botanical knowledge was J.D. Hooker (�8�7-�9��). While sailing on board James Cook’s Endeavour, he notably improved the floristic knowledge of the southern hemisphere. Hooker’s publications compiled as The Botany of the Antarctic Voyage (Hooker �844-�8�0), were almost as epoch-making as Darwin’s Origin of species (Thiselton-Dyer �909).

By the second half of the 19th century a huge amount of floristic knowledge had accumulated. This knowledge, coupled with the ecological principles developed since A. von Humboldt, permitted August Grisebach (�8�4-�879) to publish his Vegetation der Erde, which related explicitly the plant world with the regional climates (Grisebach �87�). In Grisebach’s view, Chile had to be classified into 3 regions =Gebiete: 1. a Chilean floristic core, Chilenisches Übergangsgebiet (transition zone) from 23° to 34°, that holds a “unique flora”; 2. an Antarctic region (Antarktisches Waldgebiet), ranging from �4° to ��°, characterized especially by the genus Nothofagus; and �. a tropical Andean flora that ranges from Ecuador to northern Chile (figure 2.4).

Adolf Engler (�844-�9�0), one of the most prominent scientists in botanical history, working at the Botanical Garden in Berlin, was the first one to try a synthesis of the evolution of the plant world on the earth surface (Engler 1879, 1882). He classified the world flora into four realms and 32 regions, dividing also each region into diverse provinces and districts, thus constructing the hierarchical system that forms the base of all the following classification systems. Chile was classified into the Südamerikanische Florenreich (recognizing the Nordchilenische Provinz as the Chilenische

Übergangsgebiet from Grisebach) and in the Altoceanisches Florenreich, South of ��°S. This Altoceanisches Florenreich grouped southern Chile with New Zealand’s South Island, the sub-Antarctic islands, most of Australia and the Cape region from Africa (figure 2.5).

“Engler was surprisingly perceptive in realizing that, scattered over the islands and lands of the southernmost part of the world, lay the remains of a single flora, which he called ‘the Ancient Ocean’ flora. It was over 80 years before acceptance of the movement and splitting of continent at last explained this very surprising pattern of distribution” (Cox & Moore �00�, p. ��).

�0

Figu

re �

.4 V

eget

atio

n of

the

wor

ld, a

fter G

riseb

ach

(�87

�, �

d ed

ition

�88

4)

��

Figu

re 2

.5 B

otan

ical

regi

ons a

nd re

alm

s, flo

ristic

and

phy

siog

nom

ic (E

ngle

r 188

2)

��

Later, Engler suggested Australe Florenreich would be a better name as it is characterized by the Austral-antarktischen Florenelement (Engler �899, p. �49) as well as adding a fifth kingdom, the Ozeanisches Florenreich, which was composed of the aquatic plants from the vast oceans.

Oscar Drude (�8��-�9��) worked close with A. Engler from the Botanical Garden in Dresden. Being a student from Grisebach, Drude found several difficulties in synthesizing the floristic knowledge of his predecessors with the growing ecological (physiognomic) knowledge as

Figu

re �

.� F

loris

tic re

alm

s fro

m D

rude

(�88

4)

��

systematized by Grisebach (1872). Drude first published his work Die Florenreiche der Erde based on a floristic approach (Drude 1884). He therefore defined 14 floristic kingdoms and 55 floristic regions =Gebiete. In this scheme, northern Chile to 4�°S corresponds to the Andines Florenreich and southern Chile to the Antarktisches Florenreich (figure 2.6).

Drude’s concern over about the floristic and ecological differences led him to publish separate maps for a floristic classification and for a vegetation classification (Drude 1887). Three years later he abandoned the floristic classification altogether: “The maps published in the geographical reports of 1884 about my floristic classification show the uncertainty of the boundary lines due to numerous migration routes and directions of dispersal, which overlap from one to the other realm; it is long a known fact that each attempt to draw strict floristic boundaries, is itself ruinous” (Drude �890, p. ��9). Drude decided to join the more physiognomic approach from Grisebach, and he modified the classification on the basis of the new climatological basis provided by Wladimir Köppen (1884). The floristic kingdoms still numbered 14, and South America remained unmodified.

At the beginning of the �0th century, Ludwig Diels (�874-�94�), successor of Engler in Berlin, synthesized Drude’s classification into six floristic realms, following the early proposal from Engler (�88�) but obviating the oceanic realm (Ozeanisches Florenreich), and dividing Engler’s Altoceanisches Florenreich into an Antarktis, an Australis and a Capensis (figure 2.7). Diels (1908) was the first to raise the African Cape region to the category of a realm. Interestingly he considered the Australisches Florenreich (Australis) as comprising only Australia and Tasmania -

Figure �.7 Floristic realms from Diels (�908)

�4

- and considered Malesia and New Zealand as part of the Paläotropis. South America and Central America including Mexico and Baja California was part of the Neotropis, but southernmost South America retained its designation as a realm, the Antarktis. Diels (�908) little book was reprinted five times until 1958, and his realm classification was retained adding only more details at the regional scale by Mattick (�9�4) and later popular authors like Good (�974) and Takhtajan (1978). Diels’ proposal modified by the mentioned authors is still preferred in all modern German phytogeography textbooks (e.g. Richter �997, Schroeder �998, Frey & Lösch �004).

English botanist Ronald Good’s (�89�-�99�) The Geography of the Flowering Plants (Good 1947), was to become one of the most popular books in the field, reaching to four editions and two reprints from �947 till �974. He followed Diels with the � realms, dividing them in �7 regions (figure 2.8). Chile south of 41°, was classified into the Antarctic kingdom, and as the Neotropical kingdom to the north. The scheme is very similar to that of Russian botanist Armen Takhtajan, which became very popular after its translation into English. In a first version he maintained the 6 realms from Diels and the 37 regions from Good (Takhtajan 1961) (figure 2.9), but in his definitive proposal he reduced the regions to 35. Diel’s scheme of 6 realms stayed unchanged (Takhtajan 1978) (figure 2.10). In his first classification he considered Central Chile as part of the Neotropical region (Takhtajan 1961) (figure 2.9), but in his later work he classified all the southern cone S of 25° into the Holantarctic kingdom (Takhtajan 1978) (figure 2.10).

The basic scheme of 6 floristic realms proposed by Diels (1908) stayed unchanged during the 20th century; only at the beginning of this century, Cox (�00�) took the task to deeply reanalyse both floristic and faunistic long standing schemes (the faunistic regions date back to Sclater [1858] and Wallace [1876]). For the global flora the proposition was a rearrangement of the six former floristic realms into five: the Holartic, South American, African, Indo-Pacific, and Australian.

Figure �.8 Floristic regions from Good (�947)

��

Figure �.9 Floristic regions from Takhtajan (�9��)

Regards the former Antarctic kingdom he wrote: “… the consistency of the plant geographical system is better served by transferring some of the regions of the Antarctic Kingdom to the South American Kingdom and the rest to the Australian Kingdom, in each case noting their individual historical and ecological characteristics”.

Cox (�00�) discuss the Takhtajan’s Holantarctic kingdom listing the �� endemic families and �4 endemic genera proposed by Takhtajan (�978). Takhtajan’s Holantarctics is based at the family level mainly on American endemic families (e.g. Thyrsopteridaceae, Lactoridaceae, Gomortegaceae, Aextoxicaceae). The dominance of American families seems to give reason to Cox while transferring them to the South American kingdom.

The last word stays in Morrone (2002), who challenged Cox’s proposal and compiles the floristic and faunistic knowledge in one synthetic classification. The result is a scheme of only 3 biotic realms: the Holarctic kingdom, the Tropical kingdom (=East Gondwana), and the Austral kingdom (=West Gondwana). Morrone (2002) related the classification to the history of these biotas, as was Engler’s early intention (�879, �88�). In fact the result is remarkably similar to Engler’s, but grouping the paleotropis and neotropis in one tropical realm.

In Morrone’s proposal the austral kingdom is composed by S Australia, NZ, S Africa and S South America, extending through the Andes till �°N in Colombia. He argues that a “single biogeographical scheme for all organisms, to serve as a general reference system, would be a desirable goal”. Morrone briefly mentions recent cladistic biogeographical analysis for his proposal, but do not provide explicit account of biotic similarities between the territories classified into the Austral kingdom, as did Cox & Moore (�00�, p. ���) for intercontinental relationships.

��

Figu

re 2

.10

Bot

anic

al re

alm

s fro

m T

akht

ajan

(197

8). I

= H

olar

ctis

, II =

Pal

eotro

pis,

III =

Neo

tropi

s, IV

= C

apen

sis,

V =

Aus

tralis

, VI =

Hol

anta

rctis

.

�7

2.3.1. The regional classification

Specific floristic classifications at the regional (national) level are very scarce in Chile, being the efforts traditionally concentrated in vegetation mapping (see next section 2.4) The first plant geographical map for Chile accompanied Reiche’s analysis of the distribution of the Compositae family in the country (Reiche 1905) (figure 2.11). On a second map he also proposed possible migration routes for these taxa (Reiche 1905) (figure 2.12). The first (and only) intend of a synthetical cartography for the Chilean flora is expressed in the two maps that accompanied Reiche’s Plant geography (Reiche 1907) (figure 2.13, 2.14). Here Reiche proposed distribution ranges and limits for some key taxa in his view. In a second map he divided the country in several floristic units, integrating floristic and physiognomic knowledge. Reiche proposed the limit between the antarctic and neotropical reamls at around 4�°S. In spite of appearing so simple, it is notably the first intend of a floristic cartography for Chile. 50 years later, Schmithüsen drew a schematic representation of the different floristic elements composing the woody flora between 30° and 42°S. (Schmithüsen 1956) (figure 2.15). The floristic cartographic task at the national level seems since then to be frozen in time, having been replaced by more general biogeographical classifications (e.g. Cabrera & Willink 1973; Rivas-Martínez & Navarro �000; Morrone �00�, �00�; see a discussion in Ribichich �00� for the task in adjacent Argentina).

Figure 2.11 The first plant geographical map for Chile (Reiche �90�)

�8

Figure �.�� Possible migration routes for the Chilean Compositae (Reiche �90�)

Figure �.�� Geographic ranges and distribution limits of selected taxa (Reiche �907)

�9

Figure �.�4 Plant geographical divisions of Chile (Reiche �907)

Figure 2.15 Diagram of floristic elements in Central Chile (Schmithüsen �9��)

1. General floristic area of neotropical origin

2. Forest flora of south-he-mispheric-subtropical origin

�. Laurophyllous neotropical and suth-hemispheric origin

4. Neotropical sclerophyllous element

�. Temperate rainforest of neotropical and south-hemis-pheric origin

6. Neotropical endemic flora from La Serena

7. Decoduous forest of suban-tarctic origin

8. Evergreen forest of subant-arctic origin

�0

2.4 Excursus: vegetation maps and Vegetationsbilder

Contrary to the floristic cartography, the vegetation cartography has seen a good development in Chile. Mainly on the base of the early climatic classification (Köppen �9�0), followed by bioclimatic proposals for the country (di Castri �9�8, di Castri & Hajek 1976, Quintanilla 1974, Amigo & Ramírez 1998), the vegetation cartography has developed in hand of Schmithüsen (1956) (figure 2.16), Hueck (1978), Pisano (1977, 1981, 1983), Quintanilla (�98�, �98�, �988), Gajardo (�994), and Luebert & Gajardo (�00�). These efforts and the use of global climatic surfaces on a GIS-based platform allowed Luebert & Pliscoff (�00�) to publish the most accurate bioclimatic and vegetation synthesis to date. The early works of C. Skottsberg deserve special attention (Skottsberg �9�0a, �9��). Due to the impressive floristic knowledge of the eminent botanist, his maps can be considered a good synthesis of floristic and physiognomic information (figure 2.17). Remarkably are also Skottsberg’s three volumes on the Natural History of Juan Fernández and Easter Island (Skottsberg �9�0-�9��), as well as his Vegetationsbilder, that spread the images of Chilean landscapes at the beginning of the �0th century (Skottsberg 1906, 1910b) (figure 2.18).

Figure �.�� Schmithüsen‘s vegetation map (�9��):1= north-Andean vegetation; 2= desert; 3= semi-desertic sc-rub; 4= fog-forest; 5= xeric scrub; 6= sclerophyllous mator-ral; 7= temperate forest; 8a= Valdivian rainforest; 8b= north-patagonic rainforest; 8c=subantarctic rainforest; 9= tundra; 10=subantarctic deciduous-forest, 11=east-patagonic steppe;

12=south-Andean vegetation.

��

Figure �.�7 Phytogeographic map from Skottsberg (�9�0a)

��

Figure �.�8 Vegetationsbild from Skottsberg (�9�0 b)

��

2.5 Geographic ranges in the latitudinal profile

The distribution of the genera along the latitudinal profile in Chile is based on the collections of the National Herbarium (SGO), that have been partially transferred to a data base with geographic coordinates (see Muñoz-Schick & Moreira-Muñoz �00�). The data base is a work in progress, and ca. �0 000 records have been checked so far, from ca. �00 000 specimens of native vascular plants registered at SGO. Dot maps have been done as an event theme in ArcView �.� GIS program. Every genus has been checked for false or lacking coordinates, and for evident wrong determinations. Also very helpful for plotting the distribution has been the revision of several regional floras and checklists (Skottsberg 1916, Moore 1983, Henríquez et al. 1995, Marticorena et al. 1998a, 2001) and local floras and checklists (e.g. Muñoz-Schick 1980, Teillier et al. 1994, �00�; Richter �99�, Rundel et al. �99�, Arroyo et al. �998, �000, �00�, �00�; Luebert & Gajardo �000, �00�; Luebert et al. �00�, Villagrán �00�).

A summary of the maximal latitudinal range that occupy the families and genera in continental Chile is presented as figures 2.19, 2.20, 2.21, 2.22). The taxa are organized from North to South on the base of the average of the distribution. The latitudinal extention of the family ranges represent the averaging distribution of the composing genera (extracted in MS Access).

The genera were further classified into five classes:

a) �0� genera occur in Chile in a very small range (<� latitude degree) or just in few localities. Many of these genera are endemics (e.g. Robinsonia, Cuminia) or have an occurrence only in the Chilean oceanic islands (e.g. Psilotum, Stipa, Vittaria), but others show as well such a restricted range in the continent (e.g. Androsace, Achyrocline, Grabowskia, Avellanita, Menodora, Pouteria);

b) ��� genera show a small distribution range between � to � degrees of latitude (e.g. Aphanes, Traubia, Guindilia, Orites);

c) ��4 genera have a medium- small range of distribution from � to �0 latitudinal degrees (e.g. Luciliocline, Hebe, Quillaja, Aa, Lepidoceras);

d) ��� genera show medium-large ranges of distribution, from �0 to �0 latitudinal degrees (e.g. Acacia, Podocarpus, Cissus, Oziröe, Azara);

e) ��� genera have ranges larger than �0 latitudinal degrees (e.g. Senna, Griselinia, Azorella, Ourisia, Calandrinia, Schinus). The most widely distributed genera in the latitudinal profile are Perezia, Baccharis, Colobanthus, Juncus, and Senecio, which occupy the whole latitudinal profile from Parinacota (�7°��’) to Cabo de Hornos (��°).

�4

Also the distribution of the taxa was summarized in �0 degree latitudinal bands, from 17,6° to 27° (arid tropical zone) (figure �.�0); �7° to �7°S (Mediterranean-type zone) (figure 2.21); and 37° to 56S° (temperate zone) (figure 2.22).

Figure �.�9 Latitudinal ranges for Chilean families

-57 -47 -37 -27 -17

MyricaceaeBalanophoraceaeCleomaceaeKrameriaceaeNyctaginaceaeCactaceaeMalesherbiaceaeCucurbitaceaePapaveraceaeMalpighiaceaeHyacinthaceaeRafflesiaceaeThelypteridaceaeHydrocharitaceaeZygophyllaceaeAnacardiaceaeVerbenaceaeTecophilaeaceaeTyphaceaeAizoaceaeAcanthaceaeLythraceaeValerianaceaeAmaryllidaceaeLedocarpaceaePteridaceaeZosteraceaeLoasaceaeAzollaceaeAristolochiaceaeSolanaceaeAmaranthaceaeHemerocallidaceaePortulacaceaeAlstroemeriaceaeFabaceaeCaricaceaeMalvaceaeEquisetaceaeApocynaceaeBignoniaceaeCaryophyllaceaeSapindaceaeLaxmanniaceaeAsteraceaeDioscoreaceaePlumbaginaceaeOrobanchaceaePassifloraceaeOleaceaeSapotaceaeConvolvulaceaeBromeliaceaeVivianiaceaeArecaceaePoaceaeSalicaceaeLamiaceaePolygonaceaePiperaceaeBrassicaceaePhrymaceaeEuphorbiaceaeMarsileaceaeScrophulariaceaeLinaceaeBoraginaceaeQuillajaceaePolemoniaceaeAlliaceaeMolluginaceaeIcacinaceaeGeraniaceaeGentianaceaeCampanulaceaeGratiolaceaeRosaceaeCeratophyllaceaeUrticaceaeEphedraceaeGoodeniaceaeMonimiaceaeFrancoaceaePolypodiaceaeOnagraceaeDryopteridaceaeIridaceaeLauraceaeApiaceaeAgavaceaeLoranthaceaeJuncaceaePhytolaccaceaeRutaceaeVitaceaeGomortegaceaeFrankeniaceaeViolaceaeCrassulaceaeOxalidaceaeGriseliniaceaeAextoxicaceaeElatinaceaeCalceolariaceaePotamogetonaceaeHaloragaceaeRhamnaceaeBerberidopsidaceaePolygalaceaeElaeocarpaceaeAlismataceaeOrchidaceaeLardizabalaceaeCalyceraceaeEscalloniaceaeAspleniaceaeRuppiaceaeRubiaceaeMyrsinaceaeLinderniaceaeHypericaceaeAraucariaceaeMyrtaceaeCyperaceaeCoriariaceaeGesneriaceaeTropaeolaceaeLemnaceaeHydrangeaceaePlantaginaceaeAtherospermataceaeBerberidaceaeDennstaedtiaceaeProteaceaeLentibulariaceaeLophosoriaceaeRanunculaceaeSantalaceaeCorsiaceaeJuncaginaceaeCupressaceaePhilesiaceaeCelastraceaeLomariopsidaceaeCunoniaceaePodocarpaceaeGrossulariaceaeGunneraceaeWinteraceaeBlechnaceaeAraliaceaeRestionaceaeHymenophyllaceaeDesfontainiaceaeSamolaceaeNothofagaceaeMisodendraceaeSchizaeceaeLuzuriagaceaeIsoetaceaeEricaceaeThymelaeaceaeGleicheniaceaeGrammitidaceaeDroseraceaeCentrolepidaceaeAsteliaceaeSaxifragaceaeOphiglossaceaeStylidiaceaePrimulaceaeLycopodiaceaeTetrachondraceae

��

Figure �.�0 Latitudinal ranges for Chilean genera with ave-rage distribution north of �7°S

-57 -47 -37 -27 -17

StangeaMniodesDissantheliumCremolobusStenomessonDielsiochloaCoreopsisAlchemillaAchyroclineWoodsiaChlorisSpilanthesTaraTrichoneuraTecomaNeowerdermanniaDunaliaPityrogrammaPlucheaBrowningiaMorellaHaplorhusLophopappusSalpichroaHaageocereusDistichiaTrixisEremodrabaPolylepisCorryocactusAnthochloaTunillaPlaziaNeuontobothrysOmbrophytumGomphrenaReicheellaDiplostephiumHeterospermaPhilibertiaOreocereusMetharmeLucilioclineMastigostylaEudemaChersodomaHelogyneXenophyllumLampayaTripogonAaMunroaCuatrecasasiellaCotulaBoutelouaExodeconusLeptochloaParastrephiaUrbaniaEnneapogonPycnophyllumMancoaSteviaVillanovaDaleaAllioniaUrmeneteaNotholaenaNasaAnatherostipaPalauaCleomeAphyllocladusNitrophilaNamaAlternantheraPortulacaLycopersiconTiquiliaSporobolusSalviaDrymariaTigridiaDomeykoaWerdermanniaIpomopsisBalbisiaAcantholippiaHuidobriaMicrosterisEremocharisDinemandraFuertesimalvaReyesiaMaihueniopsisSchkuhriaErechtitesZephyraEnceliaGeoffroeaDiclipteraCaesalpiniaCrotonBacopaFagoniaPerityleEulychniaGrabowskiaWedeliaOxyphyllumCumulopuntiaAloysiaKrameriaGypothamniumDeuterocohniaZameioscirpusRostrariaWerneriaHoffmannseggiaOrobancheCynodonVerbesinaDoniophytonOxychloëPitraeaPolyachyrusNototricheMalacothrixAmblyopappusTetragoniaCopiapoaLenziaProsopisLyciumChuquiragaBakerolimonEchinopsisMalesherbiaPennisetumSicyosCristariaArgemone

��

Figu

re �

.��

Lat

itudi

nal r

ange

s for

Chi

lean

gen

era

with

ave

rage

dis

tribu

tion

in C

entra

l Chi

le b

etw

een

�7°S

and

�7°

S

-57 -47 -37 -27 -17

LippiaSkytanthusTagetesOphryosporusMirabilisOziroeAlonaBartsiaBidensIvaniaTessariaPilostylesEriosyceChorizantheArgyliaMaireanaMentzeliaErrazuriziaOxythecaFlaveriaThelypterisElodeaCressaCardionemaCruckshanksiaLeontochirCaiophoraParonychiaViguieraTillandsiaEragrostisPintoaTrichoclineCerastiumParietariaCistantheMontteaAristidaPlumbagoHeliotropiumTarasaBomareaBalsamocarponGrindeliaSalixMiqueliopuntiaBoerhaviaPhrodusChaetantheraBahiaBulnesiaLupinusSennaSpergulariaVerbenaGymnophytonTyphaProustiaNolanaPellaeaMalvellaHomalocarpusAgalinisDinemagonumCestrumKurzamraGlandulariaPleurophoraMyrcianthesCalliandraCyphocarpusPectocaryaCuscutaLimoniumCordiaPhragmitesCentaureaPolycarponMontiopsisConantheraGalinsogaPuyaAmbrosiaMenonvilleaHeterozosteraChiropetalumMuhlenbergiaLinumVulpiaAzollaAnisomeriaCiclospermumLeucocoryneAristolochiaAraeoandraGuynesomiaBridgesiaAgeratinaSchizopetalonBrachycladosPatosiaCalyceraPasitheaSchizanthusNicotianaDodonaeaMicrophyesHeleniumPleocarphusVasconcelleaEpipetrumEquisetumMarsileaMicroserisTecophilaeaCicendiaCyperusPaspalumAsterisciumFlourensiaCryptanthaCarpobrotusLeunisiaTrichopetalumGethyumPachylaenaLastarriaeaZoellneralliumLlagunoaFabianaMathewsiaTropidocarpumBarneoudiaPorlieriaTweediaPassifloraLemnaCheilanthesSchinusPolypogonMenodoraSolanumLotusDistichlisMelicaConvolvulusRhodophialaTeucriumTrifoliumTriodanisPouteriaAcaciaMuehlenbeckiaFacelisAndeimalvaTraubiaGayophytumBipinnulaDioscoreaTriptilionPlaceaLarreaVivianiaLaretiaEryngiumDichondraAristeguietiaStachysJubaeaTropaeolumErodiumAlonsoaMoschariaTrevoaPhycellaMarticoreniaMiersiaPleopeltisDiposisMulinumCentauriumMutisiaAdiantumHaplopappusPteromonninaTetillaGuindiliaCorrigiolaStemodiaSalpiglossisSpeeaSphaeralcea

-57 -47 -37 -27 -17

AdenopeltisPeperomiaMelospermaLigariaMimulusWeberbaueraCalydoreaNassellaStenandriumMicropsisAvellanitaPsilocarphusCalopappusQuillajaLithreaDennstaedtiaGochnatiaPodanthusRhombolytrumGlinusLinanthusMargyricarpusCryptogrammaGiliaKageneckiaLobeliaLudwigiaQuinchamaliuDiplolepisBeilschmiediaScyphanthusBuddlejaRetanillaCitronellaDowningiaPiptochaetiumChaptaliaMinuartiaLepechiniaOtholobiumSolariaPlectritisOchagaviaWahlenbergiaOenotheraClarkiaBlennospermaWolffiellaPozoaHydrocotyleNothoscordumBowlesiaColliguajaCeratophyllumJunelliaCamissoniaAphanesCryptocaryaGilliesiaNoticastrumPhalarisArtemisiaCastillejaMecardoniaCissarobryonImperataNastanthusStuckeniaTristerixLoasaAstragalusEuphorbiaEphedraPanicumPeumusSellieraJaravaPicrosiaAtriplexDescurainiaAcrisioneEscalloniaLapageriaHypochaerisGlycyrrhizaLuciliaBromidiumDaucusUtriculariaSolivaCentipedaLilaeopsisLegrandiaChenopodiumHypselaJaborosaGnaphaliumCalothecaWendtiaHerreriaLardizabalaMikaniaPilulariaPelletieraSaturejaCalandriniaLepuropetalonConyzaLeucheriaEleocharisMoschopsisRanunculusGamochaetaPhaceliaPitaviaAustrocedrusCalystegiaGymnachneCynoglossumHerbertiaCissusPhylloscirpusArenariaCystopterisDrabaGentianaLepidiumLimosellaSileneValerianaVestiaAdesmiaMaihueniaCalceolariaEvolvulusGeraniumHordeumEccremocarpuBromusSisymbriumCorynabutilonUrticaGomortegaCalamagrostisAlismaGutierreziaAzorellaEpilobiumWolffiaHybanthusAmaranthusBaccharisColobanthusJuncusPereziaSenecioFrankeniaFestucaScirpusHelictotrichonNavarretiaBlepharocalyxStellariaAspleniumCatabrosa

�7

Figu

re �

.��

Latit

udin

al ra

nges

for C

hile

an g

ener

a w

ith a

vera

ge d

istri

butio

n in

the

tem

pera

te z

one

sout

h of

�7°

S

-57 -47 -37 -27 -17

DichantheliumCrassulaGaliumOxalisPoaViolaSaginaOritesGriseliniaPlantagoSpartinaOurisiaBrachysteleAlstroemeriaAextoxiconSarmientaElatineCortaderiaMyriophyllumDanthoniaSarcocorniaApiumBerberidopsisAristoteliaPratiaDiosteaTetraglochinGlyceriaLumaSolidagoGamochaetopsisElymusCrinodendronSaniculaCliococcaAmsinckiaSuaedaLilaeaTriglochinPrumnopitysDasyphyllumSigesbeckiaNotantheraPodagrostisBelloaFrancoaJovellanaDesmariaLaureliaChascolytrumAzaraNardophyllumZannichelliaRuppiaChevreuliaRhaphithamnusPerseaPuccinelliaPolypodiumSagittariaPotamogetonLeptocarphaColletiaPolygonumRhamnusCarexGentianellaBulbostylisRhynchosporaMyrceugeniaLinderniaGratiolaHedyotisHypericumElytropusChusqueaMegalastrumSophoraRorippaAraucariaFasciculariaPleurosorusRelchelaLeptophyllochloaAcaenaLysimachiaErcillaCentellaAmphibromusGreigiaOlsyniumBoquilaCoriariaPhytolaccaLeptostigmaBlumenbachiaLepidocerasUgniHollermayeraSisyrinchiumHypolepisLegenereAnagallisAntidaphneValdiviaHydrangeaLomatiaAustrocactusViciaHabenariaMitrariaGevuinaBerberisComberaSolenomelusCallitricheTrisetumLibertiaFragariaFonkiaEucryphiaLophosoriaRytidospermaSaxe-gothaeaAmphiscirpusAmomyrtusLastheniaPterisLathyrusAsterantheraFitzroyaGamocarphaTrichomanesArachnitisPolygalaPileaAnarthrophyllumLatua

-57 -47 -37 -27 -17

MyosurusFuchsiaMaytenusElaphoglossumDiscariaOvidiaPlagiobothrysTristagmaAnemoneAntennariaMontiaPhleumCaldcluviaWeinmanniaDeschampsiaPolemoniumEuphrasiaMadiaRumexErigeronLaureliopsisAlopecurusArjonaDysopsisNassauviaRumohraOsmorhizaGunneraRibesNierembergiaChloraeaCalthaCollomiaCardamineLuzulaGavileaPodocarpusPolystichumDrimysKoeleriaMyoschilosBlechnumAsterArmeriaGaultheriaHymenophyllumNerteraUnciniaHymenoglossumAgoserisAgrostisOreopolusPseudopanaxApodasmiaOphioglossumCampsidiumDesfontainiaScutellariaNoccaeaTepualiaSamolusEmpetrumNothofagusGeumMisodendrumSchizaeaTaraxacumHuanacaHieraciumLagenophoraBoopisLuzuriagaEmbothriumSaxifragaCodonorchisIsoetesPinguiculaAdenocaulonSchoenusHippurisChiliotrichumMyrteolaOnurisAnthoxanthumOreobolusRubusLycopodiumMarsippospermumPterocactusMagallanaSchizeilemaGleicheniaMacrachaeniumGrammitisDonatiaDroseraPilgerodendronPrimulaLittorellaLepidothamnusPhilesiaOrtachneSerpyllopsisGaimardiaLeptinellaTribelesHistiopterisAsteliaNanodeaCarphaTapeiniaPhilippiellaAbrotanellaTetronciumLebetanthusBenthamiellaPhyllachneGrammospermaXerodrabaDrapetesEriachaeniumRostkoviaHebeMyosotisTetrachondraLecanophoraBolaxLepidophyllumHamadryasSaxifragodesChiliophyllumOreomyrrhisBotrychiumAndrosaceSaxifragellaChrysospleniumHuperziaLandoltia

�8

�9

� Geographic Relationships of the Chilean Flora

70

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

7�

3 Geographic Relationships of the Chilean Flora

Since the beginnings of plant geography, the relationships between floras have been analysed by classifying the taxa into floristic elements (Christ 1867, Engler 1882, Wangerin 1932, Good 1947, Takhtajan 1978). Wulff (1950) described five different approaches to define floristic elements, but recognized that “most investigators have been inclined to believe that the geographical factor is of primary importance and that the term element should be applied to it” (Wulff �9�0, p. �04, see also Wangerin 1932). In any case Wulff (1950) was the opinion that geographical [floristic] elements are fundamental for an understanding of a flora, and that an analysis of a flora should begin with these elements. More recently, floristic elements have been analysed for the North American flora north of México (Qian 1999), the East Asian flora (Qian et al. 2003), the flora of the Bolivian Andean valleys (López �00�), and the Ecuadorian superpáramo (Sklenar & Balslev �007). In the case of the Chilean flora, the first attempt is the one of Reiche (1907). He was able to identify seven elements or Kontingente (table �.�). Later, Villagrán & Hinojosa (�997) applied a similar classification to the woody genera of the Chilean temperate forests. These authors described eight floristic elements, including fossil taxa (table 3.1).

Table �.� Floristic elements in the Chilean Flora

Reiche �907 (Kontingente) Villagrán & Hinojosa �997

� Tropical-American Neotropical (amplio / disyunto)

� Andes Chile/Argentina Endémico bosques subantárticos / Endémico de Chile

� California/Mexican -

4 Antarctic Australasiano (cálido/antártico)

� Boreal (Europa, S Chile) Amplio

� Pantropic Pantropical

7 Introduced flora -

The present work is the second attempt, after Reiche (�907), to analyse the geographical relationships of the whole Chilean vascular flora at the genus level. Comparing the global distribution of the genera, seven floristic elements were distinguished, plus several generalized tracks that represent wide disjunct distributions. Generalized tracks have been integrated into biogeographical analysis by Croizat (�9��, �9�8)�8, and have been successfully used for the analysis and interpretation of disjunct distribution patterns (e.g. Katinas et al. �999 for the assessment of the Andean biota; Crisp et al. 1999 for the biogeographic analysis of the Australian flora; Katinas et al. 2004 for the analysis of North American Onagraceae). Recently Mihoc et al. (�00�) applied track analysis for assesing the evolution of the Andean genus Adesmia (series Microphyllae). Floristic elements have been usually recognized at the genus level (e.g. Good �947), and this approach will be followed here. The global distribution for each genus has been obtained from

7�

Wielgorskaya (�99�) and Mabberley (�997). For complementing these global accounts, new available monographs and systematic papers have also been consulted to get an appropriate image of the global distribution of the genera. As resumed in section �.�, by current knowledge, the Chilean flora is composed of 813 genera. After analysing the distribution of each genus, 7 floristic elements were identified, plus nine generalized tracks that represent wide disjunct distributions, (see table 3.2, and figures 3.1 to 3.7. Tracks are further represented in figure 4.2 and both elements and tracks are summarised in appendix A.

Table �.� Floristic elements for the vascular plant Chilean genera (update 07 February �007)

Floristic elements N° Genera %�. Pantropical 88 ���. Australasiatic �9 7

�. Neotropical ��� �7

4. Antitropical ��� �9�. South-temperate 8� �0

�. Endemic 8� �0

7. Cosmopolitan ��4 ��

Total 8���00 %

Table �.� Generalized tracks for the vascular plant Chilean genera (update 07 February �007)

Tracks (follow subsections in the text) N° of genera�.�.� Austral-antarctic track ��3.2.2 South Pacific tropical track ���.�.� Circum-austral track �

�.�.� Wide Neotropical track �4�.�.� Andean track ����.�.� South Amazonian track �9

�.4.� Wide antitropical track 843.4.2 Antitropical disjunct Pacific track ��3.4.3 Circum-Pacific track 9

3.1 Pantropical floristic elementGenera grouped in this element grow mainly in tropical regions (pantropics), sometimes extending into subtropical and/or temperate areas.This element includes 88 genera. The families with the greatest number of genera are Asteraceae (8 genera) and Poaceae and Fabaceae (� genera both). The most species-rich genera are Dioscorea, Conyza, and Calamagrostis. Few genera occur in the whole country (Calamagrostis, Conyza), or are restricted to northern Chile (e.g. Notholaena, Spilanthes, Gomphrena). Most are found in central-south Chile, disjunct to the rest of their distribution (e.g. Cryptocarya, Cissus, Glinus,

7�

Passiflora, Dodonaea, Dennstaedtia, Sigesbeckia, Pelletiera, Pouteria, Beilschmiedia, Wolffiella, Mikania, Hedyotis, Podocarpus).

Note: Megalastrum and Lippia are included in this element despite being restricted to the American and African tropics. Alonsoa was included here, too, even though the majority of its species are found in the Neotropics, because two of its species occur in South Africa.

Podocarpus (Podocarpaceae), sensu Bader �9�0 Acacia (Fabaceae)

Figure 3.1 Examples of the pantropical floristic element

Lobelia (Campanulaceae), sensu Meusel & Jäger �99� Nicotiana (Solanaceae), sensu van Steenis & van Balgooy �9��

Bacopa (Gratiolaceae), sensu Dawson �9�8 Cotula (Asteraceae), Meusel & Jäger �99�, Bremer �994

3.2 Australasiatic floristic elementThis element comprises genera from the South Pacific, i.e. Australasia as well as South America, and the Pacific islands (figure 3.2). Some genera are restricted to temperate subantarctic latitudes, others extend their distribution into tropical regions in South-East Asia or in South America. Webster (�99�) drew the distribution limits for several australasiatic genera in Central America (figure 4.3).The australasiatic floristic element comprises 59 genera. The families with the greatest number of genera are Asteraceae (4) and Poaceae, Apiaceae, Cunoniaceae, and Proteaceae, all with � genera. Within these genera, three generalized tracks based on superimposing distributions have been identified.

74

Azorella distribution, sensu Baumann �988 Berberidopsis distribution, sensu Ronse De Craene �004

Rytidosperma distribution, sensu Linder & Barker �000

Oreomyrrhis distribution, sensu Mathias & Constance �9��

Hebe distribution, sensu van Steenis & van Balgooy�9��Coprosma distribution, sensu van Steenis & van Balgooy �9��

Jovellana distribution, sensu Dawson �9�8, Heads �994

Ourisia distribution, sensu Meudt & Simpson �00�

Leptinella distribution, sensu Bremer �994

Lagenophora distribution, sensu Cabrera �9��

Figure 3.2 Examples of the australasiatic floristic element

7�

3.2.1 Austral-antarctic track

This track comprises genera occurring in southern South America, New Zealand, Eastern Australia and Tasmania. It was previously described as South Pacific track by Crisp et al. (�999). It numbers �� genera, e.g. Luzuriaga, Eucryphia, Prumnopitys, Jovellana, Pseudopanax. Most of these genera are restricted to the southern temperate forests, but some of them reach northernmost Chile (e.g. Hypsela, Azorella, Cortaderia) and some of them even reaching Ecuador. Some genera reach Central Chile, disappear in the Atacama and reappear in the northern Andes (e.g. Fuchsia, Lomatia, Prumnopitys). Haloragis is an austral genus that reaches the Juan Fernández islands, but not the continent.

3.2.2 South Pacific tropical track

This track comprises genera occurring in southern South America, New Zealand, Eastern Australia and/or Tasmania, some also extending to New Guinea, Malesia, and even to East Asia. It numbers �� genera, e.g. Araucaria, Nothofagus, Lagenophora, Abrotanella, Coprosma, and Hebe. Many genera are restricted to the southern temperate forests, but a few reach northernmost Chile (e.g. Colobanthus, Muehlenbeckia, Pratia). Some reach central Chile, disappear in the Atacama and reappear in the northern Andes (e.g. Citronella, Uncinia, Oreobolus). Doodia is widespread in the Pacific till Isla de Pascua, not in continental Chile. Dicksonia, Arthropteris, Coprosma, and Santalum are austral genera that are represented in the Juan Fernández Islands, also absent from the continent�9.

3.2.3 Circum-austral track

This track comprises genera occurring in southern South America, Australasia, and extending further into the Indic Ocean, reaching Madagascar or South Africa. These are only � genera, Nertera, Rumohra, and Weinmannia. This latter genus disappears in northern Chile and its maximal species-richness is found between the northern Andes and Central America.

3.3 Neotropical (American) floristic elementStrictly speaking, this is an American element, but neotropical has been used for a long time in the phytogeographical literature. This element includes ��� genera, mostly from the Asteraceae (��), followed by Solanaceae (��) and Poaceae (��), Cactaceae (��), Brassicaceae (7) and Apiaceae (7). Three generalized tracks have been identified within this element:

3.3.1 Wide Neotropical track

This track comprises genera from South America, extending to Mexico, to south-western USA or even to southern Canada. The main massing however, lies in the intertropics. Many species also occur in Brazil.

7�

Figures 3.3 Examples of the neotropical floristic element (distribution from monographs and checklists)

?

Alstroemeria Azara Calceolaria

Cremolobus Eremodraba Escallonia

Eudema Malesherbia Mathewsia

Mecardonia Nototriche Vasconcellea

77

The wide Neotropical track comprises �4 genera, e.g. Haplopappus, Calceolaria, Baccharis, Drimys, Nassella. Some are found in all of Chile (e.g. Baccharis, Gamochaeta, Calceolaria, Calandrinia), others just in northern Chile (e.g. Coreopsis, Tara, Bouteloua), but most genera occur in central to south-temperate Chile, disjunct from the core neotropical distribution (e.g. Piptochaetium, Stenandrium, Chusquea, Calydorea, Chaptalia, Ugni).

3.3.2 Andean track

This track comprises genera ranging from southern Chile to Colombia (and Costa Rica), but not reaching North America. It numbers ��� genera, e.g. Chuquiraga, Nototriche, Polylepis, Geoffroea. Some are found in all of Chile (e.g. Perezia, Escallonia); others only in northernmost (Dunalia, Stenomesson) or north-Central Chile (Exodeconus, Equinospsis, Lycopersicon); many distributions are disjunct between southern Chile/Argentina and northern South America (e.g. Llagunoa, Myrcianthes, Dysopsis, Blepharocalyx, Desfontainia, Myrteola). More than 80 genera have a main massing in the central Andes of Perú, Bolivia, northern Argentina and Chile. Most of them occur in a continuous range from northern Chile to Peru/Bolivia (e.g. Neowerdermannia, Oreocereus, Tunilla, Pycnophyllum, Philibertia, Balbisia, Acantholippia), others occur disjunctly between central Chile/Perú (Tetraglochin, Weberbauera, Kageneckia, Eccremocarpus, Lucilia). �9 genera are restricted to central/northern Chile and adjacent Argentina (e.g. Urbania, Urmenetea, Werdemannia, Lenzia, Kurzamra).

3.3.3 South-Amazonian track

This track comprises genera found in Chile, northern Argentina, Uruguay, Paraguay, and south-eastern BrazilIt numbers �9 genera, e.g. Colliguaja, Quillaja, Azara, Tweedia, Myrceugenia, Viviania, and Dasyphyllum. Most of these genera occur disjunctly between Central Chile and SE Brazil.

3.4 Antitropical floristic elementThis element comprises genera found both in the northern and in the southern temperate regions, but which are absent from the intervening tropics. This pattern is commonly referred to as the amphi-tropical pattern in the literature, but as W. Welß (pers. comm.) correctly noted, the most appropriate term would be antitropic, since amphitropical means both tropics. In Cox’s (�990) opinion, the most appropriate term to be used would be amphitemperate, but since it includes subtropical distributions, the most suited term is antitropical (also used by Glasby �00�, and Parenti �007).

This element includes ��� genera. The families with the greatest number of genera are Asteraceae (��), Poaceae (�0), Fabaceae (8), Polemoniaceae (7), and Boraginaceae (7).

78

Figures 3.4 Examples of the antitropical floristic element

E Asia

Hydrangea distribution, sensu Mabberley �997 Antennaria distribution, sensu Meusel & Jäger �99�

Larrea, sensu Hunziker et al. �97�

Mancoa, sensu Al-Shehbaz comm. pers.

Euphrasia distribution, sensu Van Steenis �9��, Dawson �9�8Artemisia distribution, sensu Meusel & Jäger �99�

Osmorhiza, sensu Wood �97�

Tropidocarpum, sensu Al-Shehbaz �00� Agoseris, sensu Wood �97�

Frankenia, sensu Wahlen �987

79

One of the naturalists to refer to this element was G. Treviranus (�80�), even though he did not recognise it formally. While discussing the components of his Antarktische Flor, he noticed that there was a floristic relationship between southern South America and New Zealand, and that some of this genera have their main distribution in the northern and southern temperate zones (e.g. Pinguicula, Salix, Ribes) (Treviranus (�80�, pp. ���-���). Du Rietz (�940) and others called it the pattern of bipolar plant distribution. The antitropical element was treated in depth in several papers that arose from a symposium at the beginning of the �9�0s (Constance �9��). The antitropical element includes many variable distributions patterns, some restricted, others very wide. Within this element three generalized tracks were identified.

Note: Menodora (Oleaceae) is considered as an antitropical genus, since its distribution range includes both subtropical Americas, and subtropical South Africa (this genus was not included in the track).

3.4.1 Wide antitropical track (bipolar-temperate element)

This track comprises genera found in Eurasia, North America, southern South America, some also ranging into the montane America tropics. It numbers 84 genera, e.g. Astragalus, Fagonia, Valeriana, Vicia. Many of these genera are found in all of Chile (e.g. Hypochaeris, Cystopteris, Bromus, Valeriana). Some are restricted to northern Chile (e.g. Alchemilla, Woodsia), while others are restricted to southern Chile (e.g. Rhamnus, Fragaria, Adenocaulon, Saxifraga), some of them even to the southernmost Magallanes region (Botrychium, Chrysosplenium, Androsace).

3.4.2 Antitropical disjunct Pacific track

All the genera grouped in this track have a disjunct distribution between south-western North America and South America, occurring mostly in subtropical and tropical deserts.This track numbers �� genera, e.g. Agalinis, Hoffmannseggia, Camissonia, Tiquilia, and many Polemoniaceae (Gilia, Microsteris, Ipomopsis, Linanthus). Few genera are found along the whole latitudinal gradient in Chile (e.g. Phacelia). Some occur in northern Chile and the Central Andes (e.g. Mancoa, Cistanthe, Tarasa), but many are found only in Central Chile and south-western North America (e.g. Navarretia, Blennosperma, Plectritis, Linanthus, Tropidocarpum, Lastarriaea, Errazurizia).

3.4.3 Circum-Pacific track

This track comprises genera with a disjunct distribution in North America and South America, that are also represented in Australasia.It numbers 9 genera: Lilaeopsis, Distichlis, Flaveria, Sicyos, Microseris, Soliva, Plagiobothrys, Gochnatia, and Gaultheria.

Note: Two mainly circum-Pacific genera reach South Africa: Acaena and Carpobrotus. Therefore

these two genera were not included in the circum-Pacific track.

80

Figure 3.5 South-temperate floristic element ((distribution from monographs and checklists)

Austrocedrus Embothrium Fonkia

Grammosperma Hamadryas Laureliopsis

Melosperma Menonvillea Monttea

Onuris Pilgerodendron Saxe-gothaea

8�

3.5 South-temperate floristic element This element comprises genera only found in central/southern Chile and in adjacent Argentina.This element includes 8� genera. The families with the greatest number of genera are Asteraceae (9), Apiaceae (�), and the Brassicaceae (4). Most genera are restricted to the southern temperate forests (e.g. Laureliopsis, Pilgerodendron, Fitzroya, Drapetes, Embothrium, Boquila) and Patagonia (Lecanophora, Grammosperma, Eriachaenium, Saxifragella, Xerodraba, Lepidophyllum, Chiliophyllum). Some are represented in the temperate zone but reaching more arid environments to the north, like Triptilion, Nasthantus, Trichopetalum. The genus Mulinum reaches the northest latitude at ��°S. The south-temperate element will be further discussed in section 4.4.

3.6 Endemic floristic elementThis element comprises genera endemic to continental Chile, and to the Chilean Pacific islands.It numbers 8� genera, of which �7 genera are endemic to continental Chile (e.g. Oxyphyllum, Bridgesia, Leontochir, Balsamocarpon, Trevoa, Jubaea), while �� genera are endemic to the Chilean Pacific islands, especially Juan Fernández (e.g. Dendroseris, Lactoris, Cuminia, Juania). The endemic element will be further discussed in sections 4.�, 4.4, and 4.�.

Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon

Figure 3.6 Endemic floristic element (collections from SGO)

8�

Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa

Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus

Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus

8�

Huidobria Hymenoglossum Ivania

JF

Jubaea Lapageria Latua

Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme

Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera

JF

84

Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea

Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra

Placea

Figure 3.6 Endemic floristic element (continues)

Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia

JF

8�

3.7 Cosmopolitan floristic elementThere are not many genera that can be considered as really cosmopolitan (Good �974), e.g. some semi-aquatic plants (Sagittaria, Landoltia, Wolffia), some ferns and fern allies (Isoetes, Lycopodium, Huperzia, Adiantum) or some really widespread terrestrial angiosperm genera like Senecio, Rubus, Cyperus, Ceratophyllum, Gnaphalium, Ranunculus, Geum, Juncus or Scirpus. Genera included in this element have a wide distribution in more than two continents and more than two principal climatic zones (e.g. tropical and temperate). In fact, the element should be called subcosmopolitan, but cosmopolitan is more often used.

It numbers ��4 genera, some of them occupying the whole country (e.g. Asplenium, Gnaphalium, Ranunculus, Chenopodium, Silene), some being restricted to the north (e.g. Salvia, Chloris, Portulaca), and most being found in central and southern Chile (Ceratophyllum, Samolus, Aphanes, Geum, Alisma, Glyceria, Panicum, Hypericum). A few genera are restricted to the Magallanes region (Huperzia, Landoltia).

Figure 3.7 Cosmopolitan floristic element

Apium (Apiaceae), sensu Meusel et al. �978 Bidens (Asteraceae), sensu Meusel & Jäger �99�

Eryngium (Apiaceae), sensu Meusel et al. �978 Hydrocotyle (Apiaceae), sensu Meusel et al. �978

Lycium (Solanaceae), sensu Meusel et al. �978 Mimulus (Phrymaceae), modified from Meusel et al. 1978

8�

87

4 Biogeographic analysis

88

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

89

4 Biogeographic analysis

The seven floristic elements detected contain genera which schow a diversity of geographic range sizes in Chile. In every element, there are some genera distributed in the whole country, from the southernmost Cabo de Hornos to the northern Parinacota province, with exception of the endemic element. Every element also contains a number of genera known for a very restricted range or known only from a couple of localities. A method of assessing the main massing of the genera is to calculate for every element the average distribution of the genera that compose them [northern limit + southern limit / �] (see appendix A for the latitudinal limit of each genus) Results are shown in table 4.1 and figure 4.1.

Table 4.1 Average of geographic range of floristic elements

floristic element Average (°S)

1. Pantropical -��,�

2. Australasiatic -4�,�

3. Neotropical -�9,�

4. Antitropical -��,�

5. South-temperate -4�,�

6. Endemic -��,�

7. Cosmopolitan -��,0

The neotropical element has the northernmost average (�9,�°S), while the south-temperate element has the southernmost average (4�,�°S). The australasiatic element has an average at 4�,�°S. The pantropical, antitropical, cosmopolitan and endemic elements show an average latitude between ��°S and ��°S. The fact that most elements have their average in Central Chile tends to reinforce the early view of Grisebach (�87�), Engler (�88�), and Schmithüsen (�9��) to consider this region as a transition zone with different converging elements.

After grouping the Chilean plants in seven floristic elements (Kontingente), Reiche (�907, p. ��9) proposed that the neotropical and antitropical (Reiche’s Californian) elements occur in Chile due to a migration route (Wanderungslinie) from the north to the south along the Andes down to Magallanes, until this migration was stopped at the border of the Antarctic realm (situated sensu Reiche from ~ 40° to the south along the coast). On other hand, the Antarctic element shows sensu Reiche a migration route from the south to the north, mostly

54°

48°

42°

36°

30°

24°

18°

south-temperate

neotropical

antitropical

pantropical/

cosmopolitan

endemic

australasiatic

Figure 4.1 Average range for floristic elements

90

along the coast, up to the Maule river (~ ��°). Reiche illustrated these migration routes in his map for the Compositae (Reiche 1905) (figure 2.12). Now, Reiche’s view can be complemented by analysing the disjunct distribution patterns in all the floristic elements detected.

4.1. To be or not to be disjunct Villagrán & Hinojosa (1997) classified some of the temperate woody genera in an elemento neotropical disyunto. These authors suggest that the disjunct distribution is the result of the former existence of a continuous subtropical forest from SE Brazil to central/South Chile during Oligocene/Miocene times (see chapter �).

Disjunction distribution indeed includes many taxa and is the rule rather than the exception in the Chilean flora. 152 genera compose the antitropical (disjunct) floristic element, 59 genera compose the australasiatic (disjunct) floristic element. Many of the pantropical (e.g. Passiflora, Cryptocarya, Cissus, Podocarpus) and (sub)cosmopolitan genera (e.g. Sanicula, Coriaria) also show a disjunct distribution between the Chilean and the global distribution range. At the regional scale, the neotropical and the south-temperate elements also comprise disjunct distribution ranges (e.g. Escallonia, Matthewsia, Azara, Alstroemeria for the neotropical; Monttea and Werdermannia for the south-temperate element).

The different tracks that account for the major disjunctions in the Chilean vascular flora are presented in figure 4.2.

Austral-antarctic track (31) Wide Neotropical track (64)

Pacific antitropical track (56)Panbiogeographic nodes

Circum-austral track (3)

Andean (113) and South-Amazonian (39) tracks

Wide Antitropical track (84)

Circum-Pacific track (9)

South Pacific tropical track (25)

Figure 4.� Generalized tracks for the Chilean plant disjunct ditributions (in parenthesis the number of genera composing the tracks)

9�

As noted by Moore et al. (�00�), researchers have debated the causes of the antitropical disjunctions for over a century (e.g. Bray �900; Johnston �940; Constance �9��; Raven & Axelrod �974; Hunziker et al. �97�; Solbrig �97�a,b; Carlquist �98�). Moore et al. (�00�) mention three types of hypotheses to account for the disjunction:

a) migratory hypothesis, whereby ancestral populations dispersed over short distances through the tropics across arid or semi-arid ‘‘islands’’ (stepping-stone migration) or migrated directly through an ancient, continuous arid to semi-arid tropical corridor along the Pacific coast of the Americas (Raven 1963, Solbrig 1972, Williams 1975)

b) parallel evolution of near-identical arid-adapted taxa from widely distributed tropical ancestors (Johnston �940, Barbour �9�9).

c) long-distance dispersal, probably by migrating birds; from the �9�0s on, many researchers have favoured this explanation as the main cause for the disjunctions (Raven �9��, �97�; Cruden �9��; Carlquist �98�; Simpson & Neff �98�).

The commonly preferred scenario of long-distance dispersal (LDD) which was developed since the �9�0s is well described by Raven (�9��, p. ���): “In evaluating the probabilities of LDD between North and South America it is important to note that most of the distribution pattern corresponds closely with migration routes of birds. Furthermore, an unduly high proportion of the plants involved grow in open communities such as those of seacoasts or seasonally moist places frequented by migratory birds. Small seeds occasionally adhere to birds and exceptionally may not fall off until the bird has reached a favourable habitat on the other side of the tropics. Considering the millions of birds that fly between temperate North and South America every year, some transport might happen at least at the rate postulated for the colonization of Hawaii which lies on no known migration route”.

Moore et al. (2006) further classified antitropical disjunct plants of North and South America into two general phylogenetic classes:

a) identical or closely related, disjunct species or species pairs. The morphological similarity exhibited within these species-level disjuncts suggests that they arose from very recent dispersal events. Molecular analyses have confirmed the phylogenetic closeness and likely recent trans-tropical colonization in several groups (see section �.�).

b) disjunct plant groups with multiple species endemic to each continent. Most of these groups occur in the more arid regions of the Americas: e.g. Ephedra, Hoffmannseggia, Tiquilia. In Moore et al.’s (�00�) opinion, these disjunct groups could provide evidence both of earlier dispersal events, and of multiple dispersal events.

Moore et al. (�00�) ruled out the possibility of ancient vicariance events. These alternative hypotheses are categorized as “grotesque hypotheses that have been proposed to avoid the bugaboo of long-distance dispersal” (Raven �9��, p. ���). But if grotesque is better than bugaboo is just a matter of belief and not a matter of facts… (see further discussion in section 6.1).

9�

4.2 The austral v/s the neotropical floristic realmAs mentioned in section 2.3, the Chilean flora has been alternatively classified under the neotropical and/or the austral floristic realm. We can summarize four views that are in conflict:

a) the older view of Engler (�88�), Diels (�908) and Skottsberg (�9��), which draw a boundary between the neotropical and the austral floristic realms at around 47°S (figure 2.17); b) Takhtajan’s (1978) newest view setting this line at around 23°S (figure 2.10); c) the modern proposal of Cox (2001) that aligns all of the South American flora under an American kingdom; d) and recent Morrone’s (2002) classification that situates the Chilean biota plus the whole Andes in an Austral kingdom.

The latitudinal range of several genera supports each of these views: a) the boundary at ~47°S is shown by several genera that do not overpass this limit to the north, like Tetrachondra, Grammosperma, Hebe, Oreomyrrhis, or Rostkovia. b) the boundary at ��° poposed by Takhtajan (�978) is shown by most (4�) of the genera that compose the australasiatic element. The Fray Jorge fog forest, located at latitude �0°, has long been recognized as the northern outpost of some of the subantarctic elements (Muñoz Pizarro & Pisano �947, Trocoso et al. �980, Squeo et al. �004). Some kilometres north of Fray Jorge, in Pichasca, the northernmost fossil occurrence of the emblematic subantarctic genus Nothofagus has been documented (Torres & Rallo �98�). c) Cox’s (2001) vision neglects or minimizes the australasiatic element in the American flora and therefore do not leave opportunity for comparisons. d) In defense of Morrone’s (�00�) view, many australasiatic genera reach northern Chile and further the northern Andes till Ecuador or Colombia, like Colobanthus, Hypsela, Azorella, Ourisia, Trichocline, Cortaderia, Muehlenbeckia. Some of them reaching even Central America/Mexico (e.g. Weinmannia) (figure 4.3). Furthermore, several genera are restricted to temperate Chile and are absent in the northern part, but they reappear in the northern Andes, like Rumohra, Citronella, Prumnopitys, Nertera; some of them even reaching their highest level of species richness in the northern Andes: e.g. Fuchsia, Weinmannia, and Uncinia.

So the question remains: which of the classifications fit better to the Chilean flora? With the analysis of the floristic elements in the latitudinal profile we can attempt to answer the question. Since the latitudinal

100° 90° 80°

10°

20°

30°

Podocarpus

WeinmanniaDrimys/Centropogon

Gunnera

Desfontainia/Escalllonia

Figure 4.� Northward extent of spread of Gondwana genera in tropical America (redrawn from Webster �99�)

9�

distribution of the genera, as represented in figures �.�9 to �.��, does not account for possible distribution or collection gaps, four regional floras were taken for a similarity analysis: Antofagasta (ANT), Coquimbo (COQ), Biobío (BIO), and Magallanes (MAG) (figure 4.4). Three of the regions included in the anaylsis have a published checklist (table 4.�). The checklist for BIO was gently provided by Dr. R. Rodríguez from CONC herbarium. Synonym taxa were homologized (e.g. Lagenophora=Lagenifera, sensu Cabrera �9��) . Regions analysed are dissimilar in area, but some of them harbour similar numbers of native genera (e.g. COQ=457, BIO=465), MAG showing the lowest generic richness: MAG=252.

Jaccard similarity index was applied to the data set (Zunino & Zullini �00�, Cox & Moore �00�, Lomolino et al. �00�) (appendix B). The highest floristic similarity is between COQ and BIO, that share 323 genera (table 4.3). The lowest similarity is shown by ANT and MAG, which share 97 genera. There seems to be a relationship between the similarity and the geographic distance, and in figure 4.5 both variables are represented, showing this trend of increasing similarity between nearest regions.

Table 4.2 Regions for floristic similarity analysis (see also appendix B)

Abbreviation Regions Area (km�) N° native vascular plant genera

Source

ANT Antofagasta ���.049 ��� Marticorena et al. �998a

COQ Coquimbo 40.�80 4�7 Marticorena et al. �00�BIO Biobío �7.0�� 4�� CONC, R. Rodríguez pers. comm.

MAG Magallanes ���.0�� ��� Henríquez et al. �99�

Table 4.� Floristic similarity and geographic distance different regions in Chile: Antofagasta (ANT), Biobío (BIO), Coquimbo (COQ) and Magallanes (MAG). Geographic distance have been calculated as the latitudinal difference between the geographic centroid of each region.

Compared regions Shared generaSimilarity Jaccard Distance (km)

ANT/COQ �44 0,4� 777ANT/BIO �74 0,�9 ���7ANT/MAG 97 0,�� ���7COQ/BIO ��� 0,�4 7�0COQ/MAG ��0 0,�7 �4�0BIO/MAG �78 0,�� �700

Isla de Pascua

Juan Fernández

Islas Desventuradas

BIO

COQ

ANT

MAG

Figure 4.4 Chilean continental regions included in the similarity analysis and location of Chilean Pacific islands.

94

The analysis of the floristic elements in the compared regions indicates clear trends along the latitudinal gradient in Chile: the cosmopolitan, antitropical, south-temperate, and australasiatic genera show a relative increase towards the south, while the proportion of neotropical, pantropical and endemic genera decrease towards the south (table 4.4, figure 4.6). Highly remarkable is the exchange of the neotropical and the australasiatic genera between BIO and MAG (arrow in figure 4.�).

Table 4.4 Floristic elements in each region

REG pantropical australasiatic neotropical antitropicalsouth-

temperate endemic cosmopolitan

n % n % n % n % n % n % n %ANT �8 8,89 8 �,�4 ��� ��,�� 4� �4,�9 �7 �,40 �� 4,7� 8� ��,0�COQ �9 8,�9 �� 4,8� ��4 ��,�� 7� ��,08 4� 9,�� �8 8,�7 ��7 ��,77BIO 4� 9,09 4� 8,87 �00 ��,�� 7� ��,�8 �� ��,04 �7 �,84 ��� ��,��MAG �� 4,�8 �8 ��,�4 �9 ��,�� 44 �7,�� 4� ��,7� � 0,40 84 ��,47

0 2000 4000 6000 8000 10000

0.0

0.2

0.4

0.6

0.8

1.0

Distance [km]

Jacc

ard

sim

ilarit

y

ANT/COQ

ANT/BIO

ANT/MAG

COQ/BIO

COQ/MAGBIO/MAG

Figure 4.� Similarity versus geographic distance between Chilean regions.

0,00

10,00

20,00

30,00

40,00

ANT COQ BIO MAG

Regions

%

1. Pantropical2. Australasiatic3. Neotropical4. Antitropical5. South-temperate6. Endemic7. Cosmopolitan

Figure 4.6 Floristic elements present in four regional floras (percentage)

9�

Therefore and in spite of the relative low amount of genera strictly restricted to southernmost Chile, the replacement of the neotropical floristic element by the autralasiatic element in MAG, suggests the consistency of classifying subantarctic Chile south of 47° in an austral floristic realm as earlier proposed by Engler (�88�), Drude (�884), Reiche (�907, p. �8�), Diels (�908), and Skottsberg (�9��).

4.3 Analysis of Endemism Endemism, since Augustin P. de Candolle (�8�0) coined the term, has turned out to be one of the most appealing concepts in historical biogeography, and more recently in conservation biology. Endemic taxa are restricted to a specific area, and found nowhere else. The concept is scale-dependant, i.e. taxa can be endemic to a single lagoon, to a region, or to a continent. Endemism also depends on taxonomy: taxonomical categories are hierarchical. Therefore, lower taxonomic categories, species and genera, tend to show in general a higher level of endemism than higher taxa, such as families and orders (Lomolino et al. �00�). Taxa can be endemic to a location due to three different reasons: (�) they originated in that place and never dispersed; (�) their entire range has shifted in locality after the origin; or (�) they survive in only a small part of their former range.

A geographic area that contains two or more non-related endemic taxa is formally defined as an area of endemism, a concept of vital importance in modern historical biogeography (Harold & Mooi �994, Linder �00�). In the words of Nelson & Platnick (�98�) “the most elementary questions of historical biogeography concern areas of endemism and their relationships”. Hereafter, the patterns of endemism and its hierarchical arrangement in the Chilean continental vascular flora will be analysed. Since the country has mainly natural borders, i.e. the Andes and the Pacific, it can be interpreted as a natural unit for the analysis of endemism. Reports on endemism at the genus level have been partially published by Muñoz-Schick & Moreira-Muñoz (�000).

0

10

20

30

40

50

1 2 3 4 5 6 7 8 9 10+

N° species per genus

N°

ende

mic

gene

ra

Fig. 4.7 N° of endemic genera v/s N° of species per genus

9�

Fig. 4.8 Latitudinal ranges for the �7 Chi-lean endemic genera. Small distribution ranges on top, large towards the buttom.

-57 -47 -37 -27 -17

Neuontobothrys

Metharme

Ivania

Leontochir

Marticorenia

Avellanita

Valdivia

Reicheella

Oxyphyllum

Gypothamnium

Pintoa

Balsamocarpon

Miqueliopuntia

Dinemagonum

Araeoandra

Guynesomia

Bridgesia

Pleocarphus

Tecophilaea

Leunisia

Gethyum

Placea

Traubia

Speea

Jubaea

Calopappus

Tetilla

Cissarobryon

Legrandia

Pitavia

Gomortega

Desmaria

Hollermayera

Latua

Dinemandra

Zephyra

Copiapoa

Bakerolimon

Phrodus

Cyphocarpus

Homalocarpus

Moscharia

Trevoa

Adenopeltis

Podanthus

Scyphanthus

Ochagavia

Miersia

Acrisione

Gymnachne

Vestia

Fascicularia

Leptocarpha

Huidobria

Alona

Conanthera

Anisomeria

Leucocoryne

Microphyes

Epipetrum

Peumus

Lapageria

Sarmienta

Notanthera

Francoa

Ercilla

Hymenoglossum

97

4.3.1 Geography of Endemism

The �7 endemic continental Chilean genera (section �.�) belong to �� families (appendix A). In continental Chile, the family with the highest number of endemic genera is Asteraceae (��). The Alliaceae are represented by 4 endemic genera, and the Brassicaceae, Solanaceae and Tecophilaeaceae by � endemic genera. Noteworthy is the existence of two endemic families in the continental Chilean flora: the Gomortegaceae and the Francoaceae, with one genus (Gomortega) and two genera (Francoa and Tetilla), respectively (table �.�).

The most species-rich genus is Copiapoa (Cactaceae), which contains �� endemic species, followed by Leucocoryne (��, Alliaceae), Homalocarpus (Apiaceae), and Alona (Solanaceae), both with six species. As seen in figure 4.7, most of the Chilean endemic genera are monotypic, i.e. are composed of only one species (4� genera).

The distribution ranges of the endemic genera were grouped in four categories: small (within one latitudinal degree), medium-small (� to � latitudinal degrees), medium-large (� to �0 latitudinal degrees) and large (more than �0 latitudinal degrees). Of the �7 endemic genera analysed in this study, � (8,9% of the total) belong to the smallest category, i.e. are found only within � degree of latitude (figure 4.8). These are the genera Avellanita, Ivania, Leontochir, Metharme, Neuontobothrys, and Valdivia. �8 genera (4�,8%) occur in medium-small ranges of distribution between � and � latitudinal degrees, e.g. Leunisia, Tetilla, Jubaea. �0 genera (�9,9%) feature medium-large ranges of distribution between � and �0 latitudinal degrees, e.g. Copiapoa, Miersia, Vestia. �� genera (�9,4%) feature large ranges of distribution wider than �0 latitudinal degrees,

e.g. Huidobria, Hymenoglossum, Lapageria. The endemic genus with the widest latitudinal distribution in Chile is Leucocoryne with a distribution range of �9 latitudinal degrees from ��° to 40° S.

The analysis of the generic richness indicates a latitudinal trend, with the greatest number of endemic genera (��) ranging between ��°-�4° S (figure 4.9) The number of endemic genera decreases constantly to the north and to the south of this central area.

Besides the latitudinal pattern of geographic ranges, there is also a longitudinal pattern. This is especially pronounced in Chile with its high environmental contrast between the low coastal areas and the Andes mountains. Drawing the

0 10 20 30 40

55-56

53-54

51-52

49-50

47-48

45-46

43-44

41-42

39-40

37-38

35-36

33-34

31-32

29-30

27-28

25-26

23-24

21-22

19-20

17-18

Latit

ude

(°S

)

N° genera

Fig. 4.9 Endemic genera at each latitude degree.

98

distribution range for every genus gives a good idea of the individual pattern of each endemic genus, but does not allow further analysis (figure 4.10). The division of Chile into equal-area cells and the application of Parsimony Analysis of Endemism (PAE) can provide additional results.

4.3.2 Parsimony Analysis of Endemicity (PAE)

Parsimony Analysis of Endemicity (PAE) is a biogeographical method that aims to classify areas by the most parsimonious solution based on the shared presence of taxa (Rosen �988, Morrone �994). Analogous to cladistic analysis, PAE treats areas as taxa, and taxa as characters. Although taxonomic information is not derived from their clades, it does, however, generate area clades, or patterns that may be regarded as hypotheses of area genealogy (Rosen �99�, as quoted by Nihei �00�). In cladistic analysis, characters are represented by characteristics that are uniquely shared by a set of taxa (=synapomorphies). In PAE, however, information concerning an area hierarchy (the grouping of areas) is based on synendemic taxa (Rosen �99�), i.e. endemic taxa that are found

Figure 4.�� Generic richness for each ��� x ��� km cell

Figure 4.�0 Geographic ranges of endemic Chile-an genera

Fig. 4.�� Grid (��� x ��� km) for the PAE application in continental Chile

99

in more than one area (Nihei �00�). The geographical distributions of taxa are combined into a presence/absence matrix. After application of a standard maximum parsimony analysis, area cladograms of minimal length (number of steps) are derived. Areas grouped together in these area cladograms are interpreted as areas of endemism (i.e. areas between which biotic interchange has occurred). For the PAE-analyis Chile was divided into a 111 x 111 km grid, i.e. in 81 cells (figure 4.11). The cells correspond to 1° degree latitude and were modified in longitudinal direction to get equal area cells. The distribution data consists of collection localities from the SGO herbarium, Santiago, in geographic degrees (unprojected map). Presence/absence (�/0) for each genus was plotted for each cell, on the base of a join between the grid map and the data base of collection localities from SGO (cartographic examples showed in section �.�). The matrix of presence/absence for endemic genera in each cell is shown in appendix C. An overview for the distribution of the genera in the grid space is presented in figure 4.12. The highest genus-richness lies at 33°-34° (coast), and decreases to the north and to the south.

For running a parsimony analysis of endemicity (PAE), a hypothetical area with only absences has been used to root the tree. The data matrix was analysed using the TNT cladistic program. A typical search with �00 replicates resulted in 4�0 most parsimonious trees. A TNT search resulted in only five most parsimonious trees. Consensus trees were obtained using the majority-rule, with confidence values calculated at each node (figure 4.13 shows the consensus tree of the TNT search).

The consensus tree discriminate two major areas of accumulated endemism within a group of central Chilean coastal cells (31-38°S) = a central coastal floristic block; and a group of central Chilean Andean cells (33°-39°S) = a central Andean floristic block. A third group of southern cells including Chiloé Island (44°S) can be interpreted as a transition toward the temperate zone, that features fewer endemic genera. The cells from ��°S to the north are at a basal position in figure 4.13, and are therefore considered as having the lowest accumulated endemism. The two central floristic blocks can be interpreted as core area of endemism in Central Chile (figure 4.14). The central coastal floristic block is characterized by some exclusively distributed genera like Adenopeltis, Ochagavia, Miersia, and Pitavia. The central Andean floristic block is characterized by Calopappus, Legrandia, and Leunisia. Furthermore, both floristic blocks are characterized by Peumus, Notanthera, Francoa, and Ercilla. The transition zone from �9° to Chiloé is characterized by Latua, Hollermayera, and Desmaria. Northern endemic taxa outside of the core areas of endemism are Zephyra, Reicheella, Oxyphyllum, and Metharme (not shown in figure 4.14, but see figures 3.6 and 4.8).

Pliscoff (�00�) and Rovito et al. (�004) also found a core area of endemism in central Chile based on woody taxa and Chilean Senecio species respectively. An interesting topic for discussion are possible direct connections of the endemic patterns with current climatic or topographic factors

�00

Majority rule tree (from 5 trees, cut 50)

AManAMcoALanALcoAKanAKinAKcoAJanAJinAJcoAIanAIcoAHanAHcoAGan

AGco

AFanAFcoAEanAEco

ADanADinADco

ACan

ACco

ABan

ABcoAAan

AAco

Zan

ZcoYan

Yco

XanXco

Wan

WcoVan

Vco

Uan

Uco

Tan

Tco

San

Sco

Ran

Rco

Qan

Qco

Pan

Pco

Oan

Oco

Nan

Nco

Man

Mco

Lan

Lco

Kan

Kco

Jan

Jco

Ian

Ico

Han

Hco

GanGin

Gco

Fan

Fco

Ean

Eco

Dan

Dco

CanCcoBanBcoAan

Hypothetical_area

10080

8080

80100

100

100

80

100100

100100

100100

100

100

60

100100

100100

100100

80

6060

60100

100

60

60

60

100

Figure 4.�� Consensus tree from a TNT search in program TNT, recommended by the distributors of the program for more than 80 taxa (in this case cells). Settings as default. � more parsimoniuos trees were found. Terminals are the cells of figure 4.11, differentiated in coastal (co), interior (in) and Andean (an). Arrows show groups of cells as represented in figure 4.14.

�0�

(geodiversity) (Cowling et al. �99�, Mutke & Barthlott �00�). The central area of endemism lies within the Mediterranean-type climate zone of Central Chile, which is located between �7°-�7°S (Luebert & Pliscoff �00�). Both areas match quite closely, but a direct relationship between climate and endemism is dubious, since the current mediterranean-type climate in central Chile (at �4°S) seems to be very recent (Villa-Martínez et al. �00�). This suggests, for Chile as well as for the European Mediterranean region (Verdú et al. �00�), that the endemic genera are not directly related to the current climate, but rather to the Paleogene history of the biota (Hinojosa et al. �00�, and chapter �). Landrum (�98�) proposed that the high proportion of endemic genera present in central Chile is due to the long isolation of these forests, at least �0 to �0 mya.

Central Chile as a core area of endemism can be interpreted as a panbiogeographic node (sensu Heads �004) where there occur a superposition of genera belonging to different elements that have further evolved under isolation.

At the end of this section, some problems with the application of PAE as a valid method in

Fig 4.�4 Floristic blocks based on terminal cells from figure 4.13.

Central coastal floristic block

Central Andean floristic block

Southern floristic block

�0�

historical geography are worth mentioning. In the opinion of some authors, e. g. Santos (�00�), biogeographical patterns rely on phylogeny. Therefore PAE should not be considered a truly historical biogeographical method. Consequently, this author considers that PAE should “rest in peace”. The criticism of Santos (�00�) is valid in the sense that one has to be aware of the limitations and assumptions inherent to the methods�0. But in a recent revision, Nihei (�00�) suggests that one should differentiate between dynamic and static approaches of PAE, and that with good knowledge of the methods one should be able to choose the right method for each given problem (see also Morrone �00�).

Some cladistists are extremely skeptic about the use of parsimony in biogeography (Goloboff pers. comm.) Basic concepts of cladistics, like synapomorphies and apomorphies, are still not well resolved in PAE. Therefore, specific methods for discovering and analysing areas of endemism are currently under development (Szumik et al. �00�, Szumik & Goloboff �004, Domínguez et al. �00�, see a comparison of methods in Moline & Linder �00�). Recent results obtained via PAE for some regions in Chile seem to confirm these critics: e.g. the confusion about the characteristic taxa in coastal v/s Andean areas of endemism in the Antofagasta region damages any proposed conservation strategy in Cavieres et al.’s (�00�) paper.

Anyway, as demonstrated by the bibliographical survey done by Nihei (�00� + online Appendix S�), PAE is getting very popular in biogeography, and several variations are appearing, like parsomoniy analysis of distributions (PAD), or cladistic analysis of distributions and endemism (CADE) (Nihei �00�). The coming years will surely see new theoretical and practical developments in this active research field.

4.4 The disintegration of the endemic and the south-temperate elementsThe question remains: how does it happen that central Chile has such a high degree of endemism at the genus level?

A possible answer is the gradually growing degree of isolation of the central Chilean biota since the Miocene and till the Pliocene, when the Andes reached their maximum height (chapter �). The presence of a high number of monotypic genera suggests that most of the endemic genera are representatives of old lineages that have lost must of their diversity due to extinctions.

Palaeogeographic scenarios show a ongoing exchange between neotropical and subantarctic palaeofloras from the late Cretaceous to the Pliocene (Hinojosa & Villagrán 1997), or even starting in the early Cretaceous (Barremian/Aptian). This affected the lineages of which extant angiosperms originated (Troncoso & Romero �998). The rising of the Andes during the Miocene and till the Pliocene is supposed to be one of the factors that promoted the final isolation of the populations of which the extant endemic genera derivate.

�0�

4.4.1 Disintegration of the Endemic Element

Australasiatic relationshipChilean endemic genera Gomortega, Peumus, and Lapageria are pylogenetically related to australasiatic lineages (Renner 2004, 2005; Vinnersten & Bremer 2001) (figures 4.15, 4.16). Current patterns of disjunction in many austral lineages suggest a close connection to the break-up of Gondwana (Villagrán & Hinojosa �997). Peumus, joining other extant Lauraceae taxa (Cryptocarya, Beilschmiedia, and Persea), was already a component of the middle Miocene flora in central Chile (Hinojosa & Villagrán �997). Renner (�004) proposed that Peumus diverged from the Monimia/Palmeria line about 7� my ago, by the disruption of a formerly continuous range that stretched from Chile across Antarctica and the Kerguelen plateau to Madagascar. Similarly, Gomortega is a stereotypical paleoendemic, maybe the only survivor of a much richer group. Fossil Gomortega are only known from the Late Oligocene-Early Miocene (�4-�� mya; Nishida et al. �989, quoted by Renner �004), but molecular analysis suggests an age of �00 mya for Gomortega (Renner �004). It is possible that this family was more numerous in the past and suffered from extremely high extinction rates. The alternative scenario is that the family has a very slow speciation rate (just one species existing for 100 mya!)

Figure 4.�� Phylogenetic and geographic relationships of Gomortega and Atherospermataceae

Figure 4.�� Phylogenetic and geographic relationships of Philesiaceae

Source: Renner 2004

Doryphora (E Australia)

Laureliopsis (Chile / Argentina) Laurelia novae-zelandiae (NZ)

Gomortega (Chile endemic)

Daphnandra (E Australia)

Dryadodaphne (E Australia / N Guinea)

Laurelia sempervirens (Chile endemic)

Atherosperma (Australia / Tasmania)

Nemuaron (N Caledonia)

Source: Vinnersten & Bremer 2001

Ripogonum (Australia/N. Guinea/NZ)

Lapageria (endemic Chile)

Philesia (Chile/Argentina)

�04

Neotropical relationshipChilean endemic genera Dinemandra, Dinemagonum and Leontochir have an evolutionary relationship with neotropical lineages (Davis et al. �00�, Aagesen & Sanso �00�), as well as the endemic genera of the Solanaceae (Alona, Latua, Phrodus, Vestia) and the Bromeliaceae (Fascicularia, Ochagavia), which are recognized as mainly neotropical families. Also, the Cactaceae have long been recognized as a neotropical family, and current hypotheses place its origin in central or in southernmost South America (Griffith 2004a,b) (box 7.1). The endemic genera of the Asteraceae are superficially related to a neotropical lineage, but the origin of the family as a whole has been hypothesized to have a Gondwanic origin at about 4�-�� mya (DeVore & Stuessy �99�). However, endemic genera could be much more recent, e.g. the ancestor of Moscharia (endemic) and Polyachyrus (Perú/Chile) possibly occupied a part of north-central Chile during the last uplift of the Andes in Pliocene and Pleistocene times (Katinas & Crisci �000). The sympatric distribution of Gypothamnium and Oxyphyllum in the northern coastal desert (figure 3.6) constitutes a challenge to our understanding of the factors that drove the speciation processes of these endemic genera belonging to different subtribes in the Mutisieae (Mutisiinae and Nassauviinae, respectively).

Wider relationshipsThe presence of some endemic genera closely related to North American or South African taxa is a real challenge for future research in this area. This is especially the case for the three endemic Chilean genera of Tecophilaeaceae (Conanthera, Zephyra, and Tecophilaea), a little family that has only eight genera and �� species, occurring in mediterranean areas from Africa to South America and North America. The endemic family Francoaceae (Francoa, Tetilla) is also closely related to the Greyiaceae and the Melianthaceae, two small families mainly occurring in South Africa.

Following the discussion in chapter 2, some consider ‘taxa’ at best as classification units, without any universally recognized evolutionary meaning (Hey et al. �00�). For example, Calopappus is considered by some authors as a synonym of Nassauvia (Bremer �994), and Leontochir as a synonym of Bomarea (Hofreiter �00�, but see Aagesen & Sanso �00�). Other endemic genera like Guynesomia have only recently been recognized as being different from their closest relatives (Bonifacino & Sancho �004). Furthermore, there is still a profound lack of knowledge about some endemic genera like Ivania and Metharme. Therefore, they are a good target for further systematic and biogeographical studies.

4.4.2 Disintegration of the South-temperate Element

As with the endemic element, analysing the phylogenetic relationships of south-temperate genera (distributed in souther Chile/Argentina) reveals a relationship with the other southern landmasses (table 4.�), e. g. Pilgerodendron, Lebetanthus, Huanaca, Philesia, Asteranthera, Mitraria, and Sarmienta.

�0�

Table 4.� Australasiatic relationships of South-temperate genera

Taxon Phylogenetic relationships Source

Pilgerodendron(Cupressaceae)

Genus from southern Chile/Argentina, closely related to Libocedrus (New Zealand); some authors propose to unite them in one genus.

Gadek et al. �000, but see Farjon et al. �004

Lebetanthus Epacridaceous genus from (MAG/Argentina) is closely related to Prionotes (Tasmania)

Crayn et al. �998

Huanaca(Apiaceae)

Patagonian genus close related to New Zealand Stilbocarpa Andersson et al. �00�

Philesia (+ endemic Lapageria)

the only members of the South American Philesiaceae, are supposedly splitted from the Australian–New Zealand Ripogonum at 47 +- 8,4 mya.

Vinnersten & Bremer �00�

subfamily Coronantheroideae

Members of the Gesneriaceae restricted to the Southern Hemisphere that are closely related. A Gondwanan origin for the whole family has been proposed. The small group of Coronantheroid Gesneriaceae supposedly invaded the Americas via Antarctica and southern South America and gave rise to the Gesnerioid Gesneriaceae. While the Coronantheroid Gesneriaceae became nearly extinct (the three Chilean/Argentinian genera Asteranthera, Mitraria, Sarmienta being the last survivors), the Gesnerioid Gesneriaceae evolved explosively in the American tropics.

Burtt �998, Weber �004

As example, Vinnersten & Bremer (�00�) suggest that the ancestor of Ripogonum, Lapageria, and Philesia was distributed in South America and New Zealand and possibly also in Australia. The isolation of South America from Australia and New Zealand corresponds to the split of the South American Lapageria and Philesia from the Australian–New Zealand Ripogonum estimated to 47 +- 8.4 mya. The same interpretation is possible for the split between Alstroemeriaceae and Luzuriagaceae, so it may be that termination of the Antarctic link during Eocene resulted in two vicariance events within Liliales (Vinnersten & Bremer �00�).

�0�

4.5 Plant geography of the Chilean Pacific islands

The Pacific islands offshore Chile are mostly treated as oceanic islands, following the early classification of Wallace (1880). According to the theory of equilibrium in island biogeography, the biota of these islands is the result of immigration and extinction rates (MacArthur & Wilson 1967). The Chilean Pacific islands show different situations: Juan Fernández, Desventuradas, and Easter Island (Isla de Pascua), are located at different distance and latitude in relation to the Chilean coast, and show different floristic relationships.

4.5.1 Isla de Pascua (27° 05’ S, 109° 20’ W)

Isla de Pascua (Easter Island), located at �,7�� km from Santiago, is considered the most isolated inhabited island on the planet (figures 4.4, 4.17). From a biological point of view, Isla de Pascua is the most depauperate of the Chilean Pacific islands, showing only a 7,68 % of specific endemism (Marticorena �990). There are some genera and families (mostly ferns) not present in Chile but widely distributed in the Pacific Islands and the pantropics, like Davallia, Psilotum, Vittaria, Doodia (section �.�). The long history of occupation by Polynesian folks have left a landscape and a floristic scenery that seems to be very far from the original one (prior to the human arrival) (Zizka 1991, Bork 2006). Therefore a synopsis of the native flora is very difficult, since several taxa are treated as native or alien by different authors (or idiochores v/s anthropochores sensu Zizka �99�). Based in the works of Skottsberg (�9�0-�9��) more recently revised by Marticorena (�990) and Zizka (�99�), a checklist with a total of �0 families and �9 genera was compiled (appendix D). The best represented family is the Poaceae, with � genera. Several taxa that appeared with a question mark in Zizka’s revision and considered as introduced by Marticorena (�990) have been left out from the analysis (e.g. Caesalpinia, Calystegia), while some listed by Marticorena as aliens have been retained due to the reasons exposed by Zizka, like Triumfetta (Malvaceae), or Kyllinga (Cyperaceae). The result of phytogeographic elements analysis shows a clear predominance of pantropical (4�%) and cosmopolitan genera (49%), with little presence

Figure 4.�7 Isla de Pascua (from Skottsberg �9�0-�9��).

�07

of australasiatic (7%) (Doodia, Rytidosperma) and one antitropical genus (Agrostis) (figure 4.18). The extinct palm Paschalococos disperta, known only from subfossil endocarps, is possible related to the continental endemic Jubaea chilensis.

The natural history of the island was discussed by Skottsberg (�9�0-�9��). He emphazised the floristic relatioship with the palaeotropics, and remarked that the flora is very poor, maybe due to the human influence, to allow any biogeographical conclusion.

4.5.2 Islas Desventuradas, San Félix (26°17’ S / 80°0�’) and San Ambrosio (��°��’S / 79°��’W)

This relatively small volcanic archipelago is located approximately 8�0 km off the Chilean coast (figure 4.4.). Desventuradas Islands consist of the two main islands San Félix (figure 4.19) and Isla (de) San Ambrosio (figure 4.20) plus several rocks and stacks: Islote Gonzalez and Roca Catedral. Together, the Desventuradas Islands have a surface area of only �0.� km². The topography is very rugged, with peak elevations of �9� m asl on Isla San Félix, 479 m a.s.l. on Isla San Ambrosio.

Figure 4.�8 Floristic elements Isla de Pascua

41%

7%0%3%0%

0%

49%

1. Pantropical2. Australasiatic3. Neotropical4. Antitropical5. South-temperate6. Endemic7. Cosmopolitan

Figure 4.�9 Isla San Félix. [www.cordell.org/SFX/SFX_pages/SFX_Main.html]

Figure 4.�0. Isla San Ambrosio (after Kuschel �9��)

�08

In spite of the relative sparce flora, the islands have long attracted naturalists due to the endemic genera Lycapsus, Thamnoseris (Asteraceae), Nesocaryum (Boraginaceae), and Sanctambrosia (Caryophyllaceae). Botanical descriptions of the islands flora are to be found in R.A. Philippi (�870), F. Philippi (�87�), Skottsberg (�9�7, �9��, �9��), Johnston (�9��), Gunckel (�9��), and Kuschel (�9��). More recent treatments have been done by Marticorena (�990) and Hoffmann & Teillier (�99�)��. According to these authors, the vascular flora of the islands consists of 13 families, �8 genera and �� native species. More recently Teillier & Taylor (�997) add one genus to the list, Maireana (Chenopodiaceae), with one species formerly known only from Australia. Based in the works of these authors, a checklist with a total of �� families and �9 genera was compiled (appendix D). The best represented family is the Amaranthaceae (4 genera formerly classified under the Chenopodiaceae). The floristic element that dominate in the islands is the cosmopolitan element (52%), but the endemic element is also noteworthy, reaching a 21% (figure 4.21). This percentage is higher than the one showed by the continental flora or Juan Fernández at the genus level. Furthermore, Marticorena (�990) reports a level of endemism of �0,�% at the species level, the highest for a Chilean region.

The natural history of the archipele has been analysed by Skottsberg (�9�7), coming to a similar conclusion that for the Juan Fernández Islands, in the sense that the Desventuradas flora do not show a oceanic character but a continental one. This view can be challenged by the recent report of the genus Maireana, formerly known only from Australia and hypothetized as the result of a recent long-distance dispersal event (Teillier & Taylor �997). The contemporary discovery of the genus in the coast of Atacama region (Marticorena �997) tend to support this hypothesis, since it is doubtful that a shrub stayed unnoticed for botanists till nowadays. This could be one of the few real evidences for the effective operation of long-distance dispersal (see discussion in section �.�). On the other hand, the existence of four endemic angiosperm genera and �0 endemic species reinforces the view of an old floristic history not just explicable by recent migration events.

Figure 4.�� Floristic elements Islas Desventuradas

0% 5%

11%

11%

0%

21%

52%

1. Pantropical2. Australasiatic3. Neotropical4. Antitropical5. South-temperate6. Endemic7. Cosmopolitan

�09

Noteworthy, in his thesis about the biogeography of Brassicaceae, Brüggemann (�000) suggest that the American Lepidium can have originated via long-distance dispersal from Australia or North America, but on the other hand he suggests that the Pacific taxa have been translated by Polynesian folks. Brüggemann (�000) mentions as example the absence of Lepidium from Juan Fernández, which apparently was not reached by the Polynesian. The presence of the endemic species L. horstii in the Desventuradas Islands is contrary to this suggestion.

4.5.3 Archipiélago de Juan Fernández

The archipelago, located between 667 and 850 km from the continent (figure 4.4), is formed by two main islands and one islet:

- Alejandro Selkirk Island (��°4�’ S / 80°47’W) (also known as Isla Más Afuera), 8�0 km from the American continent. Its highest elevation is Cerro Los Inocentes (1.380 m a.s.l.) (figure 4.22).

- Robinson Crusoe Island, (��°�8’ S / 78°��’ W) (also known as Isla Más a Tierra), located at around ��7 km from the continent. Its highest peak is Cerro El Yunque (9�� m a.s.l.). Close to Robinson Crusoe there are two islets: Islote Juanango, and Santa Clara, this latter located � km southwest of Robinson Crusoe (figure 4.23).

The archipelago is world wide known as having been the scenary for the real history that inspired the novel of Robinson Crusoe. Therefore the island Masafuera was renamed in honour of the Schottich seaman Alejandro Selkirk, who survived 4 years and 4 months on Masatierra. Since its official discovery in 1574 by the pilot Juan Fernández (maybe even before), Masatierra was an obligate anchor place for seamen and bucaneers after trespassing the Cape Horn, and therefore dozens of treasure histories characterize the island. But several botanists have long advice that the real treasure of this archipele is in its unique plant world.

The flora of the archipelago has been long a subject of interest to botanists (e.g. Gay 1832, Johow �89�, Skottsberg �9�0-�9��, Muñoz Pizarro �9�9, Stuessy et al. �984b, Marticorena et al. �998b,

Figure 4.�� Isla Alejandro Sel-kirk (Masafuera)

Figure 4.��. Isla Robinson Crusoe (Masatierra) (from Skottsberg �9�0-�9��)

��0

Danton et al. 2000, Danton & Perrier 2003). The native vascular flora comprises 56 families and ��0 genera. The best represented family are the Poaceae (�� genera) and the Asteraceae (9). The Fernandezian flora has 12 endemic genera (plus 3 endemic genera from the continent and the islands: Hymenoglossum, Ochagavia and Notanthera) (section �.�), and �04 endemic species (Stuessy et al. �998a). Unfortunatelly the combining invasion of browsing animals and continental plants place the island’s native flora at a competitive disadvantage (Dirnböck et al. 2003), so that at least 75% of the endemic flora is high endangered (Cuevas & van Leersum 2001, Stuessy et al. �998b). Swenson et al. (�997) reported a number of ��7 introduced species, and the number is continuously growing.

The origin of the Fernandezian flora has been the subject of considerable study and debate (e.g. Skottsberg 1925, 1936, 1956, van Balgooy 1971). Traditionally the origin for an island flora has been explained in direct relationship with it nearest continental mass by means of long-distance dispersal (e.g. Carlquist �974). Oceanic islands are often explained as geologically new territories and Juan Fernández is not the exception: Isla Robinson Crusoe has been dated at ca. 4 mya, Isla Alejandro Selkirk at �-� mya, and Isla Santa Clara at �.8 mya (Stuessy et al. �984a). The islands are supposed therefore to be the products of isolated intraplate volcanism associated with an hotspot. The relative youg age of the archipelago seems to leave no doubt for a recent origin for its flora, as carefully revised by Bernardello et al. (2006).

But the high level of endemism, the variety of floristic relationships and the limited methods of dispersal put some unresolved questions in this theme. Bernardello et al. (�00�) report that 80% of the island species have dry fuits, and fleshy fruits are comparatively uncommon, challenging the supposed ability for bird long-distance dispersal. In fact, the dispersal syndrome that prevails in the flora is autochory (i.e. autonomous passive dispersal). Bernardello et al. (2006) therefore suggest that the principal dispersal processes are anemochorous dispersal (air flotation) and epizoochory (carrying by birds attached to feathers). But since the extant bird fauna is scarce, the real opportunities for dispersal are relatively few.

This was early recognized by Skottsberg (�9��, �9��) and he therefore suggested that the origin of the island flora should be find in alternative palaeoscenarios. Skottesberg revised and discussed all available evidence in floristic and faunistic elements, in geotectonics of the Pacific, and in the continental Tertiary flora and came to the conclusion that the Fernandezian flora is not a one of oceanic but of continental nature. Skottsberg therefore proposed a tentative sketch on the history of the Fernandezian flora that is in concordance with Brüggen’s Tierra de Juan Fernández, an old submerged landmass west of today South America (Brüggen �9�0, Skottsberg �9��, p. �94; see figure 5.12).

Skottsberg’s critical vision has been systematically oversimplified by modern authors, while emphazising that he noted the close relationship with the American flora (e.g. Crawford et al. 1990, Bernardello et al. 2006). Skottsberg certainly recognized the floristic relationship between the islands and the American continent, but he also noted the closest relatioship with the western

���

Pacific and especially with Australasia. Indeed, several genera not found in continental Chile show a wider distribution in Australasia, like Coprosma (Rubiaceae), Arthropteris (Oleandraceae), Haloragis (Haloragaceae) or Santalum (Santalaceae). The analysis of floristic elements of the Juan Fernández flora shows a 14% of neotropical genera and 11% are australasiatic genera. Most important is the cosmopolitan element (�8%), and also the pantropical (�7%) the endemic (�4%) and the antitropical elements (12%) (figure 4.24).

Furthermore, including Juan Fernández (JF) into the similarity analysis done between Chilean regions (section 4.2) results in a closer floristic relationship between JF/MAG than between JF and the other Chilean regions (table 4.6, figure 4.25).

Table 4.�. Floristic similarity between regions including Juan Fernández

Regions SIMIL DISTANT/COQ 0,47 777ANT/BIO 0,�9 ���7ANT/MAG 0,�� ���7COQ/BIO 0,�4 7�0COQ/MAG 0,�7 �4�0BIO/MAG 0,�4 �700JF/MAG 0,18 ����JF/BIO 0,�� 79�JF/COQ 0,�� 89�JF/ANT 0,08 �4�9

Noteworthy, in figure 4.25 the relationship distance/similarity follows a trend, but the relationship MAG/JF escapes from this trend (outlier). This closer floristic relationship between MAG and JF appeals to two different explanations:

Figure 4.�4 Floristic ele-ments Juan Fernández

17%

11%

14%

12%4%

14%

28%

1. Pantropical2. Australasiatic3. Neotropical4. Antitropical5. South-temperate6. Endemic7. Cosmopolitan

���

a) The Magallanes biota had once a more northward distribution, till central Chile, and from there it reached the islands via long-distance dispersal. The presence of south-temperate elements in the Fray Jorge fog forest at �0°40’ has been long suggested as an evidence of the more widespread temperate forests during the Cenozoic. But the remnants of these forests are still represented in the coast of BIO and there is no apparent reason why the relation BIO/JF stays in the trend of similarity/distance, contrary to MAG/JF.

b) An alternative explanation for the floristic similarity between JF and MAG is a direct connection of the two land masses. This explanation is contrary to current geological consensus, since the islands seem to be geologically too young for this type of explanation. The islands are located on top of the Juan Fernández Ridge, that controls the tectonic and geological evolution of the southern Andes at ��°-�4°S since the Tertiary (Yáñez et al. 2001). The seafloor age assigned by Müller et al. (1997) to the seafloor offshore Chile between �8°S and 40°S, based on magnetic anomalies and relative plate reconstructions, ranges from 20 to 48 mya. Juan Fernández rests on seafloor dated at around 20 to 33 mya.

Some early geologists believed in a former land west of South America occupying the whole Pacific (Burckhardt 1902, Beloussov 1968), or at least a portion of it as a Transandiner Kontinent close to the South American coast (Muñoz Cristi �94�, Cañas �9��, Illies �9�7). Brüggen (�9�0,

0 2000 4000 6000 8000 10000

0.0

0.2

0.4

0.6

0.8

1.0

Distance [km]

Jacc

ard

sim

ilarit

y

ANT/COQ

ANT/BIO

ANT/MAG

COQ/BIO

COQ/MAGBIO/MAG

JF/MAGJF/BIO

JF/COQJF/ANT

Figure 4.�� Floristic similarity between different regions in Chile: Antofagasta (ANT), Biobío (BIO), Coquimbo (COQ) and Magallanes (MAG). The Jaccard similarity index as related to geographical distance within Chile (squares), and between Juan Fernández and Chilean continental regions (circles). The lines represent trend curves as an exponential curve. The relation JF/MAG (filled circle) is considered as an outlier and is not included in the trend curve. The arrow show the possible position of the relation JF/MAG following the trend (see discussion in the text).

���

p. �9) proposed the name Tierra de Juan Fernández for this larger continental land mass west of current South America. Miller (�970) analysed different possibilities for the disappearing for this land and concluded that the Ozeanisierung of Juan Fernández Land occurred in Late Tertiary, and that this land was not at all at the same location of today’s Juan Fernández archipelago (Miller 1970, p. 934). From floristic similarity as analysed in the present thesis, this land should have been existed much more to the South. To fit in the trendline of the relation similarity/distance shown by other Chilean regions, the Juan Fernández islands (Juan Fernández Land) could be located at the same longitude (80°W), but at 48°S instead of ��°, i.e. �� degrees latitude or around �.��0 km towards the South. (figure 4.26).

The presence of the endemic family Lactoridaceae seems to confirm the continental nature of the Fernandezian flora. The cladistic analysis done by Lammers et al. (1986) suggest that Lactoridaceae diverged sometime prior to the Maastrichtian (�9 mya). This has been corroborated recently by the analysis of Wikström et al. (�00�): Lactoris appears as a very ancient taxon in the base of the angiosperms supertree: the split between Lactoris and Aristolochia has been dated at around 8� mya.

In Lammers et al.’s (�98�) opinion “it seems unlikely that the Lactoridaceae evolved autochthonously in the Juan Fernandez Islands. A more plausible hypothesis is that the plants on Masatierra are relicts of a once more extensive continental distribution in South America and possibly other portions of the Southern Hemisphere, perhaps originating from the western Gondwanaland flora”. Indeed, microfossils related to Lactoris and referred to the fossil genus Lactoripollenites have been found in South Africa and Australia (Macphail et al. �999), thus suggesting that the Lactoridaceae were widespread across the Southern Hemisphere during the Late Cretaceous (Lammers et al. �98�). “Differences between Lactoripollenites and Lactoris pollen imply that these represent different clades within the Lactoridaceae or that the former evolved into the latter genus elsewhere in the Southwest Pacific region prior to its migration onto Masatierra in the Plio-Pleistocene (Macphail et al. �999). Crawford et al. (�00�) suggest that the species or its ancestors could have reached Masatierra as means of long-distance dispersal by wind, due to the small seeds, however the authors recognize that “the plants occur primarily in the forest understory, wich would seemingly minimize the effectiveness of wind as a means of long-distance dispersal” Crawford et al. �00�, p. �89).

Figure 4.�� Possible position of the hypotheaized Tierra de Juan Fernández, according to floristic similarity at the genus level (see figure 4.25).

��4

Other endemic taxa might be also the remnants of older groups that have evolved in a completely different palaeogeographic scenario. As example, the shrubby Fernandezian Wahlenbergia species, together with the species from New Zealand and St. Helena, are considered as the more basal members of the wahlenbergioid group, suggesting a Gondwanic origin (Eddie et al. �00�).

Stuessy et al. (�984a) were aware of the earlier view from Brüggen (�9�0) and Skottsberg (�9��), but they gave much value to the Potassium-argon dating. Thus most of the papers dealing with the evolution of the Fernandezian flora in the last two decades start from the 4 mya date. The dating of Santa Clara puts another question on the problem, since this little islet seem to be almost two million years (�,8 +-�,� mya) older than Robinson Crusoe. Stuessy et al. (�984a), in spite of their high confidence in the geological datations, do not rule out other models of Pacific aseismic ridges (e.g. Nur & Ben-Avraham �98�b). Also Stuessy et al. (�984b) recognize that the islands could have been much more extense and have been rapidly eroded during the last 4 million years (figure 4.27).

Figure 4.�7 Reconstruction of geomorphological history of Masatierra + Santa Clara. A: shape of original island at 4 mya; B: erosional patterns showing amphitheater valleys; C: present configuration showing bathymetric contours (adapted from Stuessy et al. �984b).

A B C

���

� Palaeogeography: insights into the Evolution of the Chilean Flora

���

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

��7

5 Palaeogeography: insights into the Evolution of the Chilean Flora

Stuessy & Taylor (�99�) suggest (based on van der Hammen & Cleef �98�) that there are three principal sources of information for interpretating the origin of a flora:

a) evidence of geomorphological changes in the landscape;b) the macro- and microfossil record;c) biogeographical relationships between extant floras.

Stuessy & Taylor (1995) further mention the three principal geological influences that shaped the present Chilean flora:

a) continental movements (due to plate tectonics);b) oceanic transgressions over the continental surface;c) the Andean uplift.

5.1 Continental movements: fragmenting the Earth’s surface

Plate tectonics is the current paradigm in geology (and related sciences like biogeography), after the general acceptance of Alfred Wegener’s Kontinentalverschiebungs theory. The Earth’s surface seems to rest on twelve major tectonic plates that interact at their borders. Under the paradigm of plate tectonics, the floral history of southern South America is intrinsically related to the geologic, tectonic and climatic history of the Gondwana continent.

The term Gondwana derive from the non-marine sedimentary rocks exposed in a series of graben-type basins of the Indian peninsula, that relate this land to the rest of Gondwanic landmasses concentrated today in the southern hemisphere (McLoughlin �00�). Epistemologically, the term remembers the ancient kingdom of the Gonds, the people still inhabiting the area south of the Narmada River in central India (Singh �944). Gondwanan has become synonymous with the southern hemisphere biota, however, McLoughlin (�00�) notes that the distinctive Indian Gondwana sedimentary sequences and fossils were deposited while the southern continents were united to Laurasia (Permian to early Cretaceous).

Several attempts have been made to relate the biogeographical information with the break-up of the Gondwana continent (e.g. Villagrán & Hinojosa �997, McLoughlin �00�, Sanmartín & Ronquist �004). Most authors recognize three major separation events that affected the evolution of the South American flora: the separation between W and E Gondwana between 180-150 mya, the separation America/Africa between ��9-�0� mya, and the split between Antarctica and southern South America (��-�8 mya) (table �.�).

Paleoreconstructions for Gondwana landmasses’s lateral movements since the Late Devonian (360 mya) is presented in figure 5.1. The reconstruction has been done with the program Timetrek v. �.� (Cambridge Paleomap Services Ltd.).

��8

Table 5.1 Dating major palaeogeographic events = 3 stage break-up of Gondwana

Major separation events

Age (mya)

Period Possible cause Source

(W Gondwana / E Gondwana)

�80-��0

Early to Late Jurassic

Breakup associated with development of a series of deepseated mantle plumes beneath the extensive Gondwanan continental crust (e.g in S Africa (ca. �8� mya) and the Transantarctic mountains (c. �7� mya)

Scotese & McKerrow �990, Storey �99�, Reeves & de Wit �000

Africa – South Am separation

��9-�0�

Early Cretaceous

Opening South Atlantic Ocean, due to the emplacement of Plume-related Parana-Etendecka continental flood basalts in Brazil and Namibia (��7-��7 Mya). Final break-up of Africa and S Am was completed only at 80 mya)

Turner et al. �994, Jones �987, Barker et al. �99�, Grunow et al �99�, MacLoughlin �00�

West Antarctica-S Am

��-�8 Palaeogene/Oligocene

Beginning at ~��-�0,� mya as a subsidence in the Powell Basin followed by seafloor spreading. Note in the TimeTrek reconstruction (figure 5.4) that Antarctica and America are already separated at ��0 mya, and closer again at 80 mya).

Lawver & Gahagan �998

��0 mya ��0 mya

�00 mya �4� mya

Figure �.� Palaeoreconstructions of Gondwana since Late Devonian (��0 mya). Program TimeTrek v. �.�., Cambridge Paleomap Services Ltd. (trial version for download disponible at http://www.the-conference.com/CPSL/timetrek.htm).

��9

�00 mya80 mya

�0 mya �4 mya

�0 mya 0 mya

Figure �.� (continued)

��0

McLoughlin (�00�) summarized the impact of the Gondwana break-up on the evolution of current southern biota. He traced back the origin of the floras in five major phases of development before the appearance of angiosperms (table �.�). The Permian is characterized by the Glossopteris flora, the Triassic by the Dicroidium flora (figures 5.2, 5.3).

Figure �.� Distribution of the Permian Gloss-opteris flora in Gondwana (after McLoughlin �00�). Palaeo-biomes after Willis & McEl-wain �00�

Subtropical desert

Glossopteris palaeoflora

Cold temperate

Cool temperate

Mid-latitude desert

Middle Permian (c. 260 Ma)

Figure �.� Distribution of the Triassic Dicroidium flora in Gondwana (after McLoughlin 2001). Palaeo-biomes after Willis & McElwain �00�

Subtropical desert

Dicroidium palaeoflora

Warm temperate

Cool temperate

Winterwet

Late Triassic (c. 200 mya)

Tropical summerwet

Glossopteris fossil leafs (www.paleon-tology.unibonn.de/glossopteris.htm)

Dicroidium leafs (www.rosssea.info/geology.html)

���

Tabl

e 5.

2 G

ondw

anan

flor

as: fi

ve m

ajor

pha

ses o

f dev

elop

men

t bef

ore

the

appe

aran

ce o

f ang

iosp

erm

s (se

nsu

McL

ough

lin 2

001)

Flor

asA

ge

(mya

)Pe

riod

Com

posi

tion

Pala

eoen

viro

nmen

tal c

ondi

tions

�.

Cos

mop

olita

n ea

rly la

nd p

lant

flo

ras

~4�0

-�9

0Si

luria

n –

Early

D

evon

ian

Free

-spo

ring,

her

bace

ous f

orm

s occ

upyi

ng m

oist

hab

itat (

Coa

stal

are

as, r

iver

flat

s, de

lta sw

amps

, lak

e m

argi

ns) =

Rhy

nops

ida,

Zos

tero

phyl

lops

ida,

Trim

erop

hyte

s and

ly

copo

d fo

ssils

.

Foss

ils o

f Zos

tero

phyl

lum

and

the

lyco

pod

Bara

gwan

athi

a pr

esen

t in

both

hem

isph

eres

sugg

est l

ittle

flor

istic

pr

ovin

cial

ism

(McL

ough

lin �

00�)

, but

seve

ral a

utho

rs

reco

gniz

e at

leas

t five

bio

geog

raph

ical

uni

ts (s

ee W

illis

&

McE

lwai

n �0

0�)

�. A

rbor

esce

nt

lyco

pod

and

seed

-fer

n flo

ras

in a

coo

ling

clim

ate

�90

�70

��0

Mid

dle

Dev

onia

n

Late

Dev

onia

n

Car

boni

fero

us

Her

bace

ous t

o sh

rub-

size

d ly

coph

ytes

attr

ibut

ed to

Hap

lost

igm

a, L

ecle

rcqi

a,

Arch

aeos

igill

aria

and

Pro

tole

pido

dend

ron.

Emer

genc

e of

firs

t arb

ores

cent

pla

nts:

Lep

idod

endr

on a

nd p

rogy

mno

sper

ms.

Seed

-fer

n flo

ras p

rogr

essi

vely

repl

aced

lyco

phyt

e flo

ras.

Gen

eral

coo

ling

clim

ate,

with

at l

east

four

per

iods

of

sout

hern

hem

isph

ere

glac

iatio

n. W

Gon

dwan

a ro

tate

d in

to

pola

r lat

itude

s. Pa

rts o

f cen

tral a

nd n

orth

er S

Am

wer

e af

fect

ed b

y gl

acia

tion.

E G

ondw

ana

rem

aine

d in

mid

dle

latit

udes

. Stro

ng la

titud

inal

gra

dien

t per

mitt

ed th

e in

itial

de

velo

pmen

t of a

dis

tinct

ive

S he

mis

pher

e flo

ra (M

eyen

19

87) a

nd th

e fir

st e

xpre

ssio

n of

intra

-Gon

dwan

an fl

oris

tic

prov

inci

alis

m.

�. T

he

glos

sopt

erid

flo

ra

�00-

���

Perm

ian

Gin

kgoa

les,

coni

fers

, Cyc

adal

es, C

orda

ites.

Glo

ssop

teri

s is a

gro

up o

f ext

inct

gy

mno

sper

ms.

App

ears

in th

e fo

ssil

reco

rd a

t abo

ut th

e en

d of

the

Car

boni

fero

us in

W

Gon

dwan

a.

Early

Per

mia

n G

ondw

ana

lock

ed in

dee

p gl

acia

tion.

M

iddl

e to

Lat

e Pe

rmia

n w

arm

er c

limat

ic c

ondi

tions

fa

vour

ed e

xpan

sion

of G

loss

opte

ris,

that

dis

appe

ar fr

om

all c

ontin

ents

alo

ng w

ith c

a. 8

0% o

f the

wor

ld’s

bio

ta

durin

g th

e Pe

rmia

n-Tr

iass

ic e

xtin

ctio

n..

4. T

he

Dic

roid

ium

flo

ra

��0-

�0�

Tria

ssic

Mor

e di

vers

e flo

ras d

omin

ated

by

Dic

roid

ium

(see

als

o H

erbs

t et a

l. �0

0�)

(Cor

ysto

sper

mal

es),

voltz

ialn

ean

coni

fers

, gin

kgop

hyte

s, pe

ltasp

erm

s, pu

tativ

e gn

etal

es, b

enne

ttita

lean

s, pe

ntox

ylal

eans

and

cyc

adop

hyte

s, pl

us m

any

lyco

phyt

es

and

osm

unda

ceae

n, g

leic

heni

acea

n, d

icks

onia

ceae

m d

ipte

ridac

ean

and

mar

attia

cean

fe

rns.

Maj

or ra

diat

ion

of th

e co

nife

rs.

The

inte

rior o

f Pan

gea

was

hot

and

dry

. War

m T

empe

rate

cl

imat

es e

xten

ded

to th

e Po

les.

Poss

ibly

one

of t

he h

otte

st

times

in E

arth

his

tory

. The

re w

as n

o ic

e at

eith

er N

orth

or

Sout

h Po

les.

War

m te

mpe

rate

con

ditio

ns e

xten

ded

tow

ards

th

e po

les.

�. Ju

rass

ic

pter

idos

perm

-co

nife

r flor

as

�00-

�4�

Jura

ssic

Con

ifer-

and

bene

ttita

lean

- dom

inat

ed c

omm

uniti

es re

plac

ed D

icro

idiu

m-

dom

inat

ed fl

oras

acr

oss t

he S

hem

isph

ere.

Pla

nt g

roup

s tha

t hav

e th

eir o

rigin

s in

the

Tria

ssic

reac

hed

mor

e im

porta

nce

in th

e Ju

rass

ic: c

ayto

nial

eans

, ben

netti

tale

ans,

pent

oxyl

alea

ns a

nd p

achy

pter

id se

ed-f

erns

. Als

o m

arat

taic

ean,

mat

onia

cean

, os

mnu

ndac

ean,

dis

ckso

niac

eae

and

dipt

erid

acea

n fe

rns,

equi

seta

lean

s, he

rbac

eous

ly

coph

ytes

, and

cyc

adal

eans

. Mos

t gen

era

pers

ist i

nto

the

earli

est C

reta

ceou

s. A

rauc

aria

n, p

odoc

arp,

che

irole

pida

cean

and

taxo

diac

ean

coni

fers

, cay

toni

alea

ns,

benn

ettit

alea

sn a

nd g

inkg

oale

ans d

omin

ated

bro

ad tr

acts

of b

oth

N a

nd S

he

mis

pher

e.

Con

ditio

ns g

ener

ally

war

m a

nd h

umid

, loc

al c

oal d

epos

its,

lack

of g

laci

atio

n. P

arts

of S

Am

and

S A

fr e

xper

ienc

ed

arid

con

ditio

ns. T

he c

osm

opol

itan

spre

ad o

f con

ifers

and

th

e bi

ota

in g

ener

al is

inte

rpre

ted

as a

con

sequ

ence

of

mor

e eq

uabl

e gl

obal

clim

ate

and

a la

ck o

f phy

sica

l bar

riers

to

pla

nt m

igra

tion

(=ra

nge

expa

nsio

n).

���

For the Late Mesozoic (i.e. Cretaceous) and the Cenozoic, several models for the history of the vegetation and climate have been proposed. Romero (�98�, �99�) proposed three main palaeofloristic types: a Neotropical palaeoflora, a Mixed, and an Antarctic palaeoflora. Troncoso & Romero (1998) refined this classification and recognized twelve palaeofloristic types. Hinojosa & Villagrán (1997) recognized five paleofloras (Tropical, Antarctic, Mixed, Sutropical mesic, and Sutropical xeric), later renamed as Gondwanic, Subtropical Gondwanic, Mixed, and Subtropical Neogene palaeofloras (Hinojosa 2005). The palaeoscenarios proposed by Romero (1993), Hinojosa & Villagrán (�997), Troncoso & Romero (�998), and Hinojosa (�00�) are summarized in table �.� and illustrated in figure 5.4, on the base of Time Trek palaeoreconstructions and palaeobiomes as proposed by Willis & McElwain (�00�).

Subtropical desert Tropical palaeoflora

Warm temperate

Cool temperate

Late Cretaceous (c. 70 mya)

Tropical summerwet

Mid-latitude desert Nothofagus expansion

Early Eocene (c. 50 mya)

Subtropical desert

Warm temperate

Cool temperate

Subtropical summerwet

Tropical everwet

Figure 5.4 Evolution model for the Chilean flora on the base of Time Trek palaeoreconstructions and the palaeobiomes sensu Willis & McElwain �00�) (see also table �.�)

A

B

���

Early Oligocene (c. 30 mya)

Warm / cool temperate

Tropical palaeoflora

Cool / cold temperate

Glacial

Subtropical summerwet

Mixed palaeoflora

Antarctic palaeoflora

Tropical everwet

Cold temperate / arctic

Miocene (c. 10 mya)

Winterwet

Warm temperate

Subtropical summerwet

Tropical everwet

Glacial

Cool temperate

Arctic

Pliocene (5 - 2 mya)

Desert (arid)

Subtropical summerwet

Subtropical summerwet

Tropical everwet

Glacial

Cool temperate

Arctic

Desert (hyperarid)

Andean uplift

Figure 5.4 (continued) Evolution model for the Chilean flora on the base of Time Trek palaeo-reconstructions and the palaeobiomes sensu Willis & McElwain �00�) (see also table �.�) (for the Pliocene see also Dowsett et al. �999 paleoreconstruction)

C

D

E

��4

Tabl

e 5.

3 C

reta

ceou

s to

Neo

gene

evo

lutio

n of

the

flora

s of S

outh

Am

eric

a, m

ainl

y fr

om R

omer

o (1

993)

, Hin

ojos

a &

Vill

agrá

n (1

997)

, Tro

ncos

o &

Rom

ero

(199

8),

Will

is &

McE

lwai

n (�

00�)

, Hin

ojos

a (�

00�)

.

Pala

eoflo

ras

Age

(mya

)Pe

riod

Com

posi

tion

Pala

eoen

viro

nmen

tal c

ondi

tions

�. P

rimiti

ve

angi

ospe

rms

�4�-

�00

Early

C

reta

ceou

s Fl

oras

dom

inat

ed b

y a

dive

rsity

of c

onife

r and

pte

ridos

perm

gro

ups.

Gym

nosp

erm

ae (B

rach

yphy

llum

and

Ptil

ophy

llum

), G

ingk

oale

s, C

ycad

opsi

da, C

onife

rops

ida

(Ara

ucar

iace

ae, P

odoc

arpa

ceae

, C

heiro

lepi

dace

ae).

Poss

ibly

ang

iosp

erm

foss

il po

llen

(Cla

vatip

olle

nite

s and

Li

liaci

dite

s) fr

om th

e B

erria

sian

(lim

it Ju

rass

ic/ C

reta

ceou

s)

Mild

“ic

ehou

se”

wor

ld. C

ool t

empe

rate

fore

sts

cove

red

the

pola

r reg

ions

.

�. N

eotro

pica

l (w

ithou

t N

otho

fagu

s)

(with

N

otho

fagu

s)se

nsu

Tron

coso

&

Rom

ero

�998

)

99-8

�

8�-�

�

Late

Cre

tace

ous

(Cen

oman

ian

to

Con

iaci

an)

Cam

pani

an-

Maa

stric

htia

n

Big

cha

nge!

: rep

lace

men

t of g

ymno

sper

ms b

y th

e an

gios

perm

s as

dom

inan

t. La

urac

ea, S

terc

ulia

ceae

, Big

noni

acea

e, M

onim

iace

ae (e

xtan

t ge

nera

Lau

relia

, Peu

mus

). Sc

hino

psis

= tr

opic

al fo

rest

and

a m

ore

xero

fitic

co

mm

unity

.

Neo

tropi

cal fl

ora

with

mar

gina

l pre

senc

e of

Not

hofa

gus.

Cam

pani

an fi

rst a

ppea

ranc

e of

Not

hofa

gus i

n A

ntar

ctic

a. M

aast

richt

ian

first

app

eara

nce

of N

otho

fagu

s in

the

foss

il re

cord

from

Cen

tral C

hile

and

Ti

erra

del

Fue

go. I

n sp

ite o

f its

mar

gina

l pre

senc

e, it

is th

e pe

ak o

f nor

ther

n ex

pans

ion

of N

otho

fagu

s in

S A

m, r

each

ing

�0°S

.

Six

glob

al b

iom

es a

re re

cogn

ized

at t

he e

nd

of th

e C

reta

ceou

s (W

illis

& M

cElw

ain

�00�

), fiv

e of

them

to b

e fo

und

in S

Am

(figu

re 5

.4

A)

�. T

ropi

cal

Gon

dwan

ic (s

ensu

H

inoj

osa

�00�

)

��-�

�Pa

laeo

cene

Trop

ical

fore

sts w

ith p

alm

s and

man

grov

es, M

yrta

ceae

, Lau

race

ae,

Prot

eace

ae w

ith m

argi

nal p

rese

nce

of N

otho

fagu

s. A

zona

l pre

senc

e of

gym

nosp

erm

s (C

heiro

lepi

dace

ae, A

rauc

aria

ceae

, Pod

ocar

pace

ae,

Zam

ianc

eae.

Hig

h pr

esen

ce o

f Not

hofa

gidi

tes i

n Ti

erra

del

Fue

go. A

num

ber

of fa

mili

es re

cord

ed in

Sou

th A

mer

ica

have

a p

antro

pica

l to

subt

ropi

cal

dist

ribut

ion:

Ana

card

iace

ae, A

nnon

acea

e, A

race

ae, A

reca

ceae

, Com

bret

acae

, M

yric

acea

e, O

laca

ceae

, Rhi

zoph

orac

eae,

Rut

acea

e, S

apin

daca

e, Z

amia

ceae

, so

me

of th

em w

hich

are

bes

t rep

rese

nted

in S

Am

: Bom

baca

ceae

, Lo

rant

hace

ae, M

elas

tom

atac

eae.

Als

o so

me

Old

wor

ld fa

mili

es, l

ike

Pand

anac

eae,

Cas

uarin

acea

e, C

aprif

olia

ceae

. Aus

trala

siat

ic g

ener

a (E

ucry

phia

, Em

both

rium

, Dri

mys

), an

d su

btro

pica

l Sap

inda

ceae

.

War

m a

nd tr

opic

al c

ondi

tions

dom

inat

ed in

W

Gon

dwan

a.

At t

he e

nd o

f the

Pal

aeoc

ene,

the

plan

et h

eate

d up

in o

ne o

f the

mos

t rap

id (i

n ge

olog

ic te

rms)

an

d ex

trem

e gl

obal

war

min

g ev

ents

reco

rded

in

geo

logi

c hi

stor

y, c

alle

d th

e Pa

laeo

cene

-Eo

cene

The

rmal

Max

imum

.

���

4.

Subt

ropi

cal

Gon

dwan

ic (s

ensu

H

inoj

osa

�00�

)

��-4

0Eo

cene

New

fam

ilies

like

Ara

liace

ae, G

uttif

erae

, Mel

iace

ae, S

terc

ulia

ceae

, Eu

phor

biac

eae,

Big

noni

acea

e, M

alpi

ghia

ceae

, Bra

ssic

acae

, Sol

anac

eae,

A

ster

acea

e, C

heno

podi

acea

e, a

nd E

ricac

eae.

Dom

inan

ce o

f a su

btro

pica

l clim

ate,

mor

e w

arm

than

toda

y bu

t coo

ler t

han

at th

e Pa

leoc

ene-

Eoce

ne. T

he E

ocen

e gl

obal

clim

ate

was

per

haps

the

mos

t hom

ogen

eous

of t

he

Cen

ozoi

c; p

olar

regi

ons w

ere

muc

h w

arm

er

than

toda

y; te

mpe

rate

fore

sts e

xten

ded

right

to

the

pole

s (fig

ure

5.4

B).

�. M

ixed

(sen

su

Hin

ojos

a �0

0�)

40-�

� O

ligoc

ene

Mix

ture

of t

he tr

opic

al e

lem

ents

with

ant

arct

ic e

lem

ents

(figu

re 5

.4 C

). Su

bant

arct

ic fl

ora

prev

ails

till

toda

y in

sout

hern

mos

t SA

m a

nd T

ierr

a de

l Fue

go. R

educ

tion

of fo

rmer

impo

rtant

Sap

inda

ceae

and

Lau

race

ae,

extin

ctio

n of

Mor

acea

e, A

nnon

acea

e, D

illen

iace

ae, M

alpi

ghia

ceae

, Vo

chys

iace

ae, T

iliac

eae,

Ste

rcul

iace

ae, S

apot

acea

e, a

nd S

tyra

cace

ae.

Gen

eral

coo

ling.

Sep

arat

ion

of A

ustra

lia

from

Ant

arct

ica

c. �

7 m

ya, o

rigin

of t

he

circ

uman

tarc

tic o

cean

circ

ulat

ion

and

the

E A

ntar

ctic

gla

ciat

ion.

�. S

ubtro

pica

l N

eoge

ne��

-�M

ioce

neA

ustra

lant

arct

ic c

old

elem

ent i

s rep

lace

d by

mor

e m

esic

ele

men

t. D

evel

opm

ent o

f Med

iterr

anea

n-ty

pe sc

lero

phyl

lous

veg

etat

ion

in C

entra

l C

hile

. Firs

t rec

ord

(Cer

ro L

os L

itres

, ��°

�8’)

of e

xtan

t scl

erop

hyllo

us ta

xa

from

Cen

tral C

hile

: Bei

lsch

mie

dia,

Peu

mus

, Myr

ceug

enia

, and

Cry

ptoc

arya

, to

geth

er w

ith so

me

taxa

that

cor

resp

ond

toda

y to

the

subt

ropi

cal f

ores

ts o

f th

e E

And

es =

Ath

yana

and

Cho

clos

perm

um.

Glo

bal m

id-M

ioce

ne w

arm

clim

atic

opt

imum

. W

ell-d

efine

d cl

imat

ic b

elts

stre

tche

d fr

om p

ole

to e

quat

or (fi

gure

5.4

D)

7. S

ubtro

pica

l N

eoge

ne (m

ore

xero

phyt

ic)

�-�

Plio

cene

Fr

agm

enta

tion

of fo

rmer

con

tinue

subt

ropi

cal f

ores

ts. O

rigin

of d

isju

nctio

ns

betw

een

cent

ral C

hile

and

NE

Bra

zil.

In

crea

sing

arid

ity in

subt

ropi

cal S

Am

, as a

co

nseq

uenc

e of

the

sepa

ratio

n of

Ant

arct

ica

from

S A

m, t

he g

laci

atio

n of

W A

ntar

ctic

a, th

e es

tabl

ishm

ent o

f the

Hum

bold

t cur

rent

, and

th

e fin

al u

plift

of t

he A

ndes

and

its i

ncre

asin

g ra

in sh

adow

effe

ct (fi

gure

5.4

. E).

Tabl

e 5.

3 (c

ontin

ued)

Cre

tace

ous t

o N

eoge

ne e

volu

tion

of th

e flo

ras o

f Sou

th A

mer

ica,

mai

nly

from

Rom

ero

(199

3), H

inoj

osa

& V

illag

rán

(199

7), T

ronc

oso

& R

omer

o (�

998)

, Will

is &

McE

lwai

n (�

00�)

, Hin

ojos

a (�

00�)

.

���

The Neogene evolution model for the Chilean flora (table 5.3, figure 5.4) rests on several tectonic events such as the separation of Antarctica and South America and the consequent opening of the Drake Passage and Antarctic glaciation. The opening of the Drake Passge seems to be a crucial but controversial issue, being dated at �4-�� mya (Kvasov & Verbitski �98�), at ~�0-�� mya (Brown et al. �00�), or ~4� mya (Scher & Martin �00�). These last authors suggest that the 4� mya deepening of the passage coincides with increased biological productivity and abrupt climate reversals. These authors agree with Brown et al. (�00�) that the Drake Passage opened before the Tasmanian Gateway, implying the late Eocene establishment of a complete circum-Antarctic pathway. Other models suggest a much earlier separation of South America from Antarctica, at 140-130 mya (Jokat et al. 2003), or as the TimeTrek model shows (figure 5.1), an early separation at 140-130 mya, a posterior closing till 80 mya, and a definitive separation at ~34, as recognized by Brown et al. (�00�).

Antarctica was covered by forests from the Permian onwards (Taylor et al. �99�). In the Early Cretaceous, the Antarctic forest ecosystem was dominated by a conifer-fern community similar to that in the warm temperate rainforests of present New Zealand (Falcon-Lang et al. �00�). During the Late Cretaceous flowering plants radiated throughout Gondwana changing the vegetation to one more similar to the angiosperm-dominated cool temperate Valdivian rainforests of present-day Chile: Nothofagaceae, Myrtaceae, Eucryphiaceae, Lauraceae, Monimiaceae, Araucariaceae, Cupressaceae, and Podocarpaceae (Poole et al. �00� and references therein). To this impressive record it is possible to add Late Cretaceous fossil flowers related to extant Winteraceae (Eklund �00�).

These Antarctic forests were ultimately eradicated due to the global cooling of climate during the Tertiary (Francis & Poole �00�). The nature and the timing of the extinctions caused by the climate cooling is still being debated due to the paucity of Neogene fossil sites (Ashworth & Cantrill �004). The extinction may have followed the mid-Miocene warm interval at c. �7 Ma or a mid-Pliocene warm interval at about � Ma (Ashworth & Cantrill �004). It has been suggested that, at least in the de-glaciated coastal areas of Antarctica and over the Antarctic Peninsula, a shrub tundra dominated until the Pliocene (Haywood et al. �00�). In southern South America there was a constant exchange of neotropical and Antarctic floras throughout the Cenozoic (Hinojosa & Villagrán 1997, 2005; Troncoso & Romero 1998) (figure 5.4).

5.2 Oceanic transgressions over the continental surfaceRiccardi (�988) proposed several phases of transgression-regression from the Late Jurassic (Kimmeridgian [��� mya]) to the Late Cretaceous (�� mya) that affected different regions in South America. The initial history of Cretaceous sedimentary basins is related to the initial break-up of Gondwana. Along the subduction zone of the Pacific coast, Late Jurassic/Early Cretaceous marine basins were initiated with the development of volcanic arcs and ensialic troughs, in Central Chile, W-central Argentina and S Patagonia. By ��� mya, central-W Argentina and Central and N Chile had gone through a regional uplift that established continental conditions. By ~��0 mya, the

��7

area was again downwarped. A marine back-arc basin extended in a NNW direction, from about 40°S to the latitude of Copiapó and perhaps Antofagasta (figure 5.5). Transgressive-regressive sequences were deposited till ~�40 mya, during a period of rapidly rising sea-level (Mitchum & Uliana �98�, quoted by Riccardi �988). On the contrary, regressive sequences characterize most of the Late Cretaceous, as a consequence of high plate convergence rates associated with a maximum in igneous activity, major uplift, and eastward migration of the magmatic arc (Ramos �98�, as quoted by Riccardi �988). The discrepancies of these trends with the global sea-level changes suggest that the transgressive pattern of the Andean Basin appears to have been controlled by regional tectonics in an area where local vertical movements were greater than global sea-level changes (Riccardi �988).

The transgressions since the Late Cretaceous [8� mya] have been revised by Donato et al. (�00�) in relation to the biogeography of Listroderina (Coleoptera). They suggest that Late Cretaceous-Early Palaeocene transgressions affected mostly northern South America (Colombia, Venezuela) and several regions in Argentina and Bolivia. On the contrary, a Late Oligocene-Early Miocene transgression could have affected south-central Chile between 35°-48° (figure 5.6). The model developed by Hinojosa & Villagrán (�997) considers this wide transgression having affected only E South America. Martínez-Pardo (�990) suggested Central Chile could have been affected by transgressions at �9-�0 mya, in concordance with the global Miocene warming (Zachos et al. �00�).

Figure �.� Late Oligocene-Early Miocene transgressions that possibly affected south-central Chile between ��°-48° (redrawn from Donato et al. �00�)

Figure �.� A marine back-arc basin extended from about 40°S to the latitude of Copiapó/Antofagasta at Early Cretaceous (�4�-��0 mya) (from Riccardi �988).

54°

48°

42°

36°

30°

��8

5.3 The Andean uplift

Stuessy & Taylor (�99�) notice that several authors treat the uplift of the Andes as a recent geological event, but it is a gradual process that has been active for more than �� million years from the Palaeogene (Cenozoic) onwards or even earlier. The supposedly rapid uplift of the Andes has been interpreted by various authors as the main cause of the actual habitat diversity, high levels of endemism and plant diversity in the tropical Andes (Donato et al. �00�, Mutke & Barthlott �00�). Several recent phylogenetic studies further support a direct association between the diversification of Andean plants (e.g. Hughes & Eastwood �00�, Smith & Baum �00�) and the major episodes of Andean uplift, from the early Miocene (about �0 mya) to the Pliocene (about � mya) (sensu Hooghiemstra & van der Hammen �998).

The Andean uplift should have had a great influence on climate alterations which directly influenced: a) adaptive processes, b) plant migrations, and c) speciation and extinction rates (Stuessy & Taylor �99�).

5.3.1 Final uplift of the Andes and Atacama hyperaridity

Earth scientists have attempted to determine the elevation history of the Earth’s surface by differ-ent methods: measuring proxies for barometric pressure, the thickness of overlying atmosphere, the enthalpy of the atmosphere, ground temperature, the d18O value of meteoric water, and pal-aeobotany (Ghosh et al. �00�). Analysing palaeobotanical data, Gregory-Wodzicki (�000) con-cluded that the Altiplano-Puna had attained no more than a third of its modern elevation of �.700 m asl by �0 mya and no more than half its modern elevation by �0.7 Ma. These data imply surface uplift on the order of �.�00–�.400 m asl since the late Miocene at uplift rates of 0.�–0.� mm/yr. Geomorphological evidence, i.e. lahar deposits in the Coastal Cordillera of central Chile (��°40’-�4°��’S) suggest an Oligocene–Miocene uplift of the Andes (Encinas et al. �00�).

Recently Ghosh et al. (�00�) proposed a novel method to calculate the Central Andean uplift, which they called clumped isotope thermometer. They obtained results of surprisingly rapid uplift for the Bolivian Altiplano at an average rate of �.0� +-0.�� mm per year between ~�0.� and ~�.7 mya. These results challenge the known forces responsible for the uplift, i.e. crustal shortening and isostatic compensation of thickened crust that could have led to altitudinal increase of the Altiplano at rates up to only 0.3 mm/year. This is in conflict with much geological evidence (e.g. Hartley �00�, proposed a proto-Central Andean mountain range placed between �� and 9 mya). The rapid uplift model (from 0 to 4000 m asl since the Middle/Late Miocene) proposed by Ghost et al. (�00�) counters former geological evidence and is therefore criticized by Sempere et al. (�00�) (see the response of Eiler et al. �00�).

According to Villagrán & Hinojosa (�997), there should be a synchronous relationship between the increasing aridity of the South American tropics at the end of the Miocene and beginning of the Pliocene, and the cooling of the Antarctic Ocean and the beginning of glaciation in W

��9

Antarctica. Also the final uplift of the Andes during the Plio-Pleistocene would have increased the drying effect of the Humboldt current that is active since the beginning of the Pliocene, which has also to do with the origin of the Atacama desert. Both, the cooling of Antarctica and the uplift of the Andes would have avoided the floristic interchange between the southernmost forests and other plant formations to the north. This interpretation is to some extent congruent with Hartley & Chong (�00�), who suggest that hyperaridity in the Atacama did not commence until the late Pliocene, with the implications that the rain shadow generated by the Andean Cordillera has had a minor influence on climate change, and that the upwelling cold Humboldt Current did not control the shift to hyperaridity. But in general, the exact time at which the climate in the Peru-Chile Desert became hyperarid is a topic of vigorous debate with ages ranging from �� mya (Dunai et al. �00�), �4 mya (Alpers & Brimhall �988) to � mya (Hartley & Chong �00�). Houston & Hartley (�00�), stressed that although the Atacama Desert has existed for at least 90 mya, the initial onset of hyper-aridity was most likely to have developed progressively with the uplift of the Andes as they reached elevations between 1.000 to 2.000 m a.s.l. But the coupling with the intensification of the cold upwelling Peruvian Current is dated by these authors at �� to �0 mya (Middle Miocene). On a recent paper, Hartley et al. (�00�) further propose that “the sedimentary succession in the Atacama Desert records deposition under an arid to semiarid climate from the late Jurassic (��0 mya) to the present day. Palaeomagnetic data indicate no significant latitudinal movement of this area since the late Jurassic. The present-day location of the Atacama within the dry subtropical-tropical transition zone is the principal cause of aridity. This situation is likely to have prevailed since the late Jurassic, supplemented by (�) the continentality effect (enhanced by the Gondwanan landmass), and (�) the presence offshore of a cold, upwelling current (from at least the early Cenozoic onwards and possibly earlier), resulting in conditions promoting climatic stability and desert development. Rapid and extreme climatic fluctuations during the Plio-Pleistocene were not sufficient to destabilize the climate within the Atacama. Comparison with other long-lived deserts (e.g. SW USA, Namib, Sahara and Australia) suggests that the Atacama is the oldest extant desert on Earth”. These results are in agreement with results presented by Envenstar et al. (�00�) of erosional and depositional surfaces analysed via remote sensing. Ochsenius (�999) also has suggested an old age for the Atacama, as one of several Triassic-Jurassic palaeodeserts, prior to the break-up of Gondwana. Ochsenius could further identify Mesozoic relict floras of the intra-Andean valleys from Venezuela to Ecuador, which he described as an Interandean arid track. Dillon et al. (�00�) suggest that western South America has been an arid region well over �� 000 years ago, and analysis of pollen assemblages of rodent middens also suggest that hyperariditiy has prevailed during the Pleistocene-Holocene, the last �0 000 years (Maldonado et al. �00�).

5.4 The last 30 000 years: surviving the Ice Age

The Pleistocene is characterized by several cycles of glaciation/deglaciation. The last glaciation cycle is known as the Llanquihue (LLG) glaciation. At the time of middle Llanquihue glaciation, cool, humid interstades on Isla Grande de Chiloé with Subantarctic Evergreen Forest was increasingly replaced by parkland under progressive cooling after 47,000 [�4]C yr BP (Heusser

��0

et al. �999). A closed-canopy of North Patagonian Evergreen Forest established by ��,�00 [�4]C yr BP. Later, after ca. ��,000 until �0,000 [�4]C yr BP, depending on localion, forest at low elevations became modified by expansion of a cold-tolerant element. Cool, humid interstadial conditions, punctuated by cold stadial climate, are characteristic of the last >40,000 [�4]C years of the Pleistocene at midlatitude in the Southern Hemisphere.

During the tardiglacial, between �9,400 and �4,4�0 [�4]C yr BP, the advance of the glaciers seem to have devasted �/� of the actual area of the southern forests, specially in the regions Aisén, Magallanes, insular and continental Chiloé, and the central and southern Andes (Villagrán & Hinojosa 2005). Periglacial effects like solifluction and glaciofluvial activity should have also affected the Andes, longitudinal depression, and coastal Cordillera between �9°-4�°, affecting

principally the Valdivian forest taxa (Veit �994, Denton et al. 1999, Heusser 2003) (figure 5.7).

The current disjunct range of several species in both cordilleras is a relict of a formerly wider distribution (before the cold period �0.000-�4.000 yBP), as shown by the relict presence of Fitzroya and Pilgerodendron in the central depression, Villagrán et al. �004).

Campos de Hielo Norte and Sur are the last remnants of the Pleistocene glaciations, the biggest inland icecaps after Greenland. The traditional view proposes that taxa mostly survived the glaciations in the foreland of the glaciers and on several nunataks, but this view has been recently challenged by Richter et al. (�004) and Fickert et al. (�007). These authors suggest, based on research on six active glaciers (e.g. Monte Tronador in southern Chile) that the nunatak theory offers just a small area for a survival

of plants, while the size of possible refuges would be considerably enlarged if debris-covered glaciers were considered. This hypothesis is concordant with the results of Premoli et al. (�000), which suggest

that the populations of Fitzroya survived the last glcial maximum in multiple refugia rather than in only one refugium, such as an ice-free area of coastal Chile (Single Refugium hypothesis). Multiple small refuges on the eastern side of the Cordillera are also hypothesized for the survival of Austrocedrus during the last glaciation (Pastorino & Gallo �00�).

Pleistocene and Holocene changes have disrupted species ranges, extirpated local populations, and changed selective pressures (Premoli et al. �000), but it is doubtful that they affected speciation

54°

48°

42°

36°

Figure �.7 Maximal extension of the last cycle of the Llanquihue glaciation (after Denton et al. �999, Heusser �00�).

���

processes. Some authors have emphazised the role of last glaciations in speciation, but others call this a ‘failed paradigm’ (e.g. Klicka & Zink �997). Continuing with Fitzroya, ist phylogenetic position, together with the one of Pilgerondenron have long been considered as enigmatic (e.g. Page 1990), but more recently Quinn & Price (2003) suggest that Fitzroya is closer to Diselma (from Tasmania) and Widdringtonia (from South Africa). Pilgerondenron appears phylogenetically nested within australasiatic Libocedrus, closer to L. yateensis of New Caledonia.

5.5 Alternative palaeogeographies: against scientific consensus

The palaeogeographic scenarios presented in the previous sections arise more or less from the scientific consensus, under the current geologic/tectonic paradigm. In spite of having travelled dusty waters since the first proposal of the Kontinentalverschiebungs- Theorie by Alfred Wegener (1915), the theory settled finally in the scientific community as more or less trustable truth. The theory was re-discovered by the geological community when the mechanisms by which the continents would move over the earth surface were found (e.g. Hess �9��). Now the theory of mantle plumes�� (figure 5.8) as the mechanism underlying the plate movements has settled as standard in tectonics (e.g. Storey 1995). But the theory is not free from conflict. Foulger et al. (�00�) wrote “If the plume hypothesis is abandoned, this will arguably represent the most significant paradigm shift in Earth science since the advent of plate tectonics”. Authors in the volume concentrate the discussion on the mechanisms: e.g. the nature of the driving forces, the depths of recycling, the sources of melting, and the fundamental question whether mantle plumes exist. “Without plumes, plate tectonics loses a number of capabilities. The most important is the ability to rift continents. The uplift, thinning, and dragging apart of the lithosphere as the plume head strikes will be hard for plate tectonics to replace. Rifting continents join the initiation of subduction zones far from continents, mountain building, and the sudden redirection of the Pacific plate as the primary mechanical problems of the theory” (Fischer �00�)��.

The assessment of Pratt (�000) points in the same direction: “Plate tectonics -the reigning paradigm in the earth sciences- faces some very severe and apparently fatal problems. Far from being a simple, elegant, all-embracing global theory, it is confronted with a multitude of observational anomalies and has had to be patched up with a complex variety of ad hoc modifications and auxiliary hypotheses. The existence of deep continental roots and the absence of a continuous, global

asthenosphere to lubricate plate motions have rendered the classical model of plate

Figure �.8 Mantle plumes, www.seismo.unr.edu/ftp/pub/louie/class/plate/; www.seismo.unr.edu/.../class/�00/interior.html

���

movements untenable. There is no consensus on the thickness of the plates and no certainty as to the forces responsible for their supposed movement. The hypotheses of largescale continental movements, seafloor spreading, and subduction, as well as the relative youth of the oceanic crust are contradicted by a substantial volume of data. Evidence for significant amounts of submerged continental crust in the present-day oceans provides another major challenge to plate tectonics. The fundamental principles of plate tectonics therefore require critical reexamination, revision, or rejection” �4. Dickins et al. (�99�) wrote in the same sense: “After surveying the extensive evidence for former continental land areas in the present oceans, we are surprised and concerned for the objectivity and honesty of science that such data can be overlooked or ignored. There is a vast need for future Ocean Drilling Program initiatives to drill below the base of the basaltic ocean floor crust to confirm the real composition of what is currently designated oceanic crust”. Hamilton (�00�) further proposed that current paradigms in tectonics are more tied to the modes of research and publication in the geological community rather than to empirical evidence.

Other models have been proposed, aside the mainstream geological community, such as the alternative models of Dobson (�99�). His circulation model “differs from previous models in proposing that convection drives circular plate motion while gravity drives lateral motion. Convection currents upwell at high-pressure centres, spiral outward, transfer to low-pressure cells, spiral inward, and descend at low-pressure centres. In most instances, upwelling and descent of asthenosphere occur at opposing plate centres rather than at plate margins. Cells migrate laterally in a global pattern driven by gravity. Sea-floor spreading and subduction occur because of differential rates of lateral plate motion.” (Dobson �99�, p. �0�). The palaeogeographic reconstruction of Dobson (1992, p. 200) (reproduced here as figure 5.9) looks perfectly suitable for being able to explain the close floristic relationship between Central Chile and California, because both territories appear together.

Tiny motion in the crust, measured in centimeters per year and usually attributed to plate tectonics at work, may instead be the result of differential rotation between the lithosphere and mantle as proposed by Smith & Lewis (�999). Cretaceous has also been recently challenged due to biogeographical and geological reasons. “Palaeomaps, it seems, are like the weather: If you

Figure �.9 Alternative palaeoreconstructions (Dobson �99�)

���

don’t like the alleged size or placement of a pre-Cenozoic ocean, just wait a while. It will change” (McCarthy �00� b).

Land bridgesThe idea of former land bridges in the site of today’s oceans, contrary to the current paradigm of plate tectonics, is not an old one, and like a circular idea, is constantly re-presented and re-rejected. Probably the first one who proposed and illustrated former lands between today’s continents was Hermann von Ihering (�907), precisely while studying the biogeographical connections of the Neotropical biota. He proposed ancient land connections between America and Africa and between Antarctica and Australasia, that he called Archinotis and Archhelenis respectively (figure �.�0). Also Croizat (�9��) proposed vast emerged lands between the shapes of current continents (figure 5.11). The theory of land bridges was extendedly defended by G. van Steenis (1962) in The theory of Land Bridges in Botany. The idea of the land bridges have been recently called again by several authors for explaining the biogeographical relationships between the American and African tropics (Morley �00�), the pantropics (Zhou et al. �00�) and between Tertiary disjunct relict floras from Eurasia – North America (Milne 2006).

Figure �.�0 Land bridges proposed by H. von Ihering (�907)

��4

Pacifica continentAs discussed in section 4.5.3, the idea of a Pacifica continent West of South America is an old idea. During the 80’s, under the umbrella of vicariance biogeography the idea was revitalized (Nur & Ben-Avraham 1977, 1981a; Kamp 1980). The hypothesis was firmly rejected by several

Figure �.�� Land bridges as proposed by Leon Croizat (�9�8)

Figure 5.12 Former land areas in the present Pacific and Indian Oceans (from Dickins et al. 1992, 1996)

���

authors like Cox (�990). Dickins et al. (�99�) also propose different emerged lands around the Pacific (figure 5.12). The differences between regional palaeoreconstructions and global ones are notably, especially the reconstructions based on accreted terranes and the reconstructions that place Laurentia very far from Gondwana during the Middle Ordovician.

It is true that theory-free descriptions of natural history phenomena have little scientific impact (Hull �988, as quoted by Grande �994). If we remove the Pacifica hypothesis we are left to choose another causal explanation for the repeating pattern of area (floristic) relationships, and other parts of the scenario remain intact. Even if we remove some taxa, we do not change anything significant about the whole picture. But if we remove the initial transpacific area pattern, everything collapses: “There is no repeating pattern from wich to extrapolate and for which to build complex scenarios” (Grande �994, p. 7�).

Expanding earthThe most recent recycled hypothesis that could perfectly explain the biogeographic relationships between southern Chile and Australasia is the expanding earth hypothesis (figure 5.13). First proposed by Lindemann (�9�7) and Hilgenberg (�9��), the theory loosed attention during the �0 century, in spite of being still defended by some researchers (e.g. Carey �988, Vogel �990). The theory was reanalysed specifically from a biogeographical point of view by Shields (1997, 1998) and more recently by McCarthy (�00�, �00�a). It has been very criticised by biogeographers like Cox (�990) and Briggs (�004). Mainstream textbooks treat the idea in different forms: Cox and Moore (�00�), being consequent with Cox (�990), do not mention the hypothesis in their Biogeography. But Lomolino et al. (�00�) dedicate two pages to discuss the pros and cons of the expanding Earth hypothesis (Lomolino et al. �00�, pp. �48-�49). The debate is far from being finished, and the last word stays so far in McCarthy (2007), also gaining the cover of the biogeography book recently edited by Ebach & Tangney (�007).

Figure �.�� Expanding Earth hypothesis and a close Pacific ocean (after Shields 1998, Mc-Carthy �00�). See also http://www.expanding-earth.org/

���

Bonus track: modeling expanding earth, Paola pregnant on ��.��.�004 (top); same on ��.0�.�007 (bottom)

��7

� Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography

��8

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

��9

6 Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography

“...Neither of the authors of TNT, nor the distributor, are responsible in any way for any problems the program causes to your computer, your data, your career, or your life”…………. Warning/Disclaimer from the distributors of the cladistic program TNT [http://www.cladistics.com/aboutTNT.html]

6.1 Molecular dating, the reigning paradigmThe program of molecular dating of phylogenies has becoming very popular due to the availability of better resolved phylogenies in combination with new methods for estimating divergence times

(see Sanderson �997, Huelsenbeck et al. �000, Britton et al. �00�, Sanderson �00�, Thorne & Kishino �00�). There has been also an increasing number of taxonomically identifiable fossils (Friis et al. �00�). In spite of the datings have given widely different results, more recent studies tend to converge on similar ages (Sanderson et al. �004, Bell & Donoghue �00�). Especially

datings of the lower nodes within the angiosperms, have given much older ages than those obtained from the fossil record (e.g. Wikström et al. �00�).

Molecular dating methods currently in use are classified by Rutschmann (2006) in three main classes:

�. methods using a molecular clock and one global rate of substitution (e.g. linear regression, character-based maximum likelihood clock optimization);

�. methods that correct for rate heterogeneity (e.g. linearized trees, local rates methods); �. methods that try to incorporate rate heterogeneity (e.g. heuristic rate smoothing,

penalized likelihood). Since variation in rates of nucleotide substitution along a lineage and between different lineages is now known to be pervasive, the clock model of molecular evolution have been changed to the relaxed or sloppy clock, that try to address the deviations from the clock-like model (Rutschmann �00�).

Milne (2006) described a simplified three-stages procedure from most modern molecular dating studies, reflecting the interaction from the fields of molecular systematics, mathematics and palaeobotany. These three stages are:

�. a phylogenetic hypothesis is generated, by use of data from one or more DNA markers. Every branch in a phylogeny has a length measured in number of substitutions for the sequences examined. It is not possible to convert branch length data directly into an estimate of time because substitution rates vary between lineages, a consequence of rate heterogeneity [Welch & Brohman �00�];

�40

�. substitution numbers on branches are converted into measures of relative time within the phylogeny, technique known as rate smoothing [Sanderson et al. �004];

�. conversion of relative ages into actual ages, assigning an actual age to at least one node of the phylogeny [e.g. Wikström et al. �00�]. Nodes assigned actual ages are known as calibration points, and the method normally used is dated fossils. The minimum are of that fossil is determined from its stratigraphic position, and that age is assigned to the node. Then, the minimum age of every other node is calculated as a proportion of that of the calibration node.

The molecular dating program is not free of uncertainty. Anderson et al. (�00�), after discussing the difficulties and uncertainties in obtaining a stem group age for the eudicots, finished their paper asking if the results could not be rather an artifact from constraints or method: “All our methods rely on the assumption that there is an autocorrelation of evolution rates in adjacent lineages and that some kind of smoothing is reasonable, but could it be that rates change abruptly rather than gradually? This has been suggested by Sanderson & Doyle [�00�]. In palaeozoology there has been more debate on the rate of evolution. It has been proposed that events like the Cretaceous-Tertiary boundary mass extinction and the Cambrian Explosion could be artifacts of the rock record [e.g. Benton & Ayala �00�, and references therein]. Other scientists suggest that these events are real and that we should trust the fossil record [Benton et al. �000]”.

This seems to be a matter of trusting and believing, and exemplified the still weak knowledge about evolution rates and modes of speciation. Anderson et al. (�00�) concluded questioning if dating could be done without a [molecular] clock: ”to estimate divergence times consistently, we need precise fossil dates on all nodes where rate changes occur. Of course, this is not realistic, but in any case we should avoid to calibrate with only one or a few fossils, as is often done. If we had many fossils placed more or less even over the tree the choice of method would be less important. We need more fossils and these fossils must be well dated and as widely dispersed across the phylogenetic tree as possible. Excluding any of the fossils in this study, clearly gave younger ages for the clade to which the fossil was attached“.

“Molecular dating may be affected by a variety of error sources the majority of which are not sufficiently theoretically understood” (Sanderson & Doyle 2001). These authors pointed out that nonclocklike behaviour of evolutionary rates might lead to significant deviation among results obtained with different dating methods. Different methods may introduce systematic biases, which are generally hard to detect. “If our finding that increased sampling leads to older age estimates is corroborated in the future, then current dating methods need revision”. Indeed, Heads (�00�b) did a deep revision of the whole molecular biogeographic program, and is one of the most downloaded paper from the journal Cladistics. But in spite of being vastly cited, Heads’s concerns are not yet seriously incorporated in most recent papers (Heads pers. comm.)

“Ultimately, we may well conclude that accurate divergence time estimates require multiple

�4�

reliable calibrations. That is too bad, if true, but it may be true. Until this largely empirical question is resolved, it may be desirable to evaluate existing and new methodologies in the context of a few carefully chosen systems that offer numerous fossil calibrations. If methods can be fine-tuned to succeed in problems where cross-validation is feasible, then there may be some hope to extend them to more difficult problems with fewer available fossil calibrations. Ironically, the systems in which divergence time estimation from sequence data is needed most critically are the ones with few or no good calibrations… Perhaps we should learn to walk in the context of these systems before learning to run in the real world” (Near & Sanderson �004).

In a recent opinion article Pulquério & Nichols (�007) ask “how wrong can we be?” regards to the application of molecular clocks. The authors analyze current promising approaches to solve the question of the uncertainty in the dates attributed to calibration points; e.g. Drummond et al. (�00�) propose a method that do not impose unproven assumptions about the pattern in clock-rate variation among lineages. The answer to Pulquério & Nichols’ initial question but still seems to be “we do not yet know”, and they further concluded “We await the more rigorous type of assessment with some nervousness, given that we suspect they might reveal that many past studies placed too much confidence in simple molecular clock analyses, and that their conclusions should thus be revisited” (Pulquério & Nichols �007).

6.2. Phylogeny of Chilean plants

The evolutionary history of plant life, also known as the plant tree of life is a major endeavour, based on the impressive progress made over the last �0 years or so of the DNA revolution (Palmer et al. �004). Today, “nothing in biology makes sense except in light of phylogeny” (Dobzhansky 1973, quoted by Palmer et al. 2004). In spite of conflicts and different points of view (sections 1.3, 6.1), Palmer et al. (2004) are confident to say that we are obtaining a comprehensive, robustly resolved, and accurately dated plant tree of life. Under this scenario and considering the cladograms and chronograms published by APG II (�00�), Palmer et al. (�004), Pryer et al. (�004), Davies et al. (�004), and Stevens (�00� onwards), a synthesis of the phylogeny of the Chilean vascular plants can be intended.

Lycophytes are the most primitive plants in the Chilean extant vascular flora (= Isoetes, Huperzia, Lycopodium). This group has been hypothesised as splitting from the euphyllophytes in the early-mid Devonian [ca. 400 mya] (Pryer et al. �004; Palmer et al. �004). Living euphyllophytes belong to two major clades (figure 6.1): seed plants (spermatophytes) and monilophytes (Kenrick & Crane �997, Nickrent et al. �000, Pryer et al. �004).

Monilophytes, ferns s. str., include most former groups recognized as ‘fern and fern allies’. Monilophytes have been dated back to the Late Devonian [ca. �70 mya]. Chilean representatives are classified in 21 families and 47 genera (appendix A).

�4�

The seed plants (spermatophytes) are the most diverse vascular plant group in the world and in Chile. Spermatophytes comprise cycads, ginkgos, conifers, Gnetales, and angiosperms. Extant seed plants likely number between ��0.000 and �00.000 species (Thorne �00�; but see Scotland & Wortley �00�). In Chile the seed plants are represented by 41 orders, 154 families and 763 genera (table �.�, appendix A). The extant seed plants have been shown to be a monophyletic group; that is, the entire group arose from a single common ancestor, with initial radiation on the Late Palaeozoic [ca. �00 mya] (Stewart and Rothwell �99�, Hill �00�). However, exact relationships among these lineages and the pattern and chronology of divergence remain unclear, despite the recent accumulation of molecular data sets to address the question (e.g. Magallón & Sanderson �00�, Burleigh & Mathews �004). The earliest known seed plants have been reported from the Late Devonian of West Virginia (Gillespie et al. �98�). Gnetales and modern conifer families

appeared in the Triassic to Jurassic, and angiosperms in the limit Jurassic/Cretaceous (Stewart and Rothwell �99�, Crane �99�). From the Permian through the late Jurassic many seed plant lineages

went extinct, including lyginopterids, medullosans, Callistophytaceae, glossopterids, Cordaitales, and Voltziales (Stewart & Rothwell �99�), and their relationships with extant groups remain poorly resolved. During the Cretaceous and Cenozoic, the diversity of all surviving seed plant lines except angiosperms decreased (Knoll �984, Crane �98�).

The Chilean flora is lacking extant cycads or Ginkgoales, but several fossil taxa have been described (Troncoso & Herbst �999, Herbst & Troncoso �000, Torres & Philippe �00�, Leppe & Moisan �00�, Herbst et al. �00�, Nielsen �00�). Conifers are represented by one order, � families, and 8 genera. Chilean Gnetales are represented by one order, family and genus (Ephedra).

Angiosperms are the most intensively studied group, that also offers the benefits of being relatively young and species rich (Palmer et al 2004), with at least 260 000 living species classified in 453 families (APG II �00�). The fossil record of the angiosperms extends back at least to the early Cretaceous, conservatively ��0 mya (Crane et al. �004). Molecular dating has but pushed this age to [�79-��8 mya] the Early – Middle Jurassic (Wikström et al. �00�). The most basal Chilean clades in the angiosperm tree are the following (proposed ages by Wikström et al. 2001) (figure 6.1):- Ceratophyllum [��� – �40 Mya] Late Jurassic, limit Jurassic/Cretaceous.- Monocots [��4 - ��9 Mya] Late Jurassic, limit Jurassic/Cretaceous. - Piperales [�49 - ��7 –Mya] Late Jurassic, limit Jurassic/Cretaceous, Piperaceae [9� – 90 Mya] Late Cretaceous; Lactoris/Aristolochia [97 – 8� Mya] Late Cretaceous; Peperomia/Piper [4� – 4� Mya] Eocene. - Laurales [�4�-��� Mya] Early Cretaceous, Monimiaceae splitting at 9�-8� Mya (Late Cretaceous), Lauraceae splitting from Gomortegaceae/Atherospermataceae at 87-80 Mya (Late Cretaceous), Peumus/Hedycarya [78 – 7� Mya] Late Cretaceous, Gomortega/Laurelia [��-�9 Mya] Eocene. Winterales (alt. Canellales) split from Magnoliales at ���- ��7 Mya (Early Cretaceous), Winteraceae�� from Canellaceae at 99 Mya (limit Early/Late Cretaceous), Drimys/Belliolium [�4-�9 Mya] Oligocene.

�4�

Figure �.� Angiosperm chro-nogram calibrated against the geological time-scale (adapted from Wikström et al. �00�). In grey taxa not represented in Chile.

Amborellaceae

Nymphaeaceae

Schisandraceae

Illiciaceae

Austrobaileyaceae

Chlorantales

Piperales

Laurales

Winterales

Magnoliales

Ceratophyllum

Ranunculales

ProtealesSabiaceae

Buxaceae

Trochodendraceae

Gunnerales

Santalales

Dilleniaceae

Caryphyllales

Cornales

Ericales

Garryales

Oncotheca

Gentianales

Solanales

Lamiales

Aquifoliales

Asterales

Apiales

Escallonia

Eremosyce

Berberidopsidales

Saxifragales

Geraniales

Crossomatales

Ixerba

Aphloia

Tapiscia

Brassicales

Malvales

Sapindales

Zygophyllales

Rosales

Fagales

Celastrales

Oxalidales

Malpighiales

Fabales

eu

rosid

s Ie

uro

sids II

eu

rosid

s

eu

dic

ots

as

te

rid

s

eu

as

te

rid

s II

eu

as

te

rid

s I

ma

gn

oliid

s

eu

ma

gn

oliid

s

180 160 140 120 100 80 60 40 20 P

180 160 140 120 100 80 60 40 20 P

Early Cretaceous Late Cretaceous Paleogene NeogeneJurassic

Monocots

Dipsacales

Vitaceae

Cucurbitales

Myrtales

�44

Within the Monocots, results from Janssen & Bremer (�004) suggest that considerable monocot diversification took place during the Early Cretaceous, with most families already present at the Cretaceous–Paleogene boundary. The crown nodes of Araceae, Petrosaviaceae, Orchidaceae, Arecaceae and Dasypogonaceae have been estimated to date back to the Early Cretaceous (Janssen & Bremer �004). Liliales diverged from their sister –group in the Early Cretaceous, ��7 mya and the extant lineages diverged from each other ca. 8� mya; Poales at ��� +- �� mya (Mid-Cretaceous) (Bremer �000). Bromeliaceae also appear to date back to the Cretaceous (Linder & Rudall �00�), but the uncertainties in dating the Bromeliaceae are considerable (Bremer �00�). Janssen & Bremer (�004) found an unexpected crown node age of ��� mya for Orchidaceae. “Traditionally, this family has been looked at as a very specialized and hence, probably, a young group. Considering the extensive sampling within orchids (145 genera) and its firm phylogenetic position at the base of the Asparagales, this age estimate appears to be well supported. However, a methodological bias due to the extended sampling still cannot be excluded. If our age estimate turns out to be true, the evolutionary history of this family could be seen in a new light. Orchid diversity is not necessarily due to a rapid and recent radiation, and similar patterns, for example in palms, might be hypothesized” (Janssen & Bremer �004).

Basal Eudicots have been recently dated back to the Early Cretaceous: Ranunculales estimated at ���-�47 (Wikström et al. �00�) or at ��0 mya (Anderson et al. �00�), Proteales at �44-��0 mya (Wikström et al. �00�) or at ��9 mya (Anderson et al. �00�).

Core Eudictos: Gunnerales has been dated at ��7-��� mya, Santalales at ��8-��� mya, Saxifragales at ���-��� mya, Caryophyllales at ���-�04. A recent recognized order is Berberidopsidales, composed by two monotypic families from Central Chile/Australia (Berberidopsidaceae) and Central Chile/Argentina (Aextoxicaceae). Berberidopsidales have been dated back to ��4-��4 mya (Wikström et al. �00�), the split between Berberidopsis and Aextoxicon at �00-90 mya (limit Early/Late Cretaceous).

Rosids have been dated back at �09-�00 mya, eurosids I at �0�-9� mya, eurosids II at �09-�00 mya.

For asterids, Wikström et al (�00�) obtained consistently younger age estimates, mostly to �0 to �0 my for major groups and orders compared to estimates by Bremer et al. (�004). As example, the crown node age in the analysis of Bremer et al. (�004), is ��8 my whereas Wikström et al. (�00�) estimated it to ��7 to �07 my. The crown node age of Asterales was estimated to 9� my by Bremer & Gustafsson (�997) using rbcL sequences and calibration with fossil Asteraceae pollen, and to 9� my (Bremer et al. �004). The order comprises the Asteraceae and Campanulaceae, and several small families mainly from the southern hemisphere that supposedly originated during break-up of Gondwana in the Late Cretaceous and Early Palaeogene (Bremer & Gustafsson �997).

�4�

6.3 Vicariance v/s dispersal in the Chilean FloraAs already mentioned in section 4.�, disjunct distributions has been interpreted under two highly conflicting views: vicariance v/s dispersal. During the 1980s and 1990s, under the paradigm of vicariance (cladistic) biogeography (Nelson and Platnick �98�, Humphries & Parenti �999), biogeographers inclined toward the idea that plant disjunctions resulted from the fragmentation of earlier, larger landmasses, such as Gondwana. These vicariance explanations remained dominant until the recent advent of molecular systematic techniques, particularly molecular-based dating of lineage divergences (de Queiroz 2005). In Moore et al.’s (2006) opinion “… using these techniques, much recent scholarship has demonstrated that numerous plant disjunctions are far too young to have resulted from vicariance, leaving transoceanic dispersal as the only plausible alternative [...] The realization that long-distance dispersal may have been far more frequent than previously supposed has led plant biogeographers using modern molecular tools to reexamine the relative importance of vicariance and dispersal in explaining the classic patterns of worldwide plant disjunction”. Moore et al. (�00�) consequently found that the disjunct distribution of extant species of Tiquilia is the result of at least four long-distance dispersal events from North America to South America. Taken the dispersalist universe as framework, Moore et al. (�00�) do not mention alternative palaeogeographic hypothesis, as listed by Constance (�9��). But taken seriously the concerns already expresed in section �.�, some researchers are more cautious and conclude: “Given the inherent methodological problems, absolute age of clade divergences, relevant as evidence of long distance dispersal or vicariance, cannot yet be determined with confidence” (Ladiges et al. �00�).

Recent geophysical research seems to support geographical scenarios suited for long-distance dispersal upon the southern seas (Muñoz et al. �004). For the authors, wind connectivity explains the biogeographical similarities in the southern hemisphere for groups known as good dispersers, like lichens, mosses, and ferns. They suggest that this could apply as well for angiosperms with little propagules like orchids. But there are only four fern genera composing the australasiatic element (Rumohra, Arthropteris, Dicksonia and Doodia), and only Rumohra reaches the continent, being the others only distributed in the Pacific islands. There are certainly some circumaustral taxa at the species level (e.g. Shizaea fistulosa, Hymenophyllum ferrugineum). But at the genus level only Rumohra is strictly circumaustral and most of the fern genera have a subcosmopolitan (e.g. Asplenium, Blechnum, Equisetum) or pantropical distribution (e.g. Gleichenia, Hypolepis, Hymenophyllum). Similarity in the relatively old fern group may actually be the result of ancient vicariant events (see a discussion in Wolf et al. �00�). For the orchids, described as good dispersers, there is no one single genus classified as australasiatic, challenging the supposed good dispersal ability of the family due to their very tiny seeds. Van Steenis argued at this respect: “Although dispersal of orchids may seem easy by the large amount of dust seed, successful establishment may depend on presence of its mycorrhizal fungus and insects for pollination. That the three of them, fungus spores, seeds, and insects, will travel together over long distances by chance is utterly unlikely” (van Steenis 1962). Even genera that superficially appear like good dispersers, like the ones pertaining to the Asteraceae, are not necessarily good disperses. Of the

�4�

four Asteraceae genera classified as australasiatic, three lack a pappus suited for wind dispersal (Abrotanella, Lagenophora and Leptinella). Only Trichocline, with its pappus of many scabrid bristles, could be suited for long-distance dispersal. From a dispersalist point of view one would expect exactly the contrary. This is why Heads (�999) challenged the hypothesized long-distance dispersal explanation proposed by Swenson & Bremer (�997) for Abrotanella (see also Wagstaff et al. �00�).

S. Cain already warned us about the error in convenient dispersal stories: “Long-distance dispersal operates for some organisms, and it is especially applicable to littoral species and a portion of the biota of oceanic islands. The hypothesis, however, is much too widely used; in most cases of wide disjunction, a careful investigation shows that the dispersal mechanisms, agents, and establishment requirements of the species rule out this explanation. All too frequently the assumption of long-distance dispersal is merely a careless and easy way out of a difficult problems and it leads to fanciful and even ridiculous conclusions” (Cain �944, p. �0�-�0�).

But researchers tend to ignore or minimize empirical evidence: “Hoffmannseggia fruits and seeds have no obvious adaptations for external animal dispersal and no one has ever recorded their being eaten by birds. Nevertheless, we believe that bird dispersal is the most likely explanation for the repeated pattern of long-distance dispersal from South to North America” (Simpson et al. �00�).

“Hätten die Wandervögel die ihnen oft zugeschriebene Bedeutung für Verbreitung der Pflanzen, so würden die Zugstraßen der Vögel als deren Fäcalstraßen sich floristisch ebenso darstellen, wie etwa die prähistorischen Handelstraßen aus den Fundstücken kartographisch rekonstruierbar sind. Von dem ist aber keine Rede.“ (von Ihering

�89�).

Michaux (2001) goes beyond arguing that the opposition of vicariance versus dispersal is an artifice of poorly defined concepts, and suggest in place of this simplified opposition the recognisance of five processes – modification, movement, mixing, splitting and juxtaposition – that are not logically equivalent as they operate at different time scales. As summarized in table �.�, the processes operating to shape current disjunct patterns might rather be a couple of dispersal and vicariance processes operating at different time scales (see also Sluys �994). We could add that these processes also vary at the taxonomic level under analysis. Vicariance logically predominate as explanation at the order and family levels (e.g. for Apocynaceae, Proteaceae, Apiales, Poales), and LDD tend to dominate at the genus level. But note both conflicting views in explaining the disjunct distribution in Abrotanella, Nothofagus, and Microseris.

�47

Table �.� Evolutionary processes (dispersal v/s vicariance) proposed for several Chilean taxa

Families or groups Genera or group Evolutionary process Sources

Apocynaceae - Gondwanan origin for the family Potgieter & Albert �00�

Apiaceae Oreomyrrhis Origin in Eurasia and subsequently dispersal to North America and southern Pacific Rim.

Chung et al. �00�

Araliaceae Pseudopanax Gondwanan vicariance between Australasia-South America and LDD to Hawaii

Mitchell & Wagstaff �000

Araucariaceae - Gondwanan origin Setoguchi et al. �998Asteraceae Microseris Colonization Aus from N Am Vijverberg et al. �999

Asteraceae Microseris Vicariance, widespread ancestor Grehan �007

Asteraceae Abrotanella Dispersal between Australasia – South America

Swenson & Bremer �997, Wagstaff et al. �00�

Asteraceae Abrotanella Vicariance Heads �999

Asteraceae Hypochaeris Dispersal from NW Africa across the Atlantic Ocean for the origin of the South American taxa

Tremetsberger et al. �00�

Atherospermataceae Laurelia Initial diversification at 100–140 mya, probably in West Gondwana, early entry into Antarctica, and long-distance dispersal to New Zealand and New Caledonia

Renner et al. �000

Berberidaceae Berberis South America-Old World disjunct distribution due to Cretaceous vicariance

Kim et al. �004

Boraginaceae Tiquilia three independent dispersals from North to South American

Moore et al. �00�

Brassicaceae Cardamine dispersal from South America to Australasia, or vice versa

Bleeker et al. �00�a

Brassicaceae Rorippa LDD via migrating birds explains the amphitropical disjunction between South American R. philippiana and North American R. curvisiliqua.

Bleeker et al. �00�b

Brassicaceae Lepidium Probably long-distance dispersal from western North America to South America by birds in the Pleistocene

Mummenhoff et al. �00�

Cunoniaceae - Gondwanan ancestry Bradford & Barnes �00�Cyperaceae Oreobolus LDD Autralasia to South America Chacón et al. �00�Ericaceae - Laurasian in origin.with following

dispersalsKron & Luteyn �00�

Fabaceae Hoffmannseggia Dispersals between California and Central Chile

Simpson et al. �00�

Fabaceae Sophora Origin in North hemisphere and dispersals to the South and the Pacific

Hurr et al. �999, Peña et al. �000

Gentianaceae Gentianella Dispersal from South America into Aus+NZ less than �,7 mya

Von Hagen & Kadereit �00�

Gesneriaceae Coronantheroid Gondwanic origin, Coronantheroid Gesneriaceae (Asteranthera, Mitraria, Sarmienta) relicts

Burtt �998

Gunneraceae Gunnera Early vicariance followed by recent dispersals

Wanntrop & Wanntrop �00�

Gunneraceae Gunnera Late Cretaceous vicariance Fuller & Hickey �00�

�48

Lauraceae Beilschmiedia/Cryptocarya

Gondwanan vicariance (diverged from ancestor about 90+- �0 mya.

Chanderbali et al. �00�

Lycopodiaceae - Vicariance, Permian origin of the genera Wikström & Kenrick �00�

Monimiaceae Peumus Vicariance at 7� mya, former continuous range from Chile across Antarctica and the Kerguelen plateau toMadagascar

Renner �004

Monocots (major groups)

- Gondwanan vicariance Bremer & Janssen �00�

Nothofagaceae Nothofagus Gondwanan vicariance

vicariant main massings of the four subgenera compatible with allopatric differentiation and no substantial dispersal

Swenson et. al �00�,

Heads �00�

Nothofagaceae Nothofagus Vicariance Am-Aus subgen Fuscospora, dispersal in Lophozonia

Knapp et al. �00�, Cook & Crisp �00�

Plantaginaceae Hebe at least two instances of transoceanic LDD from Australasia to South America.

Godley �9�7, Wagstaff & Garnock-Jones �998

Plantaginaceae Ourisia origin in the central Chilean Andes followed by consecutive dispersal to the southern Andes and dispersal(s) to the northern Andes and Australasia

Meudt & Simpson �00�

Poales - Gondwanan vicariance Bremer �00�

Polemoniaceae Gilia Dispersal California to South America Morrell et al. �000

Portulacaceae American origin and dispersals into Australia

Applequist & Wallace �00�

Proteaceae - Gondwanan; Mid-Cretaceous divergence between major groups

Hoot & Douglas �998

Primulaceae Primula Ancestral haplotype gave origin to S Am, Euro and N Am lineages.

Guggisberg et al. �00�

Ranunculaceae Caltha Northern hemisphere origin, followed by dispersal to the Southern Hemisphere (Gondwanaland). Vicariance is invoked to explain present-day distributions in South America, Australia, and New Zealand.

Schuettpelz & Hoot �004

Restionaceae Apodasmia Gondwanic origin followed by dispersals Linder et al. �00�

Rhamnaceae Discaria Probably Gondwanan relict Richardson et al. �000

Rubiaceae Coprosma Vicariance Heads �99�

Rubiaceae Coprosma Africa as ancestral area followed by long-distance dispersal into the Pacific

Anderson et al. �00�

Tetrachondraceae Tetrachondra LDD New Zealand – South America Wagstaff et al. �000

Winteraceae - migration from tropical Gondwana to Antarctica and Australia throughout South America

Barreda & Archangelsky �00�

Zygophyllaceae Fagonia dispersals between South America and southern Africa

Beier et al. �004

Table �.� (continuation)

�49

As revised in section 6.2, the ancestors of extant Chilean flora should have been much more widespread in Gondwana -- as is indicated by the Antarctic palaeofloras (Poole et al. 2003). Indeed, many of the elements found today in the southern continents can be traced to the Gondwana era, as a once-continuous cool-temperate flora, now scattered into a relict distribution by tectonic movements. Furthermore, some authors like to speak of the austral floras as a Gondwanan element (e.g. Barlow �98�, Nelson �98�, Hinojosa �00�).

New analytical tools like DIVA analysis or parsimony-based tree fitting have the supposed capacity to discriminate between vicariance and dispersal (e.g. Sanmartín & Ronquist �004). Recently Sanmartín et al. (�007) tested the directional dispersal in the Southern Hemisphere using event-based tree fitting. The direction of circumpolar currents predicts predominantly eastward dispersal from New Zealand to South America. But contrary to the expectations, dispersal between New Zealand and South America was more frequently inferred to be westward. The authors gave two possible explanations to their failure to detect significant patterns:

a) insufficient sample size of phylogenies that include South American taxa;b) dispersal is as likely to be eastward as it is to be westward for New Zealand–South

American events.The authors concluded that “Only once we have a better understand of dispersal process will it be possible to apply realistic estimates of dispersal frequency and asymmetry to biogeographical reconstructions“ (Sanmartín et al. �007). But this seems also valid for a better understanding of vicariance processes, that frequently have been oversimplified as a several steps break-up model, obviating the complex geological and biotic nature of a region (see next section).

6.4 Sloppy biogeography v/s harsh geology?

“Las hipótesis de relaciones entre áreas formuladas por la geología no tienen validez intrínseca superior a las que se formulen a partir de datos biológicos, es decir, de los cladogramas de áreas. A partir de este principio, de por sí totalmente aceptable, algunos autores concluyen que las hipótesis geológicas no pueden utilizarse ni para corroborar ni para refutar las hipótesis biogeográficas, en contradicción patente con uno de los principios básicos del cladovicariancismo: la idea del paralelismo entre la evolución de la vida y la de la Tierra” (Zunino & Zullini �00�, p. �90).

Recently some authors call for avoiding the circular logic in confronting biogeographical hypothesis with geological evidence (e.g. Renner �00�, Waters & Craw �00�). “Constraining nodes in a phylogenetic tree by geological events risks circularity in biogeographical analyses because it already assumes that those events caused the divergence, rather than testing temporal coincidence” (Renner �00�, p. ���).

The problem seems to be: how much geological evidence is necessary to accept a biogeographical hypothesis?

��0

In the words of Heads (�00�b): “assuming a priori that any particular geological event, such as the break-up of Gondwana, is relevant to biogeography is a fatal flaw of much biogeography, both dispersalist and vicariance… In fact, a great deal of evidence suggests biogeographic patterns involving New Zealand, New Guinea, New Caledonia etc. were determined by earth history events both prior and subsequent to the break-up of Gondwana”. Croizat (1958) was the first to call for this independent view of biogeography as a discipline with own methods and tools. He proposed that biogeographers should not base all their study in geological “well established” hypothesis. Biogeography, as a mature and independent discipline should be able to develop their own theories and views to compare with the geological theories. That’s the way Wegener (�9��) could develop his worldwide accepted theory (continental drift). At his time he was emphatically criticized by most geologists, but at the end the evidence imposed itself.

Upchurch et al. (2002), expressed it in this way “Clearly, there is not a perfect fit between the biogeographical patterns and palaeogeographical history, but there are several reasons why it would be premature to reject the biological signal: (i) palaeogeographical reconstructions are themselves hypotheses that potentially contain errors; (ii) congruence may increase as time-slicing and area selection are refined; (iii) the degree of congruence partly depends on a priori expectations regarding the effect of barriers on dispersal (e.g. phylogenetic divergence may commence before a barrier is fully developed); and (iv) the repeated area relationships are statistically supported signals that stand by themselves as patterns that require explanation”.

McCarthy (2005b) cited an interesting reflection from Johnston [1998] in his analysis of Proteaceae biogeography: “Unfortunately, at the time we wrote this paper [on Proteaceae], we were misled by conservative geologists who had not got around to accepting continental drift, and our phytogeographic understanding was much distorted by this”. McCarthy continues: “some still tend to elevate geological speculation over basic distributional realities. Implicit in papers that indulge in extravagant dispersalism and a plethora of just-right fossil absences is the notion that the basic principles of biogeography are wispy and yielding while geophysical theories are made of sterner stuff. Such papers appear to extend the legend that planetary scientists work in a field devoid of speculation, the belief that when a biogeographer and geologist confront each other on a narrow path, the biogeographer must step aside. But the question Wegener and du Toit may well have asked half a century ago is still apropos today: Does any scientist, from any field, really believe that we know more about the formation and inner workings of planets than about the locations, habits, relationships, and biomechanics of plants and animals -- about taxa that we have watched and held and explored inside and out?” (McCarthy �00�b).

Here there is a big paradox from modern biogeography: the palaeoreconstructions have been traditionally done integrating geological and paleontological information, i.e. biogeographical evidence, stratigraphic evidence, isotopic signature and palaeomagnetic data (e.g. Rapalini �00�). The integration of these different lines of evidence are full of conflict depending on the point of view and the data analysed. But usually botanists take the most accepted reconstruction and try

���

to fit the extant disjunct distributions with the major events in e.g. the splitting of Gondwana (e.g. split of Africa-America, split of Antarctica-South America, see McLoughlin �00�). But the regional tectonic reconstructions suggest that the tectonic history is much more complicated and everywhere we are dealing with geological and biotic composite areas: in Australasia (Morley �00�), New Zealand (Engler �88�, Craw �988), the Subantarctic Islands (Michaux & Leschen �00�) Tasmania (Heads �999), New Guinea (Heads �00�) and in southern South America (Crisci et al. �99�, Katinas et al. �999).

Back then to alternative palaeogeographies (section �.�): “I am fully aware that my reconstruction of Pacifica [continent] calls into question certain widely accepted dogmas of geology, and that for my failure to accept them I am accused of naïveté. However, I suspect that in the final analysis it will be the geologist who is proved naïve for his gullibility in swallowing the dogmas – hook, bait, and sinker” (Melville �98�). Or in the words of Sluys (�994): “... under a vicariance paradigm the classical pre-drift reconstruction of Pangea cannot adequately explain trans-Pacific tracks. Therefore, alternative paleogeographic models may be invoked as explanatory hypotheses: the lost continent Pacifica, island integration, a new reconstruction of eastern Gondwanaland, an expanding earth. None of these alternative models is fully compatible with all geological and biogeographic data available at present. It is stressed that biogeographic data and theories should not be made subservient to geological theories. Biogeographical data on flatworms may indicate paleogeographical relations which are worthy of examination by geologists”.

Also Linder & Crisp (�99�) found that the biogeographic pattern found in Nothofagus was not congruent with geological hypothesis and wrote: “This is not congruent with the current geological theories, nor with the patterns evident from insect biogeography. We suggest that concordant dispersal is an unlikely explanation for this pattern, and propose that the solution might be found in alternative geological hypotheses (Linder & Crisp �99�, see a recent analysis of Nothofagus by Knapp et al. �00�, and Heads �00�).

6.5 Species and speciation The concept of species and modes of speciation is intimately related to the whole task of molecular dating and consequently, to modern biogeography. Templeton (�998) argues that it is first necessary to have a definition of species before studying speciation, because “the definition of species that one uses has a major impact upon how one interprets the significance for speciation of basic evolutionary mechanisms such as gene flow or selection” (Templeton 1998, p. 32).

Howard (1998) argues that in spite of the theoretical and practical importance of a good definition for the subject of study, many biologists studying speciation have stopped following the controversy about species definition, possible for two reasons: a) most of them feel conformt with the most widely accepted definition, the so called Biological Species Concept (BSC) associated with Dobzhansky (1951) and Mayr (1963); b) on the contrary, the lack of interest could also reflects discomfort with a debate that appears to have no end. Much of the problem of the species concept

���

seems to stem from an attachment to this Biological Species Concept (BSC) as canonical truth and that knowledge of breeding compatibility has everything to do with reconstructing phylogeny. The BSC states that a species is a population of organisms that can interbreed and produce fertile offspring. It’s easy to see why it causes alarm, because it obviously can’t be applied to fossils. It’s also invalid when applied to asexual organisms (of which there are not few…)��. Complicating the discussion, Wilkins (2002) recognize more than 20 species concepts; some that conflict in part and others that are virtually incompatible. In every discipline of the natural sciences (i.e. botany, palaeontology, ichthyology), the process of diagnosis and description of a species is different insofar as the evidence is different (Parenti & Ebach in prep.).

In the words of Wilkinson: “The traditional species problem, what are species?, has no single answer because there is no single kind of thing that we call species. The species problem has often been approached with the presupposition that a single kind of entity exists in nature that corresponds to a species concept, just because the word species exists in the language of biology” (Wilkinson �990).

The modern BSC has a big disadvantage for the biogeographer, namely that the concept is separated from the spatial and temporal context, and therefore, is difficult to use in historical analysis (Zunino & Zullini �00�, p. �8). Other concepts seem to be more suited for the biogeographic analysis, like the Evolutionary Species Concept (ESC), in the sense of Wiley (�978).

The concept is intimately bounded to the modes of speciation, i.e. the practical origination of the species. For many years, the more (even the only) accepted mode of speciation was allopatric, as promulgated by Mayr (�9��). But researchers like Bush (�97�, �998) found that spatial separation of populations is not absolutely essential for genetic divergence and the formation of new species, i.e. sympatric speciation is also possible. Other modes of speciation are emerging, but still allopatric seems to be still the most vastly recognized: “Historical biogeography now had one fundamental goal – to reconstruct the sequence of events on a dynamic earth that would have passively isolated co-distributed groups of ancestral species, resulting in subsequent allopatric speciation and biotic diversification” (Riddle 2005).

It is common believe that if the range of sister taxa do not overlap, it is likely that they diverged in allopatry. On the other hand, if sister taxa exploit distinct habitat and have ranges that overlap extensively, sympatric divergence is a more likely scenario (Berlocher �998). “Unfortunately, even in well-studied groups, range maps are often incomplete, habitat utilization patterns are poorly understood, and relevant taxa are not available for genetic and morphological analysis… Until we correct this situation, the relative importance of nonallopatric speciation in generating species diversity will remain highly conjectural” (Howard �998, p. 440).

A different but certainly much related issue is the understanding of macro-evolution, i.e. the evolution of higher taxonomix categories like genera and families. But modern biogeography

���

concentrate the focus on just one taxonomic level: the species, as the basic unit in biology and biogeography, more than higher taxonomical categories that are treated as more or less artificial. Heads defines this tendency as speciescentrism: “The modern view is well exemplified by Cracraft [�00�] who included the following in his seven great questions of systematic biology: What is a species?, How many species are there?, Where are the Earth’s species distributed?, and How have species’ distributions changed over time?” Heads find equally myopic speciescentrism in Hubbell’s (�00�) Unified Neutral Theory of Biodiversity and Biogeography (Heads �00�a, p. �0�). In spite of efforts made by authors like Salomon (�00�), the synthesis between modes of evolution and bigeography seems to be still far.

Modes of speciation have been studied in the Chilean flora since the seminal papers of B.B. Simpson (�97�). She studied the modes of speciation in several groups of Perezia coming to the conclusion that the Prenanthoides group suffered phyletic evolution (i.e. genetic stasis). Simpson suggests that this might be the rule for the biota located in the Nothofagus forest habitat from ��° to 4�°S. On the other hand, Magellanica group shows a case of true speciation (i.e. the splitting of an ancestral stock into several species by means of geographical isolation = allopatry). The Magellanica group’s evolution is related to the last glaciations, in which the species suffered reduction of the population size and the presence of barriers to gene flow. These results suggests that the taxa living in higher elevations (above the timberline) have been pushed to rapidly adapted to this new environments. But phyletic evolution sounds paradoxical, since it considers evolutionary stasis for a long period, with an abrupt change conducent to speciation only during the Pleistocene (sensu Simpson �97� for Prenanthoides group).

Hershkovitz et al. (�00�) found recently the contrary situation, also working with Asteraceae: the generally lowest elevation species of Chaetanthera appear to be the most evolutionarily derived. The authors describe these results as “contrary to intuition”, and suggest that the “lower elevation taxa would have required a secondarily evolved tolerance to the increased aridity developing on the western slope of the Andes that became intense from the Pliocene onwards” (Hershkovitz et al. �00�, p. ��).

In support of Simpson’s findings are the results with another Asteraceae genus, Hypochaeris, which suggest that populations from the coastal Cordillera are older than the Andean ones (Muellner et al. �00�).

The speciation mode described as phyletic evolution by Simpson (�97�) has been more recently treated as anagenetic speciation by Stuessy et al. (�00�). In anagenetic speciation “a founder population arrives on an island and simply diverges through time without further specific differentiation” (Stuessy et al. �00�). This anagenetic speciation has been suggested as an important mode of evolution in the endemic vascular plants of the Juan Fernandez Islands (Stuessy et al. 1990). The Fernandezian flora has been one of the best study cases for speciation on islands (Stuessy & Ono �998). Carlquist (�974) mentioned Juan Fernández as one of the best scenarios for

��4

testing adaptive radiation (= occurrence of congeneric species in different habitats), exemplified in the genera Dendroseris and Robinsonia. The author found a paradox in the existence of this radiation since “the small size of the islands …seems too limited to promote adaptive radiation” (Carlquist �974, p. �0�). Sanders et al. (�987) attempted to characterize this habitat differences (primarily edaphic factors) but were not able to demonstrate clear-cut differences. On the contary, Stuessy et al. (�990) suggested that the signs for adaptive radiation are equivocal and the species-richness of this genera should be rather explained by anagenesis (see also Stuessy et al. �00�).

Noteworthy, Hughes & Eastwood (�00�) still treat the whole Andes as a scenery (archipelago) of island radiation. Gentry (1982) proposed a combination of speciation modes, in wich at a finer scale, microgeographic (allopatric) speciation, perhaps even more or less sympatric, is probably the rule rather than the exception in the Andes. Smith & Baum (�00�) postulate that periodic hybridization events (i.e. sympatric speciation) coupled with pollinator-mediated selection and the potential for microallopatry may have acted together to promote diversification in montane Andean taxa, such as Iochrominae (Solanaceae) (the genus Dunalia occurs in northernmost Chile).

6.5.1 Hybridization “…as the principles of cladistic analysis were examined more closely, even this extremely parsimoniuos view of biological systematics turned out to have some problems… hybrid species result in ambiguous cladograms – and, contrary to what zoologists frequently claim, hybrid species are not at all rare. They are common among plants” (Hull �00�, p. ���).

As proposed by Smith & Baum (�00�) for a clade of Andean Solanaceae, the modes of speciation could be a continue mixture of (micro)allopatric and sympatric episodes of hybridization. It appears that these events have patially obscured the underlying divergent phylogenetic history, having clouded the branching pattern in some parts of the tree. Some authors like Dickerman (1998) have developed modified parsimony methods to detect hybridization or horizontal gene transfer. Descent patterns in a phylogenetic system with a single hybrid event can be described as the sum of two gene trees, each describing the history of part of the genetic material composing the system. Systems with more than one hybrid event will require a larger set of trees. This set of gene trees is called a phylogenetic forest. Dickerman’s (�998) method returns a reticulate hypertree in which some taxa have more than one immediate ancestor, as one would expect if hybridization or horizontal transfer has occurred. Ané and Sanderson (�00�) developed another method that returns either a single tree or a collection of trees. In the opinion of Dickerman (�998), the dominance of the tree model in phylogenetics can be attributed to two principal strengths: (a) algorithmic and conceptual simplicity of trees and (b) the expectation that certain evolutionary processes generate treelike descent patterns. “Algorithmic simplicity, as exploited by computer programs for phylogenetic inference, remains one of the reasons tree analysis is so prevalent in biology. Admitting the possibility of hybrid taxa with two or more immediate ancestors implies the need to consider graphs of descent relationships more complicated than trees” (Dickerman

���

�998). In the same line, Posada & Crandall (�00�) and Winkworth et al. (�00�) found that the complex evolutionary processes that characterize plant species radiations are not likely to be well represented by bifurcating tree models. These authors propose that phylogenetic networks provide a better tool because they are capable of graphically representing the competing signals in a data set, and are therefore more suited for biogeographic analysis and interpretation. That the process of horizontal gene transfer is not to be diminished is shown by the recent results from Davis et al. (2005), which report for the first time the horizontal gene transport from an angiosperm to a fern (from root-parasitic Loranthaceae to the fern Botrychium virginianum).

Hershkovitz (�00�) discussed the role of hybridization as a possible explanation for the morphological and molecular diversity of Andean Portulacaceae. He wrotes “cladogenesis de facto represents a priori an ad hoc assumption, because the most popular phylogenetic methods constrain for cladogenesis without justification as to why this process should be preferred, much less assumed, in the group in question”. But “It is perhaps more the rule than the exception that, at least in plants, the conditions necessary and sufficient for hybridization exist. … In phylogenetics, hybridization is discriminated against in part because it introduces tremendous mathematical complexity into phylogenetic reconstruction and in part because the relevant parameters are poorly understood or cannot be easily evaluated using common molecular systematic approaches [Linder & Rieseberg �004]”. Hershkovitz (�00�) found for the Andean Portulacaceae, “the unexpected similarity of the marker sequences in morphologically divergent taxa may itself result from hybridization”. He therefore suggests that “…interpretation of molecular data derived from one or a few samples per taxon and few DNA sequences must allow for the possibility of past gene flow…[]…If ecological instability and novelty of habitats favour hybrid success, then current and historical environmental parameters must be considered favorable to hybridization of Andean Portulacaceae”.

The existence of x Margyracaena, an hybrid genus between an endemic species (Margyricarpus digynus) and the introduced genus Acaena in Juan Fernández, is a good example of the real possibilities of the hybridization process.

6.5.2 Monophyletic v/s polyphyletic

L. Constance, while analizing the antitropical (amphitropical) disjunct pattern recognized “on the common assumption that each species, genus, or natural family has subsequently radiated in all directions to occupy suitable territory, we should be led to expect a series of relatively continuous distributions of such taxa over the earth’s surface, much in the sense of Willi’s concept of Age and Area” (Constance 1963, p. 109-110). He continues “…Attempts to explain the broken ranges of living or extinct organisms have led to postulate polytopic origins, parallel and convergent evolution, orthogenesis, shifting poles and an oscillating equator, long-distance dispersal, land bridges, and continental drift, among others”. Constance ruled out some explanations: “…modern bicentric or vicarious species can, then, scarcely be the relicts of widely and continuously

���

distributed ancestral populations. Nor am I able to accept the idea of a polyphyletic or polytopic origin of these, although such an explanation presented no difficulty to the pre-Darwinian Special Creationists. In their more modern forms, postulated polytopic origins simple push the problem backward to the presumed existence of a more continuously distributed ancestor, which was able to give rise to “look-alike” descendents in several or many portions of its range [Briquet, Süssenguth, Pachosky, Rosa, Schröter, Du Rietz, Wulff]… (Constance 1963, p. 111)�7 . In fact the multiple origins hypothesis have been systematically dismissing since Darwin. But early botanists like A. Engler or O. Drude accepted the possibility of multiple origins for major groups (quoted in Bews �9��, p. 4). Guppy advanced the theory of differentiation and pointed out: “if the differentiation hypothesis is correct, no natural order could have been developed on the lines implied by the Darwinian theory, which, as interpreted in recent works, begins with the variety and terminated with the order, a process that reverses the usual methods of Nature…Yet such a process, as is there implied is common enough in the plant world, but it accounts not for the natural orders but for the oddities of plant forms …It is here termed a specializing process in contrast with that of differentiation; but it is the differentiation process that has been the principal determining cause of diversification in plants” (Guppy quoted by Bews 1921, p. 3).

In this sense, Hu (�9�0) proposed a polyphyletic system of classification of angiosperms, and more recently some authors still proposed that flowering plants evolved from multiple, unrelated seed plant lineages, like in the Polyphyletic-Polychromic-Polytopic hypothesis (Wu et al. �00�) and Nair’s Triphyletic theory (Nair 1979). While carpels, double fertilization, and flowers are viewed by some to be too complex to evolve more than once from totally unrelated seed plant lineages, evidence from studies of phenotypes of homeotic genes suggest otherwise. The activities of insects may have hastened and stimulated the development and evolution of seed plant reproductive organs multiple times, in many evolutionary lines (Miller �00�).

The words of Gilmour sound still valid: “In any case, I think it is doubtful whether the expression degree of relationship can be validly applied in connexion with the process of evolution. Singer has pointed out that the word inheritance, as used in biology, has been derived from the concept of legal inheritance in human affairs, and that many of the legal associations of the word have been carried over and tend to obscure its biological meaning. The same process has, I suggest, occurred with the word relationship. The word as commonly applied to the products of the sexual reproductions of human individuals has quite a definite meaning. Thus in this sense two brothers are more nearly related than two cousins. Here, the degree of relationship depends on the repetition of a uniform process, namely mating and birth. In evolution, however, we are not dealing with a uniform process, but with an immensely complicated tangle of many different processes, and it is surely inadmissible to talk of plants being more nearly or more distantly related to each other from an evolutionary point of view. For example, take the case of a population of plants spreading in a certain direction. At one point part of the population may become isolated by a natural barrier and may develop into a distinct morphological type with a different genotypic constitution, while at another point altered conditions may induce a doubling of the chromosomes resulting in a

��7

second distinct morphological type. One cannot, I think, say that these derivative populations are equally related to the original population, nor that one is more nearly related to it than the other. They have certainly both been derived from the original population, but by quite different methods, and one cannot, I think, speak of degree of relationship in connexion with the process …These considerations are in harmony with the view, which is now accepted by many biologists, that the course of evolution cannot be adequately represented by a diagrammatic phylogenetic tree, as is done for human and animal pedigrees, but rather by a network in three dimensions; and, if it is not pressing an analogy too far, one might add, a network with meshes of different sizes and cords of different thicknesses” (Gilmour �9��, p. �0�).

In this sense, Heads opinates: “it seems safer to assume that ancestors in general are not necessarily single species but may be polymorphic complexes, with many different states already present for each character” (Heads �00�b). Furthermore, Stuessy do not ruled out the possibility of a “polyphyletic origin of life on Earth” (Stuessy �00�, p. �7).

6.6 Re-inventing an origin for the land plants

The green algae known as stoneworts (Charales) are suggested to be the extant sister group to all land plants, although the phylogeny is still not conclusive (Lewis & McCourt �004, McCourt et al. �004). Megafossil evidence for the land plant crown group comprises taxa from the Middle Silurian, about 4�0–4�0 mya (Kenrick & Crane �997) or early Late Silurian (Wellman & Gray �000). However, microscopic spores from the Ordovician have supposedly derived from parent plants that were bryophyte-like if not in fact bryophytes (Wellman & Gray �000, Wellman et al. �00�). The fossil-based age estimate for these microfossils is mid-Ordovician, about 47� My ago. It is only from the Late Silurian onwards that the microfossil/megafossil record can be integrated and utilized in interpretation of the flora. The fossil record of plant megafossils is poor and biased, with only a dozen or so known pre-Devonian assemblages.

Heckman et al. (�00�) challenged the more traditional view: analysing �0 protein sequences, the authors estimated a Precambrian age for land plants of 70� +/- 4� mya. This result contrasts sharply with palaeobotanical estimates (Kenrick & Crane �997, Wellman et al. �00�). Sanderson (�00�) applied a penalized likelihood approach with two alternative and internal calibration points, and different genes than Heckman et al. (�00�). Sanderson (�00�) results suggest the land plant crown group to be of Ordovician age (48� or 490 mya depending on calibration point used). The author also conducted analyses invoking a molecular clock assumption. This increased the incongruence with the Heckman et al. (�00�) results, pushing the land plant origin forward into the Early Silurian (4�� and 4�� mya, depending on calibration point).

��8

6.6.1 The origin of the vascular plants

Traditionally, vascular plant evolution was seen as a successive series of incremental increases in complexity, from simple bryophytic ancestors through vascularized spore producers, more complex seed plants, and ultimately the angiosperms (Pryer et al. �004). During the past �0 years the phylogenetic relationships are beginning to be more clear and the predominant paraphyletic former schema is changing to a more comprehensive system of evolutionary relationships. Monilophytes and lycophytes are all spore bearing and ‘seed-free’, and therefore, were traditionally lumped under terms like ‘pteridophytes’ or ‘ferns and fern allies’, that united paraphyletic assemblages of plants.

It is now believed that a deep phylogenetic dichotomy occurred in the early-mid Devonian (ca. 400 mya), separating the lycophytes from the euphyllophytes (Gensel & Berry �00�, Pryer et al. �004) (Fig. �). Lycophytes all possess lycophylls (leaves with an intercalary meristem) and comprise three main clades: homosporous Lycopodiales (clubmosses), heterosporous Isoetales (quillworts) and Selaginellales (spikemosses). Extant lycophytes are mostly diminute plants, but many fossil members (e.g. Lepidodendron) were large arborescent forms that dominated the Carboniferous landscape and are today the major component of coal deposits (Stewart & Rothwell �99�).

The most diverse of the monilophytes are the leptosporangiate ferns, a group of more than ��000 extant species. The earliest known occurrence of fossil leptosporangiate ferns is in the Early Carboniferous (Galtier & Philips �99�); by the end of the Carboniferous six families were present. In subsequent major radiations in the Permian, Triassic, and Jurassic, several families with extant representatives replaced these Carboniferous families (e.g. Osmundaceae, Schizaceae, Matoniaceae, and Dipteridaceae) (Rothwell �987). The more derived polypod ferns dramatically diversified in the Cretaceous, acompanying the great angiosperm diversification (Schneider et al. �004).

The extant seed plants (the Spermatophyta) have been hypothesized to be a monophyletic group, supposedly having arose from a common ancestor and with an initial radiation in the Late Palaeozoic (Stewart & Rothwell 1993). The five lineages recognized within the seed plants have been shown to be monophyletic by most studies. The earliest known seed plants have been reported from the Paleozoic (Late Devonian, about �70 mya) of West Virginia (Rothwell et al. �989, Kenrick & Crane �997).

But there is still a high level of uncertainty: “If the gymnosperms are indeed monophyletic, their sister-group the angiosperms must date from the same period, the Carboniferous. This leaves a gap of over ��0 million years with no fossil record of angiosperms – a period longer than their entire known fossil history. This could be either because the gymnosperms are not a natural group, or because the stem lineage of the angiosmerps lacked distinguishing angiosperm synapomorphies” (Hill �00�).

��9

6.6.2 Angiosperms origin(s)

The origin of flowering plants is considered by some to be the “Holy Grail” (Miller 2006), or “Darwin’s abominable mystery” of botany (e.g. Crepet �998, Bowe et al. �000, Davies et al. �004). Janssen & Bremer (�004) wrote: “From fossil evidence, a major radiation of angiosperms is obvious in the mid-Cretaceous ��0–90 mya [Lidgard & Crane �990, Herendeen & Crane 1995]. With the diversification burst of angiosperms setting in no earlier than about 115 mya, there still remains a considerable gap between the fossil record and our molecular dating, with the majority of lineages already present by that time. At this time we cannot decide whether this is an indication of an incomplete fossil record or an argument against currently available molecular dating methodologies”.

Due to common results from the molecular clock approach that yielded age estimates grossly inconsistent with the fossil record, Bell et al. (�00�) investigated the age of angiosperms using Bayesian relaxed clock (BRC) and penalized likelihood (PL) approaches. These methods allow the incorporation of multiple fossil constraints into the optimization procedure. Bell et al.’s (�00�) results indicate that widely divergent age estimates can result from the different methods (�98–��9 mya), different sources of data (�7�–��� mya), and the inclusion of temporal constraints to topologies. Most dates, however, are between �80–�40 million years ago, suggesting a Middle Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil angiosperms by about 4�–� million years. These dates are consistent with other recent studies that propose the hypothesis that the angiosperms may be older than the fossil record indicates. Wikström et al. (�00�) recognize “the calibrated phylogeny here is a working hypothesis and should be viewed as such”. In Bremer’s opinion (�00�): “The analysis by Wikström et al. (�00�) was focused on the age of angiosperms in general and their tree was calibrated with a single reference fossil in the rosid order Fagales. Thus, dating within parts of their tree topologically far away from the reference node, for example, within the monocots, is unreliable” (Bremer �00�).

But information on the fossil record of angiosperms has expanded dramatically over the past twenty-five years, particular due to the discovery of numerous mesofossil, floras with fossil flowers, that has added a completely new element into the study of angiosperm history (Crepet et al. �004, Friis et al. �00�, �00�). But the exact phylogenetic origin of the angiosperms itself remains as enigmatic as ever and, in some cases, newly introduced techniques from molecular biology have given confusing results (Friis et al. �00�): “In particular, relationships between the five groups of extant seed plants remain uncertain, and it has sometimes proved difficult to reconcile estimates of the time of divergence between extant lineages made using a ‘molecular clock’ with the fossil record. One result, however, is becoming increasingly clear: a great deal of angiosperm diversity is extinct. Some groups of angiosperms were evidently more diverse in the past than they are today”. It is also possibly that some early enigmatic fossils represent lineages that diverged from the main line of angiosperm evolution below the most recent common ancestor of all extant taxa (Friis et al. �00�). Contrasts among different morphology-based analyses can

��0

be ascribed to different character selection, different sized data matrices, and disputes over homology assessment (Crepet �998). Even nucleic acid sequences produce different hypotheses of relationship depending on the specific sequences and methodologies employed. Analyses combining different data sets are promising but have not yet resolved these issues (Nandi et al. 1998). “It is ironic that there is now unprecedented potential in the field of systematic botany, while the most fundamental relationships remain unsolved” (Crepet �998).

Concrete fossils supporting the hypothesis of a Jurassic origin for the angiosperm are found in China, in the genus named Archaefructus (Sun et al. 1998) later classified as the monotypic family Archaefructaceae (Sun et al. �00�). The fossils are a combination of strongly magnolialean characters and a notable nonmagnolialean one: a missing perianth, an unusual condition found only in some species in the families Chloranthaceae and Piperaceae. Friis et al. (�00�) have criticized the dating of the fossil (to Early Cretaceous instead of Late Jurassic), and the phylogenetic position as sister clade to the rest of extant angiosperms. But reanalysing the reanalysis done by Friis et al. (�00�), specially correctly coding the leave form of Cabomba, still resolves the genus as the most basal in the angiosperm phylogeny (Nixon pers. comm.). In Crepet’s words “I have learned that the discovery of a few specimens of a new fossil taxon is seldom a unique event--there will be new specimens of Archaefructus and the kinds of characters critically needed are also the kinds likely to be preserved. Given the potential informative value of this taxon and the recent pace of innovation in studies of angiosperm systematics and paleobotany today, I predict that the great “abominable mystery,” [the origin of the angisperms] with us for over �00 years, will not last another �0” (Crepet �998). Crepet also recognized (�000): “there is some irony in the fact that despite stunning progress on many fronts, ultimate resolution still depends on the stochastic nature of fossil discovery”.

According to Lu & Tang (�00�): “The evidence from molecular clocks, fossils and geographic distribution data on the origin time of angiosperms has been greatly accumulated in the past two decades. However, fossil evidence is only the integrated embodiment for the preserved parts of plants and geological fossilization conditions, but is not, and unlikely to be, the indicator of the exact age of the groups or species. In addition, we have to consider the evolutionary history of the fossils. The application of molecular clocks is another approach, but it carries even greater errors. Besides the two lines of evidence, researches on modern distribution patterns, the formation of the plant groups, and combination of plant evolution with the earth history as well as the theory of plate tectonics, can undoubtedly improve the reliability in inferring angiosperm origin time. Analyses of �� important spermatophyte (mostly angiosperms) families or genera at different evolutionary levels have suggested that the origin time of angiosperms could be dated back to the Early Jurassic or Late Triassic… We consider that the basal angiosperm groups, i.e., members of ANITA grade belong to the primitive groups because of their many plesiomorphies. But they only share few synapomorphies, such as globose pollen grains, indicating that they may have already diverged into different lineages during the early stages of angiosperm evolution. Therefore, ANITA is a complex group originated from different lineages” (Lu & Tang �00�).

���

In view of the difficulty of finding ancestors for the angiosperms, yet also considering their sudden appearance and explosive evolutionary success, Stuessy (�004) proposed a transitional-combinational theory: “This theory suggests that angiosperms evolved slowly from seed ferns in the Jurassic beginning first with the carpel, followed later by double fertilization, and lastly by the appearance of flowers. These three fundamental transitions may have taken more than �00 million years to complete. The extant angiosperms did not appear until Early Cretaceous, as attested by micro- and macrofossils, when the final combination of all three important angiosperm features occurred. This combination provided the opportunity for explosive evolutionary diversification, especially in response to selection from insect pollinators and predators, plus attendant modifications in compatibility and breeding systems” (Stuessy 2004). The theory suggests viewing all gymnosperms, other than extinct seed ferns from which carpels arose, as having had no direct phylogenetic connections to modem angiosperms.

“The angiosperms undoubtedly originated long before the Cretaceous period. The specialised character and astonishingly modern facies of many Cretaceous angiosperms confirm our belief in an antiquity of angiosperms antedating by many millions of years, probably by several geologic periods, the first appearance of recognisable pioneers of the present ruling dynasty in the modern world (Seward �9��, cited by Takhtajan �98�, p. ���). Arldt (�9�8) and Berry (�9�0) already suggested the Triassic as the time for the appearance of the angiosperms. Several later authors postulated a Triassic or even Permo-Triassic origin (Camp �947, Thomas �947, Axelrod �9��, Zimmermann �9�9). Others like Golenkin (�9�7, quoted by Takhtajan �98�) even referred their origin to the Permo-Carboniferous, asserting that an early Mesozoic origin seems the moist likely for the angiosperms; since they could scarcely have attained such a high morphological diversity if they had arisen later that the Triassic. Croizat

Figure �.� Traditional view of vascular plant evolution (from Cornet �00�)

���

suggested an origin of the angiosperms “during the Permian as the aftermath of the so called Permo-Carboniferous Glaciations” (Croizat �9�0, p. �7��).

In concordance with Takhtajan (�9��) and earlier authors, recent molecular phylogenetic and molecular clock analysis suggest a pre-Mesozoic age for the divergence of the angiosperm lineage from other seed plants (Taylor et al. 2006). This finding greatly predates the confirmed fossil record of the angiosperm crown group. In addition, molecular phylogenetic studies have not supported the morphologically based conclusion that gnetophytes are the extant sister group to angiosperms. Taylor et al. (�00�), examining the presence of oleanane in extant and extinct taxa came to the conclusion that “if oleanane originated once in seed plants then the angiosperm stem lineage would have diverged from other seed plant lineages by the late Paleozoic”.

Figure �.� Alternative view of angiosperm origins (from Cornet �00�)

Miller (2006) briefly revised 20 theories and hypotheses for the origin of the angiosperms, while proposing a novel hypothesis based on the coevolution of flowering plants and insects, the Thigmomorphogenetic/Coevolutionary Hypothesis: “Angiospermy is a loosely defined reproductive syndrome that developed, in at least two cyclic episodes involving hybridization

���

between unrelated seed plants, over more than �00 million years of Geologic Time. Several disparate taxonomic subclasses, orders, families, genera, and species of extinct Permian seed plants probably possessed the abilities for thigmoanatomic and thigmomorphologic change leading down the long evolutionary pathway to the eventual attainment of flowering forms, even as early as Triassic time” (Miller �00�).

Results obtained by Martin et al. (�989, �99�) suggest that separation of monocotyledonous and dicotyledonous lineages took place in Late Carboniferous (�00 mya), and Wolfe et al. (�989) place the age of angiosperm origin at ��0 mya. Sanmiguelia, a Late Triassic fossil described by Brown (�9��) has been interpreted by Cornet (�989) as a very primitive angiosperm that combine both monocot and dicot characters, but this interpretation has been neglected, not so much for morphological reasons but for the general believe that angiosperms arose only in the Cretaceous (Daghlian 1981, Martin et al. 1993) (figures 6.2, 6.3).

Cornet (�00�) asks “why is there still so much resistance to the recognition of pre-Cretaceous fossils that could belong to the angiosperms or to their non-gymnospermous precursors if angiosperms did not evolve from any known group of gymnosperms (i.e., Gnetales, Bennettitales, Cycadales, Pteridospermales, and Coniferales)? Has the Cretaceous origin of angiosperms become a myth perpetuated by fear of uncertainty? Why can’t paleobotanists be more honest and say that they don’t have the answer yet to angiosperm origins, rather than bias the subject by saying “early” angiosperms and “primary” radiation in the Cretaceous - or worse by shifting the age of possible Jurassic strata to conform to a Cretaceous origin? If angiosperms evolved in the Triassic, Early Cretaceous angiosperms would be middle age. And if the genetic roots of angiospermy extend all the way back to the Silurian (see diagram above), that would make Cretaceous angiosperms “late bloomers”.

��4

���

7 Back to Postmodern Biogeography

���

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlan-gen, March-December �00�

��7

7 Back to Postmodern Biogeography

“Postmodernism hit geography like a tidal wave, and the consequences were predictable. The postmodern movement engendered intense excitement in a few scholars who were inspired by its provocations. More generally, it met with active hostility from persons who perceived their intellectual authority to be threatened, with incomprehension on the part of persons who failed to negotiate its arcane jargon, and with indifference from the majority, who willfully ignored what they perceived as the latest fad … Despite a combination of active antipathy and crushing inertia, postmodernism has flourished, because it constitutes the most profound challenge to the three hundred years of post-Enlightenment thinking. Postmodern thought holds that rationalism has failed both as an ideal and as a practical guide for social action and that Enlightenment desiderata, such as decisive theoretical argument and self-evident truth, are no longer valid. Postmodernism is no overnight sensation; in its current form it has echoed through academic corridors for three decades. Nor is it likely to disappear in the foreseeable future, despite the dismissive edicts of authoritarian academic gurus. Postmodernism is something that geographers are going to have to get used to” (Dear & Wassmansdorf �99�, p. ���).

7.1 Biogeography as a social science

The monoboreal relic hypothesis was summarized by Thiselton-Dyer (�909, p. ���) in the following words: “the extraordinary congestion in species of the peninsulas of the Old World points to the long-continued action of a migration southwards. Each is in fact a cul-de-sac into which they have

poured and from which there is no escape”. According to this hypothesis, the affinity between austral floras would thus mainly be the result of a common origin from the North. According to Du Rietz (�940, p. �44), the monoboreal relic hypothesis was worked out in great detail with regard to animals by Matthew (�9��), who came to the conclusion that “the principal lines of migration in later geological epochs have been radial from Holarctic centres of dispersal” (Matthew �9��, p. �7�), and that… “the Antarctic and southern lands being unfavourably situated for the evolution and dispersal of dominant races and contributing but little to the cosmopolitan fauna of the emergent phase” (Matthew �9��, p. ��0). Thus the bicentric austral distribution of Marsupialia in America

and Australia is, according to Matthew, “probably to be regarded as due to a very ancient dispersal from the north” (Matthew �9��, p. ���). Matthew’s views on biogeography were replicated and advanced by G.G. Simpson and the fellows of the so called New York School of biogeography. These ideas were nicely illustrated by means of caricatures accompanying Simpson’s book Evolution and Geography (box 7.�). A discussion on Matthew’s biogeographic program has been done by Nelson & Ladiges (�00�) (see also an early review by Barbour �9��). A critique is to

W.D. Matthew

��8

Box 7.� George Gaylord Simpson

be find in the writings of Livingstone (1992, 1994, 2002), one of the most lucid (post)modern geographers: “A map constructed by Griffith Taylor provides me with my final point of departure. The map was entitled Zones of migration showing the evolution of the races (figure 7.1) and it appeared as an accompaniment to the paper he published in the Geographical Review. What is immediately noticeable is the polar zenithal equidistant projection that Taylor exploited to portray his racial philosophy, a projection that effectively pushed the tropics to the margins of cultural significance… the map was intended to give visual expression to the idea that the most primitive

G.G. Simpson

��9

varieties of the human race occupied the tropical zones… That Taylor’s map was part of a more general cartographic discourse relegating the tropics to cultural insignificance is clear from two related cartographic ventures. The projection that Taylor called upon to depict his racialised conception of evolutionary history has earlier been used by his mentor William Diller Matthew in �9�� to portray both mammalian migration and the distribution of the human races” (Livingstone 2002) (figure 7.2). The implications were plain. As Matthew himself declared: “the higher races of

man are adapted to a cool-temperate climate [where] they reach their highest physical, mental and social attainments” (Matthew �9��, p. 4�). Livingstone continues noting the reprint of Taylor’s map in Huntington’s book The character of races, author who otherwise wrote “the people of the tropics are in reality the children of the human race. They represent our primitive ancestors… It is not to be expected that such people should ever rise very high

Figure 7.1 Griffith Taylor‘s map: Zones of migration showing the evolution of the races (1921).

Figure 7.� Matthew‘s map: migration and the distribution of the human races (�9��).

�70

in the scale of civilisation” (Huntington �9�4, as quoted by Livingstone �00�). Huntington himself explains that his scientific task: “it seemed to me wise to show how the principles of climate change and natural selection probably apply to the earliest human development…” (Huntington �9��, p. ix). But in Livingstone’s opinion: “the cartography accompanied a discourse of climate’s moral economy that flourished because it facilitated the elaboration of a moral topography that was crucial to the project of the racial ideology, and because it employed scientific language to diagnose and treat the sickness of a colonial regime” (Livingstone �994). The social implications/interpretations of this program will not be discussed here, but are relevant for our view that biogeographic research is not independent from the social environment that set the basis for the geography as a modern science. Theoretical and empirical associations between Social Darwinismus and Nationalsozialismus have been largely discussed by Mann (�980), Hettlage (�98�) and others as reviewed by Müller (�99�).

Let’s just briefly revise the role of Darwin and his Origin of Species in the evolution of modern geography. Livingstone (1993, p. 179) begins these reflections asking why Darwin is mostly ignored in the history of geography, with the notable exception of Stoddart (�98�) (we can add Livingstone himself, and more recently Kennedy �00�). On the contrary, Darwin is a permanent theme in biogeography, as the obligate reference point (e.g. Davies et al. �004) or as the main receptor of the critics (Croizat �9��). Darwinismus entered into geography through Friedrich Ratzel (�844-�904), who visited Ernst Haeckel’s courses at Jena during �8�9. Haeckel is known as one of the most convinced defender of Darwin’s ideas (Livingstone �99�, p. �98). Ratzel’s work was also greatly influenced by the naturalist, explorer and evolutionary theorist Moritz Wagner. He was an early devotee of Darwin’s theory, but he believed that Darwin had failed to appreciate the significance of migration and geographical isolation in the processes of speciation (Wagner �8�8). “Migration, isolation, space, and environmental determinism were all part and parcel of the Wagnerian scheme of things. And it was precisely these themes that dramatically surfaced in Ratzel’s new anthropogeographie… Ratzel’s Anthtropogeographie can best be read as an attempt to situate the new science of human geography within the naturalistic framework of Wagner’s Migratiosgesetz, which he portrayed as the [most] fundamental law of world history” (Livingstone 1992, pp. 199-200). But Wagner was convinced that direct organic modifications in response to environmental conditions could be affected and their benefits transmitted to succeeding generations, adopting rather a Neo-Lamarckian position, that provided “the theoretical sustenance on with the Ratzelian programme could thrive” (Livingstone �99�, p. �0�). This Ratzelian program has been vastly and differently reinterpreted and misunderstood (see conclusions chapter in Müller �99� and his claims against Stoddart �98�). Let’s allow Ratzel to speak himself about his relation to Darwinismus, and we will find many elements that will be later crucial in the development of biogeography�8:

“Aber wenn man nun die wirkliche Verbreitung der Lebewesen ansieht, kann man doch allen diesen Hilfsmitteln der passiven Wanderung nicht die große Wirksamkeit zusprechen, die seit Darwin und Wallace so viele ihnen zugeschrieben haben“

�7�

„Da sich Länder und Meere auf der Erde ununterbrochen verschoben haben und noch heute unter unseren Augen sich verschieben, so verändert sich also beständig der Lebensraum für die wasserlebenden und ebenso die landlebenden Organismen“�9.

„Das Suchen nach einen Mittelpunkt kann uns dabei offenbar nur verwirren. Besonders bei der Erforschung des Ursprungs eines Volkes oder eine Rasse darf man nicht vergessen, daß man weder nach Punkten (Ursprung) noch nach Linien (Wegen), sondern nach

Räumen oder Gebieten zu fragen hat“.

Ratzel’s (�90�) book Der Lebensraum: eine biogeographische Studie, is one of the first publications explicitly applying the concept, and therefore the German geographer can be considered, together with Merriam (�89�), as one of the founders of the discipline (see Ebach & Goujet �00�, Parenti & Ebach in prep.).

We can not satisfactorily end this discussion about Darwinismus due to the big impact from the theory in all the biological sciences and modern life in general. Darwinism has been periodically deconstructed (e.g. Moore 1991, Hull 2005), and the task is far from being resolved: “… graduate students today are often understandably wary of getting into any topic remotely associated with Darwin and his science, if only because they assume that it has been worked out very thoroughly by now…[]…even on the most familiar topics in this area, there are plenty of opportunities for new inquiries, interpretations and themes” (Hodge �00�, p. ���).

Some neodarwinists have been very critical against the course of postmodern science, e.g. Richard Dawkins has applauded the constant efforts of A. Sokal in reveal intellectual impostures from postmodern social scientists (Sokal & Bricmont �998, reviewed by Dawkins �998). In fact, the Sokal Affair is already a classic in revealing the thin boundary between critical science and pseudo-science or nonsence. Certainly, as Dawkins wrote, one must be well aware of the extremes in which intellectuals can fall to maintain an academic position. But these valid critics do not liberate Dawkins from the challenging views against his own view about evolution. Dawkins Selfish Gene (�989) emphasize the idea that the organism is a survival machine for its genes, a “robot that has a brain, eyes, hands, and so on, but also carries around its own blueprint, its own instructions” (Dawkins in Brockman �99�, p. 7�). Dawkin’s is graphically described as an ultra-darwinist whose vision has been criticised mostly by palaeontologists like Gould, Eldredge and Vrba (see the up to date discussion by Gert Korthof [www.wasdarwinwrong.com]). Eldredge, for example, propose that Neo-Darwinism is a “gene centered and essentially reductionistic approach to evolutionary explanation” and a “distortedly oversimplified view of the natural world” (Eldredge �99�, p. 4). Eldredge challenges the idea that genes are wholly responsible for who we are. He takes up the contradiction between the competitive struggle between organisms

Friedrich Ratzel

�7�

and the cooperative behavior of social animals, such as ants and monkeys (a vision similar to the early one of Kropotkin �90�). Eldredge argues that since an organism can reproduce only if it can survive, the central concern must be maintaining fitness to the natural environment. A cooperative nature creates social structures that are fusions of economic and reproductive life. Moreover, humans are less likely than any other creature to follow selfish biological rules. Learned behavior, not instinct, dominates our reproductive conduct. Therefore, genes do not drive evolution – large-scale ecosystems change does, by means of the so called punctuated-equilibrium model (Eldredge & Gould �97�, Eldredge �004). Milton, in his critique on Richard Dawkins writes: “As I sit and read Dawkins continual cry of ‘It must have happened this way – what other rational explanation is there?’ I am sometimes almost tempted to agree with him. But then I think of Antoine Lavoisier, secretary of the Academie des Sciences who disbelieved in meteorites and who told his fellow academicians, ‘Gentlemen, stones cannot fall from the sky, because there are no stones in the sky.’…[]…A wider scientific perspective -– a very much wider perspective -– was needed before Lavoisier and his contemporaries could grasp how it can be simultaneously true that there are no stones in the sky, and yet stones can and do fall from the sky” (Milton �00�).

Worth of final consideration for our purposes is the question if Social Darwinism is as result of the theory, or this being a common misconception. Rather the contrary, well expressed by R. Williams: “… the biology itself has from the beginning a string social component, Indeed, my own position is that theories of evolution and natural selection had a social component before there was any question of reapplying them to social and political theory” (Williams �97�, as quoted by Livingstone �99�, p. �8�).

In the same line are Gieryn’s (�999) Cultural Boundaries of Science, or Kohler’s (�00�) Lanscapes and Labscapes, in which the former explores how boundaries of science are sought out in complex entanglements with political and cultural forces, and the latter questions if Clement’s famous field ecology quadrats might be the reflection of Nebraska’s cultural landscape in the mind of the ecologist (Anker �00�).

“a better explanation for the cultural authority of science lies downstream, when scientific claims leave laboratories and enter courtrooms, boardrooms, and living rooms. On such occasions, we use maps to decide who to believe—cultural maps demarcating science from pseudoscience, ideology, faith, or nonsense…[]…Was phrenology good science? How about cold fusion? Is social science really scientific? Is organic farming?” (Gieryn �999)

After centuries of disputes like these, Gieryn finds no stable criteria that absolutely distinguish science from non-science. “Science remains a pliable cultural space, flexibly reshaped to claim credibility for some beliefs while denying it to others” (Gieryn �999).

In this respect also fits Cornet’s claims regards the neglecting of pre-Cretaceous angiosperm fossils by the scientific community (section 6.6): “Funding is paramount for survival in the academic

�7�

world of scientific research ... Without money, a scientist cannot do research, and without a job or income, the scientist cannot do professional science… [But] despite the discovery of angiosperm-like fossils in the Jurassic and Triassic, funding for searches in the Jurassic will not occur - until those at NSF are satisfied that the oldest angiosperms will not be found in the Cretaceous. That will require a paradigm shift and/or a major indisputable discovery in the earliest Cretaceous of fossil angiosperms too advanced to be the oldest angiosperms (if molecular chronology can be accepted by paleobotanists, and the Magnoliales and Winterales are recognized as existing by the late Neocomian, that milestone has already been passed). A paradigm shift will require the lawn nearest the porch light to be searched completely again and again until it is accepted that the key must lie beyond the search area. In other words, the Early Cretaceous does not contain a record of all basal angiosperms, despite claims to the contrary [cf. Crepet �000]. Until that happens those who think angiosperms have a more extensive pre-Cretaceous record have to fund their research out of pocket and with little support from their peers. For now, belief in the Cretaceous origin of angiosperms = funding, and funding means Darwinian survival” (Cornet 2002).

This kind of political and funding interest that permeate science is the same discussed by Endersby (�00�) regards the media coverage gained by the APG group (section �.�) and by Hamilton (�00�) regards the assumptions in current geologic paradigms like plate tectonics (section �.�).

This is in concordance with the vision of Griffith (2004b) regards the classical interpretation of cactus evolution. The author suggests that our understanding on cactus evolution is still influenced by the cultural context of the 1500s, when the exotic members of the family were first seen in Europe. “The striking, but culturally determined, exoticness of the Cactaceae still impacts our concept of what is relictual and derived for the family” (Griffith 2004b). The traditional view, codified in the early 20th century by Britton & Rose, proposed that leafy cacti are primitive or relictual; on the contrary, stem-succulent cacti are derived. Britton & Rose (�9�9) proposed the genus Pereskia, due to its similarity to other woody flowering plants, the nearest cactus relative to the other families. The most derived are the Cactoideae leafless plants like Echinopsis. The Pereskia-primitive idea has been so influential that “recent authors have sometimes misinterpreted new data to be consistent with Britton & Rose, even when the data are more readily interpreted as contradictory” (Griffith 2004a). That means that the evolutionary interpretations are bounded to the contextual bias (e.g. the horticultural landscape from traditional botanical gardens and colleges). Griffith challenges this view and proposes that the nearest relatives of Cactaceae were not broadleaf dicots superficially similar to Pereskia; rather, the Cactaceae likely evolved from diminutive, often geophytic Portulacaceae, including the genera Anacampseros, Talinum, Talinella and Portulaca (Hershkovitz & Zimmer �997, Applequist & Wallace �00�). Furthermore, southern South American Maihuenia, with its terete, succulent leaves and cushion habit, seems to form a deep, subfamilial lineage (Wallace 1995, Griffith 2004b) (box 7.1). Walters expressed the same view of Griffith in a more general sense: “…we have no reason to think that angiosperm classification would be substantially the same if botany had developed in, say, New Zealand in the nineteenth century instead of medieval and post-medieval Europe” (Walters �9��).

�74

As Edwards et al. (�00�) recognize, there have been several hypotheses regarding the geographical origin of the cacti, all naturally based on where the presumably basal members of Cactaceae currently reside. Buxbaum (�9�9) cited both the Caribbean and central South America as likely areas, due to the presence of Pereskia and of opuntioid and cactoid lineages that he considered ‘ancestral’.

Wallace and Gibson (�00�) hypothesized a central Andean origin for the family. They assumed that Andean Pereskia, certain Opuntioideae, and the cactoid Calymmanthium are basal cactus lineages, and all reside in Peru, Bolivia, and northern Argentina. Leuenberger (�98�) suggested a late Cretaceous origin of the group. Recent studies (Hershkovitz & Zimmer �997, Nyffeler �00�, Edwards et al. �00�), suggest that the cacti can’t be that old because sequence divergence between major clades is limited. Edwards et al. �00� assert that both Opuntioideae and Cactoideae originated in the central Andean region of Peru, Bolivia, and northern Argentina. They appeal to the Andean orogeny as the cause of diversification for many plant lineages, as early authors already recognized (e.g. Raven & Axelrod �974). Early uplift in the central Andean region (��–�0 mya) is hypothesized to have occurred under a fluctuating arid/semi-arid climate regime (Hartley 2003), which presents a likely scenario for early cactus diversification. Edwards et al. (2005) further suggest that the placement of Cacteae (an exclusively North American lineage with its center of diversity in Mexico) among the earliest diverging lineages of Cactoideae, relatively early movements out of Southamerica across the continent.

Leuenberger (�98�), in his monograph about Pereskia, concluded that northwestern South America was a more reasonable location, suggesting a late Cretaceous origin of the group. Centering the origin far away from Africa might explain the poor representation of cacti in that continent. In fact, all major lineages of the cacti, i.e. Pereskioideae, Maihuenioideae,

Opuntioideae, and Cactoideae, occur mostly or exclusively in South America. Furthermore, the closest relatives of the cacti from the ‘portulacaceous cohort’ (Applequist & Wallace �00�) have their highest diversity on continents of the former Gondwana (Hershkovitz & Zimmer �997). This is generally taken as circumstantial evidence that the family Cactaceae originated in South America (e.g. Buxbaum �9�9). Hence, various groups of Pereskia, Opuntioideae, and Cactoideae invaded Central and North America and the Caribbean from their postulated northwestern South American center of origin (Leuenberger �98�). The presence of a Rhipsalis species in tropical Africa, Madagascar, and Sri Lanka (Barthlott �98�, Barthlott & Taylor �99�) led some authors to propose that this distribution indicates an old vicariance between South America and Africa (e.g. Backeberg �94�) or even an origin of the cacti in the Old World (Croizat �9��). This would imply that the cacti originated before the split of the southern continents during the late Cretaceous and that all other cacti

Box 7.� Some notes on the biogeography and evolution of the Chilean Cactaceae

Chilean Cactaceae genera are classified under the following subfamilies and tribes.

Subfamily Tribe Genera

Maihuenioideae MaihueniaOpuntioideae Cumulopuntia, Maihueniopsis, Miqueliopuntia, Pterocactus, Tunilla

Cactoideae Trichocereeae Echinopsis, Haageocereus,Oreocereus

Notocacteae Austrocactus, Copiapoa, Eriosyce, Eulychnia, Neowerdermannia

Browningieae Browningia

Pachycereeae Corryocactus

�7�

Maihueniopsis

Tunilla

Miqueliopuntia

Opuntioideae

#

#

##

Cumulopuntia

Pterocactus

Eriosyce

Austrocactus

Eulychnia

Neowerdermannia

Copiapoa

Notocacteae

BrowningieaeBrowningia!

Pachycereae$ Corryocactus

!$

Echinopsis

Oreocereus

Haageocereus

Trichocereeae

MaihuenioideaeMaihuenia

$$$

$

Box 7.� (continuation)

�7�

54°

48°

42°

36°

30°

24°

18°

NotocacteaeCopiapoaEriosyceEulychniaNeowerdermanniaAustrocactus

!

#$

$!

$

$

####

##

#

#

#

$

!

!

!

54°

48°

42°

36°

30°

24°

18°

Opuntioideae

CumulopuntiaMaihueniopsisMiqueliopuntiaTunillaPterocactus

#

!"

$

##

!

!!

!

!

!

!!

"

"

""

"

""

"

$$

$

$

"

!

#

$

54°

48°

42°

36°

30°

24°

18°

TrichocereeaeEchinopsisHaageocereusOreocereus

!

#$

!!

!!!

!

!!!

!!

!

!

!!

$$#

#

#

that might have naturally occurred in Africa got extinct. More recently, however, this distribution pattern of Rhipsalis baccifera, which is characterized by having very sticky seeds (Barthlott �98�), has been explained as the result of relatively recent long-distance dispersal by birds (Gibson & Nobel �98�, Barthlott & Hunt �99�). Maxwell (�998) enters into the discussion: “I consider the bird-dispersal scenario to be as dead in the water as any rain forest bird that tries to fly the Atlantic. This really leaves only the vicariance explanation as a viable option… Another question must be asked: Why, of all the cacti that have juicy fruits (and are therefore potentially attractive to birds) should it be Rhipsalis that has this wide distribution?” (Maxwell �998).

While there is growing consensus concerning the spatial origin of cacti in northern South America, there is a disagreement about the temporal aspect of cactus origin. Traditionally, a

Late Cretaceous origin of cacti, ��–90 mya following the breakup of the western part of the Gondwana supercontinent, has been favoured (e.g. Gibson & Nobel �98�, Mauseth �990). This time frame would allow explanation of the absence of endemic cacti in the Old World, while maximizing the time for the evolution of the various distinctive morphological features of extant cacti (Hershkovitz & Zimmer �997). Hershkovitz & Zimmer (�997) proposed a much more

recent origin of cacti in the Late Paleogene (�0 mya). Nyffeler (�00�) adds that the small amount of sequence divergence found in the data set of chloroplast markers is indicative of a recent origin of the major radiations in Cactaceae.

Edwards et al. �00� still suggest a basal split in Cactaceae in a northern clade comprising of Pereskia species and a southern clade comprising also Pereskia species and Maihuenia and other Cactoideae and Opuntioideae. Edwards et al. �00� recognize that their sampling and resolution within Opuntioideae are insufficient to make inferences about the geographic distribution of its basal members, and recognize recent results that placed its earliest diverging lineages in Chile and Argentina (Griffith 2004a).Griffith (2004a) further propose to avoid the cultural bias in cactus evolution interpretation, through data sources that “do not involve form”, i.e. DNA sequence data.

Box 7.� (continuation)

�77

A summary about the recent results within the Cactaceae is as follow:

1. the nearest relatives of Cactaceae were not broadleaf dicots superficially similar to Pereskia; rather, the Cactaceae likely evolved from diminutive, often geophytic Portulacaceae, including the genera Anacampseros, Talinum, Talinella and Portulaca (Hershkovitz & Zimmer �997, Applequist & Wallace �00�).

�. Some studies still consider Pereskia as a deep lineage (Nyffeler �00�, whereas others interpret it as a derived group, relative younger than the Opuntioideae (Griffith 2003). Furthermore, Pereskia appears as paraphyletic in latest studies, and more studies about the possible ecological constrains related to the evolutionary history are needed (Edwards et al �00�).

�. A plausible model of evolution is that some common ancestor between the Portulacaceae and the ‘proto-cacti’, a xerophytic lineage arose which was capable of radiating and speciating in America, most likely after the split up of Gondwana. If Rhipsalis participated in this early evolution is still a controversy.

4. Maihuenia form a deep, subfamilial lineage (Wallace 1995, Griffith 2004b).

�. Maihueniopsis appears to circumscribe a suite of early Opuntioid characters including diminutiveness, early deciduous leaves and geophytism (Griffith 2004a).

�. Pterocactus is a deep lineage within the Opuntioideae (Griffith 2004b).

Due to the absence of Pereskia in Chile, the most ancestral Chilean genus is Maihuenia. Geographic ranges of Maihuenia, Maihueniopsis and Pterocactus superpose in southern Argentina and Chile. The northern limit for Chilean Opuntioideae is represented by the genus Maihueniopsis, Tunilla and Cumulopuntia that reach southern Peru and Bolivia. The southern limit in Opuntioideae is represented by Pterocactus that apparently reachs the Magellan strait in Argentina and shows the southern limit for the family.

basal Opuntioideae

Maihuenia

Other Opuntioideae + Pereskioideae + Cactoideae

Biogeographical scenario for the evolution of Cac-taceae (compare with Fig. � in Edwards et al. �00�)

Box 7.� (continuation)

�78

7.2 Biogeography toward a science of qualities

“If accepted, these views of language and science have tremendous implications. They demolish religion, science, and any other means of “knowing” imaginable. Each person is consigned to play their own language “game,” never truly understanding the extent to which all thought is shaped and determined by the prison of syntax and grammar. Nevertheless, despite its unattractiveness, Deconstruction and its cousin Postmodernism do have a certain logic and appeal. They are, without a doubt, carefully arranged and skillful critiques of the current means of knowing. Either way—whether Postmodernism is nonsense or “truth”—the dominance of Modernist thinking seems severely challenged, and there is little doubt that the reverence this culture has had for science will necessarily change as a result of this opposition” (Kau �00�).

Going back to the beginning of the thesis, the main question in modern and postmodern biogeography still remains: how is it possibly to change the course toward “the grand view, the ultimate synthesis across space and time”? (sensu Lomolino & Heaney �004). Does not the tendence run just in the opposite way, due to the fragmentation of the discipline and of natural sciences in general? Biogeography seems to suffer the crisis of reductionism of modern science, and there is no way of turn the tendency. Recent summaries of biogeographical methods and approaches (e.g. Crisci et al. �00�, Morrone �00�) serve more for feeding the critics about the supposed identity crisis of biogeography (Riddle �00�). Several attempts are currently under development to find a more integrative research program in biogeography (e.g. Salomon 2001, Donoghue & Moore �00�), but they seem to have little impact on the majority of biogeographers, each protecting his/her own scientific niche or summit. “We believe that the best means for advancing the frontiers of our science is to foster reintegration and reticulations among complementary research programs. The new series of synthesis –more complex, scale-variant, and multi-factorial views if how the natural world develops and diversifies– may be less appealing to some researchers, but it is likely to result in a much more realistic and more illuminating view of the complexity of nature” (Lomolino & Heaney �004, p. �). Parenti & Ebach (in prep.) propose a novel approach, a research agenda they call Comparative Biogeography, where they stress “the qualitative nature of biogeography, although many biogeographers use quantitative methods”. Comparative Biogeography is “a method or approach that incorporates Systematic Biogeography, biotic relationships, their classification and distribution” (Parenti & Ebach in prep.). Comparative biogeography thus retain several aspects of former chorology (see section �.�), showing real possibilities for its renewal... just under a new name?

This is in concordance with Hengeveld’s Dynamic Biogeography (Hengeveld �990). The study of dynamic biogeography can be of a quantitative or qualitative nature. These approaches depend on an extensive knowledge of the ecology, biology and local distribution patterns of species and communities, and of interactions of those communities. Both methods are equally sound and should be used in a complimentary fashion. Quantitative methods are a valuable tool in confirming or rejecting initial conclusions and theories based on a qualitative approach.

�79

But the meaning of Qualitative Biogeography goes beyond the specific approaches and join the novel challenge of a science of qualities, that is conscious of the assumptions and limitations inherent to the methods (being these quantitative or qualitative), and more important, that is conscious of the history of the discipline and of the whole scientific task.

Goodwin (�994) and Reason & Goodwin (�999) propose a radical change in the whole research endeavour of the natural sciences, taking into account recent advances in complexity theory. They suggest that this line of thinking leads toward a science of qualities based on participation and intuition, with remarkably similarities to the kinds of knowing which are seen as central in constructionist and participatory approaches to social and organizational life. “Conventional scientists begin to get very nervous when this type of procedure is described as science. They are suspicious of the intuition, and they mistrust qualitative observation. As far as the intuition is concerned, they need have no anxieties: it is a universally recognised subjective component of scientific discovery. It is the intuitive faculty that makes sense of diverse data and brings them into a coherent pattern of meaning and intelligibility, though of course the analytical intellect is also involved in sorting out the logic of the intuitive insight [...] However, scientists are trained to pay attention only to quantities. As people and as naturalists they are aware of qualities, which are often the primary indicators of change. But as scientists they factor them out of their consciousness. This restriction is based on a convention that has worked extremely well for simple systems, but it has severe limitations in the face of complexity. It is time to move into a science of qualities” (Reason & Goodwin �999). A science of emergent qualities involves a break with the positivist tradition that separates facts and values and a re-establishment of a foundation for a naturalistic ethics (Collier �994).

Complexity theory is well tied to the new developments in evolutionary biology, the so called evo-devo, that has gained an unusual impulse during the late 80s and 90s via authors like S.J.Gould, N. Eldredge, F. Varela, or S. Kaufmann, as reviewed by Brockman (�99�).

The re-thinking of the theory of evolution is taken a direction in which natural selection is being abandom, or at least integrated with the still less known principles of the auto-organisation of life (Varela �000, García Azkonobieta �00�). This development will soon encompass systematic problems in plant biology (e.g. Frohlich �00� in the recent volume on monocots evolution). This is a good sign toward the better integration between evolutionary biology, cladistics and systematics. Goodwin classify this advances as a new biology: “The new biology is biology in the form of an exact science of complex systems concerned with dynamics and emergent order. Then everything in biology changes. Instead of the metaphors of conflict, competition, selfish genes, climbing peaks in fitness landscapes, what you get is evolution as a dance. It has no goal. As Stephen Jay Gould says, it has no purpose, no progress, no sense of direction. It’s a dance through morphospace, the space of the forms of organisms” (Goodwin in Brockman �99�, p. 97)�0

�80

Did this development in evolutionary biology (i.e the new biology) already reach the new biogeography? (in the sense of Lomolino & Heaney �004). Are evolutionary scenarios been taken explicitly in the biogeographical discussion? And if not, will this happen during the next decade? Some authors are still skeptic about the recent developments: “…contemporary biogeography, especially phylogeography, is simply boring to read because the mindless slaves that produce it ignore Horace’s dictum sapere aude (Epistle I, �) –”dare to be wise”– and are not bold enough to produce anything new. Their only novelties are timid variations, and the trite stories they tell involve the same old ideas that have been worked and reworked for over �000 years. Ancient teleology is replayed every night on television, and mysterious means of dispersal and perfect adaptations are extolled as wonders of nature” (Heads �00�a, p. ���). One possibility for the sceptical is just turn off the switch (figure 7.3)… We just can, with all these concerns and conflicts in mind, keep going, asking and (re)searching. Let’s take Reason & Goodwin’s positive view, together with Gyerin pragmatic one: “…may the best science win…” (Gieryn 1999, title of chapter �).

Today a successful scientific career in evolutionary biology often seems to require few original or fundamental discoveries and much creative scenario building based more on other theories than on descriptive data. I don’t want to speculate on wether this is good or bad, but I believe that without discovery of more repeating patterns in systematics, those of us interested in evolutionary process and its connection to phylogenetic pattern will be stuck in a rut… Of course it is possible that we may never discover such constants in biology, and a smoother synthesis of pattern and process will never be made, but I remain an optimistic. I also unambiguously identify my optimism as a result of my beliefs rather than of scientific evidence. Distinguishing between one’s beliefs and one’s scientific evidence may be one of the most difficult aspects of evolutionary studies” (Grande 1994, p. 79).

Figure 7.� Valparaíso (photo courtesy M. Richter)

�8�

8 Conclusions: toward a biogeographic synthesis of the Chilean Flora

�8�

From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December �00�

�8�

8 Conclusions: toward a biogeographic synthesis of the Chilean Flora

Only one conclusion seems possible… The sciences of geobotany and geozoology carry a heavy burden of deductive reasoning. What is most needed in these fields is a complete return to inductive reasoning, with assumptions reduced to a minimum and hypotheses based upon demonstrable facts and proposed only when necessary. In many instances the assumptions arising from deductive reasoning have so thoroughly permeated the science og geography and have so long been a part of its warp and woof that students of the field can distinguish fact from fiction only with difficulty” (Cain �944, pp. ��0-���).

• The extant vascular flora of Chile is composed, under current knowledge (updated 09 February �007) of 59 orders, �79 families, 8�� genera, and about 4 ��� species.

• This account considers three endemic families (Gomortegaceae, Francoaceae, and Lactoridaceae), and 83 endemic genera registered in the Chilean vascular flora.

• Best represented families in Chile are the Asteraceae, Poaceae, Apiaceae, Brassicaceae, and Fabaceae; species-richest genera are Senecio, Adesmia, Oxalis, Viola, and Haplopappus.

• The Chilean vascular flora has its roots in the Silurian (~430 mya), the angiosperm flora in Jurassic/Cretaceous times (~150 mya), or maybe already in the Permian (~260 mya)?

• The Chilean flora has geographical relations with diverse floras. Seven floristic elements were identified: pantropical, australasiatic, neotropical, antitropical, south-temperate, endemic, and (sub)cosmopolitan, some of them refined as 9 generalized tracks representing mostly disjunct distributions. The best represented floristic element is the neotropical, followed by the antitropical and the cosmopolitan.

• The neotropical element has the northernmost distribution average at �9,�°S, while the south-temperate and australasiatic elements show the southernmost average at 4�,�°S and 4�,�°S respectively. The other elements show an average distribution in Central Chile between ��,�° and ��°S.

• The placement of the Chilean flora in the global classification have been, since the earliest intends, a controversial task. The replacement of the neotropical floristic element by the autralasiatic element in Magallanes region, show the consistency of classifying southernmost Chile in an austral floristic realm.

�84

• North and Central Chile belong to the neotropical floristic realm, while Central Chile represents a core area of endemism or a panbiogeographic node.

• Integrationg palaeobotanical and geological data under the current plate tectonics paradigm, the most logic model for the Chilean flora is an interchange of tropical floras with australasiatic floras, with later development of a subtropical more xeric flora.

• The geographic ranges of many Chilean plants have been reduced by the glaciations of the Pleistocene, but it is doubtfull that this relative recent events had an impact on speciation processes.

• The Pacific offshore islands have been traditionally classified as oceanic islands, but the floristic composition of Juan Fernández and its floristic similarity with the Magallanes region, is a strong sign to consider it as a continental flora, maybe the remnant of an ancient flora of an older terrestrial landmass to the West of current South America (recognized by some geologists as the Pacifica lost continent).

• Alternative palaeogeographic scenarios for the islands and for the continent as well, should not be rouled out till much of the evidence is confronted.

• Also alternative views in the evolution of plants (specially angiosperms) can change dramatically our understanding of biogeography and the relation to palaeogeographic events.

• Conflicts in tectonic hypotheses, systematics, and molecular dating, directly affect any proposal in biogeography. Being biogeography fragmented in the last decades into a plethora of methods, a synthetical biogeography will be still elusive and murky for decades. This synthesis will not come from the scientific consensus but from a deep reanalysis of the conceptual and epistemological basis of the discipline, in connection with new developments and debates in complexity theory and postmodern science.

• The biogeography and evolution of the southern hemisphere’s flora will continue

fascinating naturalists as it has been the case for more than ��0 years. The view from the top of Cerro La Campana in Central Chile and from El Camote in Juan Fernández releases the inspiration for an intuitive (re)search in qualitative biogeography.

• The plant geography of Chile is (and will be ever) a work in progress.

�8�

“Trapped between fundamentalists, who believe they have found truth, and relativists, who refuse to pin it down, the bewildered majority in between continues to hope there is a truth worth looking for, without knowing how to go about it or how to answer the voices from either extreme. We need a new Guide for the Perplexed – a way of understanding and identifying truth which can survive the postmodern era” (Fernández-Armesto �997, p. �).

The challenge of doing a synthesis of the Chilean Plant Geography is as difficult as 100 years ago as done by Reiche (1907). Many aspects may still not reflect the truth of the Chilean Plant Geography, but the risk hides precisely in the thought that one has reached such an elusive and dogmatic think like the truth. Some scientists are loyal to their respective schools and scholar background, other are just loyal to their intrinsic curiosity. Some scientists continue the long tradition of discovering new worlds, and in the meantime the discoveries turn to the continuous re-invention of the world (Kennedy �00�).

“But every ambitious exercise in critical geographical description, in translating into words the encompassing and politicized spatiality of social life, provoke a linguistic despair. What one sees when one looks at geographies is stubbornly simultaneous, but language dictates a sequential succession, a lineal flow of sentential statements bound by the most spatial of Earthly constraints, the impossibility of two objects (or words) occupying the same precise place (as o a page). All that we can do is re-collect and creatively juxtapose, experimenting with assertions and insertions of the spatial against the prevailing grain of time. In the end, the interpretation of postmodern geographies can be no more than a beginning.” (Soja �989, p. �).

Personally, my intact curiosity is back to the starting point, equipped with a backpack (what I am saying... five backpacks!) hitchhiking on the Pan-American highway toward the South.

�8�

Footnotes� Hugh M. Raup (a botanist) did a review of the subject for the Annals of the Association of American Geographers. Aside the title of the article, he widely use the term plant geography… Raup’s case as a botanist publishing in a geographical journal shows graphically that the differences in the field are very permeable.

� Areology is also the study of planet Mars, as counterpart to Geology (www.duden.de).

� Note that von Wettstein’s work can be seen as the precursor of the modern concept of allopatric speciation.

4 Many people use the terms ‘systematics’ and ‘taxonomy’ as synonyms, but Stuessy (�979, �00�: ��) clearly illustrates the differences. Griffiths proposed to call the more general concept ‘metasystematics’ (Griffiths 1974: 89.)

� “A character of an organism is a feature that we judge useful in classification (i.e. in making comparisons to establish the degree of relationship) and a character state is one particular aspect of this character. For example, colour of petals might be a character and character states might be yellow, white and pink. One cannot compare characters for establishing relationships; rather we compare character states. Through this process, groups are formed, and these groups are called taxa” (Stuessy �00�, p. �7).

� news in the English newspaper The Independent of ��rd November �998, as quoted by Endersby (�00�)

7 „I continued thus at the adult age my childhood baublery with maps: I connected cities of equivalent size by straight lines, in order to determine whether in the railway or road system certain rules were recognizable, whether there would be a regular transportation network, on the other hand, in order to measure the distances between equally large cities. My maps filled with triangles, often equilateral triangles - the distances between equally large cities were thus almost the same among themselves - which united building hexagons. I continued to state that in South Germany the small towns have very often the very exactly distance of �� km among each other. My goal was marked out: to find the laws that explain the number, size and distribution of cities…” (Christaller 1968, p. 96).

8 Hennig draw this nice comparison in spite of being himself a very bad geographer… see Croizat’s critic on Hennig’s explanation of a progression rule while confounding the islands upon the example rests (Croizat �97�).

9 “We are living on a time in which postmodernism is constantly confronting the hegemonic speech of science to denounce its excesses and to expose its limits”.

�0 Some geographers recognize and promote the critical discussion but still not agree on the appropriate definition (e.g. Spedding & Lorimer, in their course ‘Critical approaches to Geography’ have a lecture titled, “Post-whatever Physical Geography: does it (or should it) exist?” [http://www.abdn.ac.uk/~geo��7/crap.html]

�� “All phytogeographical divisions have to be considered as provisional or approximate. Opinions shall be expressed, but to discuss divisions, limits or nomenclature, or to try to impose ideas in a dogmatic form, will be simply a waste of time and paper without greater benefit for science”.

�� http://www.mnhn.cl/botanica/Herbario/index.html

�� Muñoz Pizarro (�9��) considered also many naturalized taxa.

�87

�4 not considering ferns and fern allies.

�� J.I. Molina is the first Chilean naturalis (see box 2.1), Joseph Banks and George Forster joined James Cook in his first and second great voyages respectively.

�� “I thus hold divisions of the sphere by areas, suggested until now, for artificial systems…. they just harmed science”

�7 “my work remainder completely different from that of which my father thought, because the documents had become more numerous, and my ideas had singularly moved away from those which had reigned in science for twenty years”

�8 Croizat apparently took the term from Van Steenis (Croizat �9��)

�9 Due to human pressure, Santalum fernandezianum is all but extinct from Juan Fernández (Islands).

�0 In fact, in the analysis of the Chilean endemic genera, the exercise of subtracting one or two genera from the data set yielded different consensus trees (not shown).

�� See also: http://www.worldwildlife.org/wildworld/profiles/terrestrial/nt/nt0403_full.html

�� Mantle plume = deep seated structures originating near the core-mantle boundary or originating at relatively shallow depths in the mantles as a response to thermal incubation beneath large continents (Hawkesworth et al. �999). �� Fischer, J.M. (�00�) Beyond the Plume Myth www.newgeology.us

�4 Also the website: “Problems with Plate Tectonics” http://ourworld.compuserve.com/homepages/dp�/lowman.htm

�� Winteraceae have often been regarded as the ‘‘most primitive’’ extant family of angiosperms (Cronquist �98�, Endress �98�). The phylogenetic position based on molecular data indicates that the family is nested in the Canellales (APG II �00�, Soltis & Soltis �004).

�� Brazeau, M. (�00�) http://lancelet.blogspot.com/

�7 Constance does not give the reference to these authors’ papers, but their work is worth of attention.

�8 It is a pity that Müller (�99�) in his excellent revision of Ratzel’s program does not explicit analyze this great piece: Der Lebensraum: eine biogeographische Studie (Ratzel �90�).

�9 Note that this was written �� years before the proposal of continental drift by Alfred Wegener.

�0 Also available at http://www.edge.org/�rd_culture/bios/goodwin.html

�88

ReferencesAagesen L & M Sanso (�00�) The Phylogeny of the Alstroemeriaceae, based on morphology, rps�� Intron, and rbcL sequence data. Systematic Botany, 28, 47-�9.

Al-Shehbaz IA (�00�) A synopsis of Tropidocarpum (Brassicaceae). Novon, 13, �9�-�9�.

Albach DC, HM Meudt, & B Oxelman (�00�) Piecing together the ‘‘new’’ Plantaginaceae. American Journal of Botany, 92, �97–���.

Alpers C & GH Brimhall (�988) Middle Miocene climatic change in the Atacama Desert, northern Chile: Evidence from supergene mineralization at La Escondida. Geol. Soc. Amer., �00, ��40-����.

Amigo J & C Ramírez (1998) A bioclimatic classification of Chile: woodland communities in the temperate zone. Plant Ecology, 136, 9–��.

Anderson CL, JHE Rova & L Andersson (�00�) Molecular Phylogeny of the Tribe Anthospermeae (Rubiaceae): Systematic and Biogeographic Implications. Australian Systematic Botany, 14, ���–�44.

Anderson CL, K Bremer & EM Friis (�00�) Dating phylogenetically basal eudicots using rbcL sequences and multiple fossil reference points. American Journal of Botany 92,�7�7-�748.

Andersson L, M Kocsis & R Eriksson (�00�) Relationships of the genus Azorella (Apiaceae) and other hydrocotyloids inferred from sequence variation in three plastid markers. Taxon, 55, �70-�80.

Ané C & MJ Sanderson (�00�) Missing the forest for the trees: phylogenetic compression and its implications for inferring complex evolutionary histories. Systematic Biology, 54, �4�–��7.

Anker P (�00�) Boundary work from the science war zone. Metascience, 12, 40�-404.

APG [Angiosperm Phylogeny Group] (1998) An Ordinal Classification for the Families of Flowering Plants. Annals of the Missouri Botanical Garden, 85, ���-���.

APG II [Angiosperm Phylogeny Group II] (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Botanical Journal of the Linnean Society, 141, �99–4��.

Applequist WL & RS Wallace (�00�) Phylogeny of the portulacaceous cohort based on ndhF sequence data. Systematic Botany, 26, 40�–4�9.

Arldt T (�9�8) Die Entwicklung der Kontinente und ihrer Lebewelt (�d. ed.) Berlin.

Arroyo MTK, C Castor, C Marticorena, M Muñoz Schick, L Cavieres, O Matthei, FA Squeo, M Grosjean & R Rodríguez (1998). The flora of Llullaillaco National Park located in the transitional winter-summer rainfall area of the northern Chilean Andes. Gayana Botánica 55, 9�-��0.

Arroyo MTK, O Matthei, C Marticorena, M Muñoz Schick, F Perez & AM Humaña (2000) The vascular plant flora of the Bellotos del Melado National Reserve, VII Region. Chile: a documented checklist. Gayana Botánica 57, ��7-��9.

Arroyo MTK, C Marticorena, O Matthei, M Muñoz Schick & P Pliscoff (2002) Analysis of the contribution and efficiency of the Santuario de la Naturaleza Yerba Loca, 33º S in protecting the regional vascular plant flora (Metropolitan and Fifth regions of Chile). Revista Chilena de Historia Natural, 75, 7�7-79�.

Arroyo MTK, O Matthei, M Muñoz-Schick, J Armesto, P Pliscoff, F Perez & C Marticorena (�00�) Flora de cuatro reservas nacionales en la Cordillera de la Costa, ��º-��º S, VII Región, Chile y su papel en la protección de la biodiversidad regional. Biodiversidad y Ecología de los Bosques de la Cordillera de la Costa de Chile (ed. by C Smith, JJ Armesto & C Valdovinos), pp. ���-�44. Editoral Universitaria, Santiago.

Ashworth AC & DJ Cantrill (�004) Neogene vegetation of the Meyer Desert Formation (Sirius Group) Transantarctic Mountains, Antarctica. Palaeogeography, Palaeoclimatology, Palaeoecology, 213, ��-8�.

Attwell CAM & FPD Cotterill (�000) Postmodernism and African conservation science. Biodiversity and Conservation, 9, ��9–�77.

�89

Axelrod DI (�9��) How old are the angiosperms? American Journal of Science, 259, 447-4�9.

Backeberg C (�94�) Cactaceae. Jahrbücher der DKG, Teil �, Juni �94�: ��, Cactaceae Lindley; Systematische Übersicht (Neubearbeitung) mit Beschreibungsschlüssel.

Bader FJW (�9�0) Die Verbreitung borealer und subantarktischer Holzgewächse in den Gebirgen des Tropengürtels. Nova Acta Leopoldina, 23, �-�44.

Barbour M (�9�9) Patterns of genetic similarity between Larrea divaricata of North and South America. American Midland Naturalist, 81, �4–�7.

Barbour T (�9��) Some Remarks upon Matthew’s ‘Climate and Evolution’. Annals of the New York Academy of Sciences, 27, �-�0. [http://www.wku.edu/~smithch/biogeog/ BARB�9��.htm]

Barker PF, IWD Dalziel, BC Storey (�99�) Tectonic development of the Scotia Arc region. The geology of Antarctica. (ed. by RJ Tingey) Oxford Monographs on Geology and Geophysics, 17, ���–�48.

Barlow BA (1981) The Australian flora: its origin and evolution. Flora of Australia vol. 1 (ed. by Bureau of Flora and Fauna), pp. ��-7�. Canberra, Australia.

Barreda V & S Archangelsky (�00�) The southernmost record of tropical pollen grains in the mid-Cretaceous of Patagonia, Argentina. Cretaceous Research, 27, 778-787.

Barthlott W (�98�) Biogeography and evolution in neo- and paleotropical Rhipsalinae (Cactaceae). Sonderb. Naturwiss. Verl. Hamburg, 7, �4�-�48.

Barthlott W & DR Hunt (�99�) Cactaceae. The families and genera of vascular plants vol. II. (ed. by K Kubitzki), pp. ���-�97. Springer, Berlin.

Barthlott W & NP Taylor (�99�) Notes towards a monograph of Rhipsalideae (Cactaceae). Bradleya, 13, 4�-79.

Baumann F (1988) Geographische Verbreitung und Ökologie südamerikanischer Hochgebirgspflanzen. Physische Geographie (Zürich), 28, �-�0�.

Beier BA, JAA Nylander, MW Chase & M Thulin (�004) Phylogenetic relationships and biogeography of the desert plant genus Fagonia (Zygophyllaceae), inferred by parsimony and Bayesian model averaging. Molecular Phylogenetics and Evolution, 33, 9�-�08.

Bell CD & MJ Donoghue (�00�) Phylogeny and biogeography of Valerianaceae (Dipsacales) with special reference to the South American valerians. Organisms Diversity & Evolution, 5, �47-��9.

Bell CD, DE Soltis & PS Soltis (�00�) The age of the angiosperms: a molecular timescale without a clock. Evolution 59, ��4�–���8.

Beloussov VV (�9�8) Some General Aspects of Development of the Tectonosphere. Rep. 28. International Geol. Congress, 1, 9-�7.

Benton MJ & FJ Ayala (�00�) Dating the tree of life. Science, 300, ��98-�700.

Benton MJ, MA Wills & R Hitchin (2000) Quality of the fossil record through time. Nature, 403, ��4–��7.

Berlocher SH (�998) Can Sympatric Speciation via Host or Habitat Shift Be Proven from Phylogenetic and Biogeographic Evidence? Endless Forms: Species and Speciation (ed. by DJ Howard & SH Berlocher), pp. 99-���, Oxford University Press.

Bernardello G, GJ Anderson, TF Stuessy & DJ Crawford (2006) The angiosperm flora of the Archipelago Juan Fernandez (Chile): origin and dispersal. Canadian Journal of Botany, 84, ����–��8�.

Berry EW (1920) Palaeobotany: a sketch of the origin and evolution of floras. Ann. Rep. Smithson. Inst., 1918, �89-407.

Bews JW (�9��) Some General Principles of Plant Distribution as illustrated by the South African Flora. Annals of Botany, 25, �-��.

�90

Blackburn TM & KJ Gaston (eds.) (�00�) Macroecology: Concepts and Consequences. British Ecological Society Annual Symposium Series. Blackwell Science, Oxford.

Bleeker W, A Franzke, K Pollmann, AHD Brown & H Hurka (�00�a) Phylogeny and biogeography of Southern Hemisphere high-mountain Cardamine species (Brassicaceae).Australian Systematic Botany, 15, �7�–�8�.

Bleeker W, C Weber-Sparenberg & H Hurka (�00�b) Chloroplast DNA Variation and Biogeography in the genus Rorippa Scop. (Brassicaceae). Plant Biology, 4, �04-���.

Bonifacino JM & G Sancho (�004) Guynesomia (Asteraceae: Astereae), a new genus from central Chile. Taxon, 53, �7�-�78.

Bork H-R (�00�) Landschaften der Erde unter dem Einfluss des Menschen. Wissenschaftliche Buchgesellschaft (WGB), Darmstadt.

Bowe LM, G Coat & CW de Pamphilis (�000) Phylogeny of seed plants based on all three genomic compartments: Extant gymnosperms are monophyletic and Gnetales‘ closest relatives are conifers. PNAS, 97, 409�-4097.

Bradford JC & RW Barnes (2001) Phylogenetics and classification of Cunoniaceae (Oxalidales) using chloroplast DNA sequences and morphology. Systematic Botany, 26, ��4–�8�.

Brako L & JL Zarucchi (�99�) Catalogue of the Flowering Plants and Gymnosperms of Peru. Reprint �99�. Monographs in Systematic Botany from the Missouri Botanical Garden, 45, �-��8�.

Bray WL (1900) The relations of the North American flora to that of South America. Science, 12, 709–7��.

Bremer K (�994) Asteraceae: cladistics and classification. Portland, Oregon. Timber Press.

Bremer K (2000) Early Cretaceous lineages of monocot flowering plants. PNAS, 97, 4707-47��.

Bremer K (�00�) Gondwanan evolution of the grass alliance of families (Poales). Evolution 56, ��74-��87.

Bremer K & MHG Gustafsson (1997) East Gondwana ancestry of the sunflower alliance of families. PNAS, 94, 9�88-9�90.

Bremer K & T Janssen (�00�) Gondwanan origin of major monocot groups inferred from dispersal-vicariance analysis. Aliso, 22, ��-�7.

Briggs JC (�004) The ultimate expanding earth hypothesis. Journal of Biogeography, 31, 8��–8�7.

Britton NL & JN Rose (�9�9–�9��) The Cactaceae. 4 volumes. Carnegie Inst., New York.

Britton T, B Oxelman, A Vinnersten & K Bremer (2002) Phylogenetic dating with confidence intervals using mean path lengths. Molecular Phylogenetics and Evolution, 24, �8-��.

Brockman J (�99�) The Third Culture: Beyond the Scientific Revolution. Simon & Schuster, New York.

Brown B, C Gaina & RD Müller (�00�) Circum-Antarctic palaeobathymetry: Illustrated examples from Cenozoic to recent times. Palaeogeography, Palaeoclimatology, Palaeoecology, 231, ��8-��8.

Brown JH (�99�) Macroecology. U. Chicago Press, Chicago.

Brown JH & MV Lomolino (�000) Concluding remarks: historical perspective and future of island biogeography theory. Global Ecology and Biogeography 9, 87–9�.

Brown RW (�9��) Palm-like plants from the Dolores Formatioon (Triassic), southwestern Colorado. U.S. Geol. Surv. Prof. Pap. 274, �0�-�09.

Brüggemann H (�000) Molekulare Systematik und Biogeographie der Gatttung Lepidium L. und Verwandter Sippen (Brassicaceae). Dissertation im Fachbereich Biologie/Chemie der Universität Osnabrück.

Brüggen J (�9�0) Fundamentos de la Geología de Chile. Inst. Geogr. Militar, Santiago, Chile.

�9�

Bunge W (�9��) Theoretical Geography. Lund Studies in Geography, Gleerup, Lund.

Burckhardt C (1902) Traces géologiques d‘un ancient continent Pacifique. Rev. Mus. La Plata, 10, �79-�9�.

Burleigh JG & S Mathews (�004) Phylogenetic signal in nucleotide data from seed plants: implications for resolving the seed plant tree of life. American Journal of Botany, 91, ��99-����.

Burtt BL (�998) Climatic accommodation and phytogeography of the Gesneriaceae of the Old World. Diversity and taxonomy of tropical flowering plants (ed by R Mathew & M Sivadasan), pp. �-�7, Mentor Books, Kerala, India.

Bush GL (�97�) Modes of animal speciation. Annual Review of Ecology and Systematics, 6, ��9-��4.

Bush GL (�998) The Conceptual Radicalization of an Evolutionary Biologist. Endless Forms: Species and Speciation (ed. by DJ Howard & SH Berlocher), pp. 4��-4�8, Oxford University Press.

Buxbaum F (�9�9) Die Entwicklungswege der Kakteen in Sudamerika. Biogeography and ecology in South America. (ed. by EJ Fittkau, J Illies, H Klinge, GH Schwabe & H Sioli), pp. �8�–���. W. Junk, The Hague, Netherlands.

Cabrera AL (1953) Esquema fitogeográfico de la República Argentina. Revista del Museo Eva Perón, Botánica 8, 87–��8.

Cabrera AL (�9��) The genus Lagenophora (Compositae). Blumea, 14, �8�-�08.

Cabrera AL & A Willink (�97�) Biogeografía de América Latina. Monografía ��. Serie de Biología. Secretaría General de la Organización de los Estados Americanos. Washington. �d revised ed. �980.

Cain SA (�944) Foundations of plant geography. Harper & Brothers, New York, London.

Camp WH (�947) Distribution Patterns in Modern Plants and the Problem of Ancient Dispersals. Ecological Monographs, 17, ��9-�8�.

Cañas J (�9��) Tectónica y sedimentaci�n de la Formación Los Molles, Prov. de Aconcagua. Comun. Escuela de Geol. (Santiago), 11, ��.

Carey SW (�988) Theories of the Earth and the Universe: a History of Dogma in the Earth Science. Stanford University Press, Stanford.

Carlquist S (�974) Island biology. Columbia University Press, New York.

Carlquist S (�98�) Intercontinental dispersal. Dispersal and Distribution: An International Symposium (ed. by K Kubitzki) Sonderbände des Naturwissenschaftlichen Vereins in Hamburg, 7, �7–47, Verlag P. Parey, Hamburg/Berlin.

Castro SA, A Camousseight, M Muñoz-Schick & FM Jaksic (�00�) Rodulfo Amando Philippi, el naturalista de mayor aporte al conocimiento taxonómico de la diversidad biológica de Chile. Rev. Chil. Hist. Nat., 79, ���-�4�.

Cavieres LA, Arroyo MTK, Posadas P, Marticorena C, Matthei O, Rodriguez R, Squeo FA & Arancio G (�00�) Identification of priority areas for conservation in an arid zone: application of parsimony analysis of endemicity in the vascular flora of the Antofagasta region, northern Chile. Biodiversity and Conservation, 11, ��0�-����.

Chacón J, S Madriñán, MW Chase & JJ Bruhl (�00�) Molecular phylogenetics of Oreobolus (Cyperaceae) and the origin and diversification of the American species. Taxon, 55, ��9-���.

Chanderbali AS, H van der Werff & SS Renner (�00�) Phylogeny and historical biogeography of Lauraceae: evidence from the chloroplast and nuclear genomes. Annals of the Missouri Botanical Garden, 88, �04–��4.

Chorley RJ (�978) Bases for theory in geomorphology Geomorphology: Present Problems and Future Prospects (ed. by C Embleton, D Brunsden & DKC Jones), pp. �-��, Oxford University Press.

Christ H (1867) Über die Verbreitung der Pflanzen der alpinen Region der europäischen Alpenkette. Denkschr. Schweiz. Nat. Ges. vol. 22.

�9�

Christaller W (�9�8) Wie ich zur Theorie der zentralen Orte gekommen bin. Geographische Zeitschrift, 56, 88-�0�.

Chung KF, CI Peng, SR Downie, K Spalik & BA Schaal (2005) Molecular systematics of the trans-Pacific alpine genus Oreomyrrhis (Apiaceae): Phylogenetic affinities and biogeographic implications. American Journal of Botany, 92, �0�4-�07�.

Collier A (�994) Critical Realism: An Introduction to Roy Bhaskar’s Philosophy. Verso, London.

Constance L (1963) Amphitropical relationships in the herbaceous flora of the Pacific Coast of North and South America: a symposium. Introduction and historical review. Quarterly Review of Biology, 38, �09-���.

Cook LG & MD Crisp (�00�) Not so ancient: the extant crown group of Nothofagus represents a post-Gondwanan radiation. Proceedings of the Royal Society B-Biological Sciences, 272, ����-��44.

Cope TA (�000) Chorology of Grasses – A Review. Grasses: Systematics and Evolution (ed. by SWL Jacobs & J Everett), pp. ���-��0, CSIRO, Melbourne.

Cornet B (�989) The reproductive morphology and biology of Sanmiguelia lewisii, and its bearing on angiosperm evolution in the Late Triassic. Evol. Trends in Plants, 3, ��-��.

Cornet B (�00�) (updated 0�/08/0�) When Did Angiosperms First Evolve? [http://www.sunstar-solutions.com/sunstar/Why0�/why.htm]

Cotterill FPD (�999). Review of Principles of Conservation Biology, �nd Edition (ed. by GK Meffe & CR Carroll), Biodiversity and Conservation, 8, �7��-�7�4.

Cowling RM, PW Rundel, BB Lamont, MTK Arroyo & M Arianoutsou (�99�) Plant diversity in Mediterranean-climate regions. Trends in Ecology & Evolution, 11, ���-���.

Cox CB (�990) New geological theories and old biogeographical problems. Journal of Biogeography, 17, ��7-��0.

Cox CB (�00�) The biogeographic regions reconsidered. Journal of Biogeography, 28, ���-���.

Cox CB & PD Moore PD (�00�) Biogeography: An Ecological and Evolutionary Approach. 7th edition. Blackwell Publishing.

Cracraft J (�00�) The seven great questions of systematic biology: An essential foundation for conservation and the sustainable use of biodiversity. Annals of the Missouri Botanical Garden, 89, ��7-�44.

Crane PR (�98�) Phylogenetic analysis of seed plants and the origin of angiosperms. Annals of the Missouri Botanical Garden, 72, 7��–79�.

Crane PR (�99�) The fossil history of the gnetales. International Journal of Plant Sciences, 157, (Suppl. S), S�0-S�7.

Crane PR, P Herendeen & EM Friis (�004) Fossils and plant phylogeny. American Journal of Botany, 91, ��8�-��99.

Craw R (�988) Continuing the synthesis between Panbiogeography, phylogenetic systematics and geology as illustrated by empirical studies on the biogeography of New Zealand and the Chatham Islands. Systematic Zoology, 37, �9�-��0.

Craw R & G Sermonti (eds.) (�988) Space, time, form. Panbiogeography symposium. Rivista di Biologia (Perugia), 8�.

Craw RC, JR Grehan & MJ Heads (�999) Panbiogeography: Tracking the History of Life. Oxford Biogeography Series. Oxford University Press.

Crawford DJ, TF Stuessy, TG Lammers, M Silva O & P Pacheco (�990) Allozyme variation and evolutionary relationships among three species of Wahlenbergia (Campanulaceae) in the Juan Fernandez islands. Botanical Gazette, 151, ��9-��4.

Crawford D, M Tago-Nakazawa,TF Stuessy, GJ Anderson, G Bernardello, E Ruiz, RJ Jensen, CM Baeza, AD Wolfe & M Silva O (�00�) Intersimple sequence repeat (ISSR) variation in Lactoris fernandeziana (Lactoridaceae), a rare endemic of the Juan Fernández Archipelago, Chile. Plant Species Biology, 16, �8�–�9�.

�9�

Crayn DM, KA Kron, PA Gadek & CJ Quinn (1998) Phylogenetics and evolution of epacrids: a molecular analysis using the plastid gene rbcL with a reappraisal of the position of Lebetanthus. Australian Journal of Botany, 46, �87-�00.

Crepet WL (�998) The Abominable Mystery. Science, 282, ����-���4

Crepet WL (�000) Progress in understanding angiosperm history, success, and relationships: Darwin’s abominably “perplexing phenomenon”. PNAS, 97, ��9�9-��94�.

Crepet WL, KC Nixon & MA Gandolfo (�004) Fossil evidence and phylogeny: The age of major angiosperm clades based on mesofossil and macrofossil evidence from Cretaceous deposits. American Journal of Botany, 91, ����-��8�.

Crisci JV, MM Cigliano, JJ Morrone & S Roig-Juñent (�99�) Historical biogeography of southern South America. Systematic Zoology, 40, ���-�7�.

Crisci JV, L Katinas & P Posadas (�00�) Historical biogeography: an introduction. Harvard University Press, Cambridge, MA.

Crisp, MD, JG West & HP Linder (1999) Biogeography of the terrestrial flora. Flora of Australia vol. 1, �d edition (ed. by A.E. Orchard), pp. ���-��7, Australian Biological Resources Study, Canberra.

Croizat L (1945) History and nomenclature of the higher units of classification. Bulletin of the Torrey Botanical Club, 72, ��-7�.

Croizat L (�9��) Manual of Phytogeography, or an account of plant dispersal throughout the world. Junk, The Hague.

Croizat L (�9�8) Panbiogeography or an introductory synthesis of zoogeography, phytogeography, and geology; with notes on evolution, systematics, ecology, anthropology, etc. (� vol.) Published by the author, Caracas, Venezuela.

Croizat L (�9�0) Principia Botanica. Published by the author, Caracas, Venezuela.

Croizat L (�9��) Space, Time, Form: The Biological Synthesis. Published by the Author, Caracas, Venezuela.

Croizat L (�97�) Biogeografía analítica y sintética (“Panbiogeografía”) de las Américas. Biblioteca de la Academia de Ciencias Físicas, Matemáticas y Naturales. Caracas.

Cruden RW (�9��) Birds as agents of long-distance dispersal for disjunct plant groups of the temperate Western Hemisphere. Evolution, 20, ��7–���.

Cuevas JG & G van Leersum (�00�) Proyecto conservación, restauración y desarrollo de las islas Juan Fernández, Chile. Revista Chilena de Historia Natural, 74, 899-9�0.

Cullen J & SM Walters (�00�) Flowering-plant families: how many do we need? Taxonomy and Plant Conservation (ed. by E Leadlay & S Jury), pp. 4�-9�, Cambridge Univ. Press.

Daghlian CP (�98�) A review of the fossil record of monocotyledons. Botanical Review, 47, ��7-���.

Danton Ph, E Breteau & M Baffray (�000) Les îles de Robinson: Trésor vivant des mers du Sud, entre légende et réalité. Nathan, Paris.

Danton Ph & Ch Perrier (2003) Liste de la flore vasculaire de l’île Robinson Crusoe, archipel Juan Fernández, Chili. Le Journal de Botanique de la Société Botanique de France, 24, �7-78.

Darwin Ch (�8�9) The Voyage of the Beagle. John Murray, London. Darwin Ch (�8�9) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. Watts & Co., London.

Davies TJ, TG Barraclough, MW Chase, PS Soltis, DE Soltis & V Savolainen (�004) Darwin’s abominable mystery: Insights from a supertree of the angiosperms. PNAS, 101, �904-�909.

�94

Davis CC, WR Anderson & MJ Donoghue (�00�) Phylogeny of Malpighiaceae: evidence from chloroplast ndhF and trnL-F nucleotide sequences. American Journal of Botany, 88, �8�0–�84�.

Davis CC, WR Anderson & KJ Wurdack (2005) Gene transfer from a parasitic flowering plant to a fern. Proceedings of the Royal Society B-Biological Sciences, 272, ���7-��4�.

Davis PH & VH Heywood (�9��) Principles of Angiosperm Taxonomy. Oliver & Boyd, Edinburgh & London.

Dawkins R (�989) The Selfish Gene. �d ed., Oxford University Press, New York.

Dawkins R (�998) Postmodernism disrobed. Nature, 394, �4�-�4�.

Dawson, G. �9�8. Las tribus y generos de Escrofulariaceas representados en Austro-América y su distribución geográfica. Rev. Mus. La Plata (Bot.), 59, �0�-��8.

de Candolle, Alphonse (�8��) Introduction à l’étude de la Botanique I, II. Librairie Encyclopédique de Roret, Paris.

de Candolle, Alphonse (�8��) Géographie botanique raisonnée, �. vols. Paris.

de Candolle, Augustin-Pyramus (�8�0) Geographie Botanique. Dictionnaire des sciences naturelles XVIII, ��9-4��.

de Queiroz A (2005) The resurrection of oceanic dispersal in historical biogeography. Trends in Ecology and Evolution, 20, �8–7�.

de Queiroz K (2006) The PhyloCode and the distinction between taxonomy and nomenclature. Systematic Biology, 55, ��0-���.

de Queiroz K & PD Cantino (2001) Phylogenetic nomenclature and the PhyloCode. Bulletin of Zoological Nomenclature, 58, ��4-�7�.

Dear M & G Wassmansdorf (�99�) Postmodern consequences. Geographical Review, 83, ���-���.

Denton GH, CJ Heusser, TV Lowell, PI Moreno, BG Andersen, LE Heusser, C Schlüchter & DR Marchant (�999) Interhemispheric linkage of paleoclimate during the Last Glaciation. Geografiska Annaler, 81A, �07-���.

DeVore ML & TF Stuessy (�99�) The place and time of origin of the Asteraceae, with additional comments on the Calyceraceae and Goodeniaceae. Advances in Compositae Systematics, vol. 1 (ed. by DJN Hind, C Jeffrey & GV Pope), pp. ��-40, Royal Botanic Gardens, Kew.

di Castri F (�9�8) Esquisse écologique du Chili. Biologie del’ Amérique Australe (ed. by C. Delamanre-Debouteville & E. Rapoport), vol. 4, 6-52. Editions du Centre National de la Recherce Scientfique, Paris.

di Castri F & E Hajek (�97�) Bioclimatología de Chile. Vicerrectoría Académica, Universidad Católica de Chile, Santiago.

Dickerman AW (�998) Generalizing phylogenetic parsimony from the tree to the forest. Systematic Biology, 47, 4�4–4��

Dickins JM, DR Choi & AT Yeates (�99�) Past distribution of oceans and continents. New Concepts in Global Tectonics (ed. by S Chatterjee & N Hotton III), pp. �9�-�99. Texas Tech. University Press.

Dickins JM (�99�) The Southern Margin of Tethys. Gondwana Nine vol. 2, pp. ����-���4. Ninth International Gondwana Symposium, Hyderabad, India, �0-�4 Jan �994. Geological Survey of India.

Diels L (�908) Pflanzengeographie. Sammlung Göschen, Göschen’sche Verlagshandlung, Leipzig.

Dillon MO, M Nakazawa & S Leiva González (�00�) The Lomas Formations of Coastal Peru: Composition and Biogeographic History. El Niño in Peru: Biology and Culture Over 10.000 Years (ed. by J Hass & MO Dillon), pp. �-9. Fieldiana Botany New Series N° 4�, Field Museum of Natural History.

Dirnböck T, J Greimler, P Lopez & TF Stuessy (�00�) Predicting future threats to the native vegetation of Robinson Crusoe Island, Juan Fernandez Archipelago, Chile. Conservation Biology, �7, ���0–���9.

�9�

Dobson JE (�99�) Spatial Logic in Paleogeography and the Explanation of Continental Drift. Annals of the Association of American Geographers, 82, �87-�0�.

Dobzhansky T (�9��) Genetics and the Origin of Species. Columbia University Press, New York.

Domínguez MC, S Roig-Juñent, JJ Tassin, FC Ocampo & GE Flores (�00�) Areas of endemism of the Patagonian steppe: an approach based on insect distributional patterns using endemicity analysis. Journal of Biogeography, 33, ���7–���7.

Donato M, P Posadas, DR Miranda-Esquivel, EO Jaureguizar & G Cladera (�00�) Historical biogeography of the Andean region: evidence from Listroderina (Coleoptera : Curculionidae : Rhytirrhinini) in the context of the South American geobiotic scenario. Biological Journal of the Linnean Society, 80, ��9-���.

Donoghue MJ & BR Moore (�00�) Toward an integrative historical biogeography. Integrative and Comparative Biology, 43, ���–�70.

Dowsett HJ, JA Barron, RZ Poore, RS Thompson, TM Cronin, SE Ishman & DA Willard (�999) Middle Pliocene Paleoenvironmental Reconstruction: PRISM2. U.S. Geological Survey Open File Report, 99-��� [http://pubs.usgs.gov/of/�999/of99-���/]

Drude O (�884) Die Florenreiche der Erde: Darstellung der gegenwärtigen Verbreitungsverhältnisse der Pflanzen: Ein Beitrag zur vergleichenden Erdkunde. Ergänzungsheft N° 74, Petermanns Mitteilungen, Gotha, Justus Perthes.

Drude O (�887) Atlas der Pflanzenverbreitung. Berghaus’ Physikalischer Atlas, Abteilung V, Gotha, Justus Perthes.

Drude O (�890) Handbuch der Pflanzengeographie. Bibliothek Geographischer Handbücher (ed. by F Ratzel), Verlag J. Engelhorn, Stuttgart.

Drummond AJ, SYW Ho, MJ Phillips & A Rambaut (2006) Relaxed Phylogenetics and Dating with Confidence. PLoS Biol, 4, e88.

Du Rietz GE (�940) Problems of bipolar plant distribution. Acta Phytogeogr. Suecica, 13, ���-�8�.

Dunai TJ, GA González López & J Juez-Larré (�00�) Oligocene–Miocene age of aridity in the Atacama Desert revealed by exposure dating of erosion-sensitive landforms. Geology, 33, ���-��4.

Ebach MC & CJ Humphries (�00�) Cladistic biogeography and the art of discovery. Journal of Biogeography, 29, 4�7–444.

Ebach MC & CJ Humphries (�00�) Ontology of biogeography. Journal of Biogeography, 30, 9�9–9��.

Ebach MC & DF Goujet (2006) The first biogeographical map. Journal of Biogeography, 33, 7��–7�9.

Ebach MC & RS Tangney (�007) Biogeography in a Changing World. Systematics Association Special Vol., CRC Press.

Ebach MC, CJ Humphries & DM Williams (�00�) Phylogenetic biogeography deconstructed. Journal of Biogeography, 30, ��8�–��9�.

Eddie WMM, T Shulkina, J Gaskin, RC Haberle & RK Jansen (�00�) Phylogeny of Campanulaceae s. str. inferred from ITS sequences of nuclear ribosomal DNA. Annals of the Missouri Botanical Garden, 90, ��4–�7�.

Edwards EJ, R Nyffeler & MJ Donoghue (�00�) Basal cactus phylogeny: implications of Pereskia (Cactaceae) paraphyly for the transition to the cactus life form. American Journal of Botany, 92, ��77-��88.

Eiler J, C Garzione & P Ghosh (�00�) Response to Comment on “Rapid Uplift of the Altiplano Revealed Through ��C-�8O Bonds in Paleosol Carbonates”. Science, 314, 7�0c.

Eklund H (2003) First Cretaceous flowers from Antarctica. Review of Palaeobotany and Palynology, 127, �87-��7.

Eldredge N (�99�) Reinventing Darwin. John Wiley, New York.

�9�

Eldredge N (�004) Why we do it: Rethinking Sex and the Selfish Gene. W.W. Norton, New York.

Eldredge N & SJ Gould (�97�) Punctuated equilibria: an alternative to phyletic gradualism. Models in Paleobiology (ed. by TJM Schopf), pp. 8�-���, Freeman Cooper, San Francisco.

Encinas A, V Maksaev, L Pinto, J P Le Roux, F Munizaga & M Zentilli (�00�) Pliocene lahar deposits in the Coastal Cordillera of central Chile: Implications for uplift, avalanche deposits, and porphyry copper systems in the Main Andean Cordillera. Journal of South American Earth Sciences, 20, ��9-�8�.

Endersby J (�00�) ‘The Realm of Hard Evidence’: Novelty, Persuasion and Collaboration in Botanical Cladistics. Stud. Hist. Phil. Biol. & Biomed. Sci., 32, �4�–��0.

Engels E-M (�00�) Ethik in den Biowissenschaften. Ethisch-Philosophisches Grundlagenstudium 2. Ein Projektbuch. (ed by M Maring), pp. ���-���, Münster, LIT Verlag.

Engler A (�879) Versuch einer Entwicklungsgeschichte der Pflanzenwelt, insbesondere der Florengebiete seit der Tertiärperiode. Vol 1. Die Extratropischen Gebiete der Nördlichen Hemisphäre. Verlag von W. Engelmann, Leipzig.

Engler A (�88�) Versuch einer Entwicklungsgeschichte der Pflanzenwelt, insbesondere der Florengebiete seit der Tertiärperiode. Vol. 2. Die Extratropischen Gebiete der Südlichen Hemisphäre und die Tropischen Gebiete. Verlag von W. Engelmann, Leipzig.

Engler A (1899) Die Entwickelung der Pflanzengeographie in den letzten hundert Jahren und weitere Aufgaben derselben. Wissenschaftliche Beiträge zum Gedächtnis der hundertjährigen Wiederkehr des Antritts von Alexander von Humboldt’s Reise nach Amerika. (ed. by Gesellschaft für Erdkunde zu Berlin), pp. �-�47.

Evenstar L, A Hartley, C Rice, F Stuart, A Mather & G Chong (�00�) Miocene-Pliocene climate change in the Peru-Chile Desert. 6th International Symposium on Andean Geodynamics (ISAG 2005, Barcelona), Extended Abstracts, pp. ��8-��0.

Falcon-Lang HJ, DJ Cantrill & GJ Nichols (�00�) Biodiversity and terrestrial ecology of a mid-Cretaceous, high-latitude floodplain, Alexander Island, Antarctica. Journal of the Geological Society, 158: 709-7�4.

Farjon A, QP Xiang & XQ Zhang (2004) The comparative study on the cuticle micro-morphology of Pilgerodendron uviferum (Cupressaceae) and its relatives. Acta Phytotaxonomica Sinica, 42, 4�7-4��.

Fernández-Armesto F (�997) Truth -a history-. Bantam Press, London.

Ferré F (�997) The Postmodern World. Principles of Conservation Biology (�d edition) (ed. by GK Meffe & CR Carroll), pp. ���-���, Sunderland, Massachusetts, Sinauer Associates .

Fickert T, D Friend, F Grüninger, B Molnia & M Richter (�007). Did debris covered glaciers serve as Pleistocene refugia for plants? A new hypothesis derived from observations of recent plant growth on Glacier surfaces. Arctic, Antarctic, and Alpine Research xx, xx

Foulger GR, JH Natland, DC Presnall & DL Anderson (eds.) (�00�) Plates, plumes, and paradigms. Geological Society of America Special Paper N° �88.

Francis JE & I Poole (�00�) Cretaceous and Tertiary climates of Antarctica: evidence from fossil wood. Palaeogeography, Palaeoclimatology, Palaeoecology, 182, 47–�4.

Frey W & R Lösch (�004) Lehrbuch der Geobotanik. �d edition. Elsevier Spektrum Akad. Verlag.

Friis EM, Doyle JA, Endress PK, & Q Leng (2003) Archaefructus - Angiosperm precursor or specialized early angiosperm? Trends in Plant Science, 8, ��9-�7�

Friis EM, KR Pedersen & PR Crane (�00�) When Earth started blooming: insights from the fossil record. Current Opinion in Plant Biology, 8, �-��.

Friis EM, K Raunsgaard Pedersen & PR Crane (2006) Cretaceous angiosperm flowers: Innovation and evolution in plant reproduction. Palaeogeography, Palaeoclimatology, Palaeoecology, 232, ���-�9�.

�97

Friis I (�998) Frank White and the development of African Chorology. Chorology, taxonomy and ecology of the floras of Africa and Madagascar (ed. by CR Huxley, JM Lock & DF Cutler), pp. ��-��, Royal Botanic Gardens, Kew.

Frohlich MW (�00�) Recommendations and goals for evo-devo research: scenarios, genetic constraint, and developmental homeostasis. Aliso, 22, �7�-�87.

Fuentes F (�9��) Reseña botánica sobre la Isla de Pascua. Bol. Mus. Nac. Chile, 5, ��0-��7.

Fuller DQ & LJ Hickey (2005) Systematics and leaf architecture of the Gunneraceae. Botanical Review, 71, �9�-���.

Gadek PA, DL Alpers, MM Heslewood, & CJ Quinn (2000) Relationships within Cupressaceae sensu lato: A combined morphological and molecular approach. American Journal of Botany, 87, �044-�0�7.

Gajardo R (�994) La vegetación natural de Chile. Clasificación y distribución geográfica. Ed. Universitaria, Santiago. Galtier J & TL Phillips (1996) Structure and evolutionary significance of Palaeozoic ferns. Pteridology in perspective (ed. by JM Camus, M Gibby & RJ Johns) pp. 4�7–4��. Royal Botanic Gardens, Kew.

García Azkonobieta T (2005) Evolución, desarrollo y (auto)organización. Un estudio sobre los principios filosóficos de la evo-devo. Tesis doctoral, Universidad del País Vasco.

Gaston KJ (�00�) The Structure and Dynamics of Geographic Ranges. Oxford University Press.

Gatterer K & W Nezadal (eds.) (�00�) Flora des Regnitzgebietes. Die Farn- und Blütenpflanzen im zentralen Nordbayern. � vols. IHW-Verlag Eching.

Gauckler K (�9�0) Das südlich-kontinentale Element in der Flora von Bayern mit besonderer Berücksichtigung des Fränkischen Stufenlandes. Abhandl. Naturhist. Ges. Nürnberg, 24, �-��0.

Gay C (�8��) La Isla de Juan Fernández. Reprint �974, Museo Nacional de Historia Natural (Santiago) Serie Educativa, 9, �-��.

Gay C (�84�–�8�4) Historia Física y Política de Chile según documentos adquiridos en esta República durante doce años de residencia en ella y publicada bajo los auspicios del Supremo Gobierno. Paris.

Gensel PG & CM Berry (�00�) Early lycophyte evolution. American Fern Journal, 91, 74-98.

Gentry AH (�98�) Neotropical Floristic Diversity: Phytogeographical Connections Between Central and South America, Pleistocene Climatic Fluctuations, or an Accident of the Andean Orogeny? Annals of the Missouri Botanical Garden, 69, ��7-�9�.

Ghosh P, NC Garzione & JM Eiler (�00�) Rapid Uplift of the Altiplano Revealed Through ��C-�8O Bonds in Paleosol Carbonates. Science, 311, ���-���.

Gibson AC & PS Nobel (�98�) The cactus primer. Harvard Univ. Press. Cambridge, Massachusetts.

Gieryn TF (�999) Cultural Boundaries of Science: Credibility on the Line. University of Chicago Press, Chicago.

Gillespie WH, GW Rothwell & SE Scheckler (�98�) The Earliest Seeds. Nature, �9�, 4��-4�4.

Gilmour JSL (1936) Two early papers on classification. Reprint 1989, Plant Systematics and Evolution, 167, 97-�07.

Glasby CJ (�00�) Polychaete distribution patterns revisited: an historical explanation. Marine Ecology, 26, ���-�4�.

Godley EJ (�9�7) Widely distributed species, land bridges and continental drift. Nature, 214, 74-7�.

Goldblatt P & J Manning (�000) Cape Plants: a conspectus of the Cape Flora of South Africa. Strelitzia, 9, �-74�, National Botanical Institute of South Africa.

Good R (�947) The Geography of the Flowering Plants. 4th edition �974, Longman, London.

�98

Goodwin B (�994) How the Leopard Changed Its Spots: The Evolution of Complexity. Charles Scribner’s Sons, New York.

Grande L (�994) Repeating Patterns in Nature, Predictability, and “Impact” in Science. Interpreting the Hierarchy of Nature: From Systematic Patterns to Evolutionary Process Theories (ed. by L Grande & O Rieppel), pp. ��-84, Academic Press, Inc.

Grant V (1998) Primary classification and phylogeny of the Polemoniaceae, with comments on molecular cladistics. American Journal of Botany, 85, 74�-7��.

Grant V (�00�) Incongruence between cladistic and taxonomic systems. American Journal of Botany, 90, ����-��70.

Gregory-Wodzicki KM (�000) Uplift history of the central and northern Andes: a review. Geological Society of America Bulletin, 112, �09�–��0�.

Grehan JR (�00�) Panbiogeography from tracks to ocean basins: evolving perspectives. Journal of Biogeography, 28, 4��-4�9.

Grehan JR (2007) A Brief Look at Pacific Biogeography: The Trans-Oceanic Travels of Microseris (Angiosperms: Asteraceae). Biogeography in a Changing World (ed. by MC Ebach & RS Tangney), pp. 8�-94. Systematics Association Special Volumes, CRC Press.

Griffith MP (2003) Relationships among the genera of Opuntioideae (Cactaceae): preliminary insights. Cactaceae Systematics Initiatives, 15, ��–��.

Griffith MP (2004a) What did the first cactus look like? An attempt to reconcile the morphological and molecular evidence. Taxon, 53, 49�–499.

Griffith MP (2004b) Early cactus evolution: the postmodern view. Haseltonia, 10, �-��.

Griffiths G (1974) On the Foundations of Biological Systematics. Acta Biotheoretica, 3–4, 8�–���.

Grisebach A (�87�) Die Vegetation der Erde nach ihrer klimatischen Anordnung: ein Abriss der vergleichenden Geographie der Pflanzen. (� ed.). � vols., Engelmann Verlag, Leipzig. (�d revised ed. �884)

Groom MJ, GK Meffe & CR Carrol (eds.) (�00�) Principles of Conservation Biology, �d edition, Sinauer, Massachusetts.

Grunow AM, DV Kent & IWD Dalziel (�99�) New paleomagnetic data from Thurston Island: implications for the tectonics of West Antarctica and Weddell Sea opening. Journal of Geophysical Research, B, Solid Earth and Planets, 96, �79��–�79�4.

Guggisberg A, G Mansion, S Kelso & E Conti (�00�) Evolution of biogeographic patterns, ploidy levels, and breeding systems in a diploid-polyploid species complex of Primula. New Phytologist, 171, ��7-���.

Gunckel H (�9��) Algunas plantas vasculares de la Isla de San Ambrosio. Revista Universitaria (Santiago), 36, �9�-��4.

Haeckel E (�8��) Generelle Morphologie der Organismen. Vol. 2. Allgemeine Entwickelungsgeschichte der Organismen. Reprint �988, Walter de Gruyter, Berlin.

Haeupler H (2005) Ein Beitrag zur Geschichte der floristischen Kartierung der Farn- und Blütenpflanzen in Mitteleuropa. Hoppea, Denkschr. Regensb. Bot Ges., 66, ���-��9.

Haeupler H & P Schönfelder (eds.) (�988) Atlas der Farn- und Blütenpflanzen der Bundesrepublik Deutschland. Ulmer Stuttgart.

Hamilton WB (�00�) The Closed Upper-Mantle Circulation of Plate Tectonics. Plate Boundary Zones (ed. by S Stein & JT Freymueller). American Geophysical Union Geodynamics Series, 30, ��9-4�0.

Hannig E & H Winkler H (eds.) (�9��-�940) Die Pflanzenareale: Sammlung kartographischer Darstellungen von Verbreitungsbezirken der lebenden und fossilen Pflanzen-Familien,-Gattungen und -Arten. Jena, Gustav Fischer Verlag.

�99

Harold AS & RD Mooi (1994) Areas of endemism – definition and recognition criteria. Systematic Biology, 43, ���-���.

Hartley AJ & G Chong (2002) A late Pliocene age for the Atacama Desert: Implications for the desertification of western South America. Geology, 30, 4�-4�.

Hartley AJ (�00�) Andean uplift and climate change. Journal of the Geological Society, 160, 7-�0.

Hartley AJ, G Chong, J Houston & AE Mather (�00�) ��0 million years of climatic stability: evidence from the Atacama Desert, northern Chile. Journal of the Geological Society, 162, 4��-4�4.

Haywood AM, JE Francis & BW Sellwood (�00�) Global middle Pliocene biome reconstruction: a data/model synthesis. Geochemistry Geophysics Geosystems, 3, �-�8.

Heads MJ (�990) Mesozoic tectonics and the deconstruction of biogeography: a new model of Australasian biology. Journal of Biogeography, 17, ���-���.

Heads MJ (�994) Biogeographic studies in New Zealand Scrophulariaceae: tribes Rhinantheae, Calceolarieae and Gratioleae. Candollea, 49, ��-80.

Heads MJ (1996) Biogeography, taxonomy and evolution in the Pacific genus Coprosma (Rubiaceae). Candollea, 51, �8�–40�.

Heads M (1999) Vicariance biogeography and terrane tectonics in the South Pacific: analysis of the genus Abrotanella (Compositae). Biological Journal of the Linnean Society, 67, �9�-4��.

Heads M (�00�) Birds of paradise, vicariance biogeography and terrane tectonics in New Guinea. Journal of Biogeography, 29, ���-�8�.

Heads M (�004) What is a node? Journal of Biogeography, 31, �88�-�89�.

Heads M (�00�a) The history and philosophy of panbiogeography. Regionalización Biogeográfica en Iberoamérica y Tópicos Afines (ed. by J Llorente & JJ Morrone), pp. �7-���, Universidad Nacional Autónoma de México.

Heads M (�00�b) Dating nodes on molecular phylogenies: a critique of molecular biogeography. Cladistics, 21, ��–78.

Heads M (�00�) Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species massings. Journal of Biogeography, 33, �0��–�07�.

Heads M & RC Craw (1984) Bibliography of the scientific work of Leon Croizat, 1932-1982. Tuatara, 27, �7-7�. [http://www.sciencebuff.org/bibliography_of_leon_croizat.php]

Heckman DS, DM Geiser, BR Eidell, RL Stauffer, NL Kardos & SB Hedges (�00�) Molecular evidence for the early colonization of land by fungi and plants. Science, 293, ���9–����.

Hengeveld R (�990) Dynamic Biogeography. Cambridge Studies in Ecology. Cambridge University Press.

Hennig W (�9�0) Grundzüge einer Theorie der phylogenetischen Systematik. Deutscher Zentralverlag, Berlin. Reprint �980, Otto Koeltz Science Publishers, Koenigstein, Germany.

Hennig W (�9��) Phylogenetic Systematics. University of Illinois Press, Chicago.

Henríquez JM, E Pisano & C Marticorena (1995) Catálogo de la flora vascular de Magallanes (XII Región), Chile. Anales Instituto de la Patagonia Ser. Cs. Naturales, (Chile), 23, �-�0.

Herbst R & A Troncoso (�000) Triassic cycadophyta from the La Ternera and El Puquén formations (Chile). Ameghiniana, 37, �8�-�9�.

Herbst R, A Troncoso & J Muñoz (2005) Las Tafofloras Triásicas de la Región de Los Lagos, Xma Región, Chile. Ameghiniana, 42, �77-�94.

Herendeen PS & PR Crane PR (�99�) The fossil history of the monocotyledons. Monocotyledons: systematics and evolution (ed. by PJ Rudall, PJ Cribb, DF Cutler & CJ Humphries), pp. �–��, Royal Botanic Gardens, Kew.

�00

Hershkovitz MA (2006) Ribosomal and chloroplast DNA evidence for diversification of western American Portulacaceae in the Andean region. Gayana Botánica, ��, ��-74.

Hershkovitz MA & EA Zimmer (�997) On the evolutionary origins of the cacti. Taxon, 46, ��7–���.

Hershkovitz MA, MTK Arroyo, Ch Bell & LF Hinojosa (�00�) Phylogeny of Chaetanthera (Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages. Molecular Phylogenetics and Evolution, 41, �94-�0�.

Hess H (�9��) History of Ocean Basins. Petrologic Studies (ed. by AEJ Engel, HL James & BF Leonard), pp. �99-��0, Washington DC.

Hettlage R (�98�) Variationen des Darwinismus in der Soziologie. Schriftenreihe der Universität Regensburg, Band 7 - Evolutionstheorie und ihre Evolution, Regensburg.

Hettner A (�9�7) Die Geographie, ihre Geschichte, ihr Wesen und ihre Methoden. Ferdinand Hirt, Breslau.

Heusser CJ (�00�) On glaciation of the southern Andes with special reference to the Peninsula de Taitao and adjacent Andean cordillera (~4�°�0‘S). Journal of South American Earth Sciences, 15, �77-�89.

Heusser CJ (�00�) Ice Age Southern Andes - A Chronicle of Paleoecological Events. Developments in Quaternary Science, �. Elsevier.

Heusser CJ, LE Heusser & TV Lowell (�999) Paleoecology of the southern Chilean Lake District-Isla Grande de Chiloe during middle-late Llanquihue glaciation and deglaciation. Geografiska Annaler Series A-Physical Geography, 81A, ���-�84.

Hey J, RS Waples, ML Arnold, RK Butlin & RG Harrison (�00�) Understanding and confronting species uncertainty in biology and conservation. Trends in Ecology and Evolution, 18, �97-�0�.

Hilgenberg OC (�9��) Vom wachsenden Erdball. Published by the author, Berlin.

Hill K (�00�) Diversity and evolution of gymnosperms. Plant Diversity and Evolution: Genotypic and Phenotypic Variation in Higher Plants (ed. by RJ Henry), pp. ��-44. CABI Publishing, Cromwell Press, UK.

Hinojosa LF (2005) Cambios climáticos y vegetacionales inferidos a partir de paleofloras Cenozoicas del sur de Sudamérica. Revista Geológica de Chile, 32, 9�-���.

Hinojosa LF & C Villagrán (�997) Historia de los bosques del sur de Sudamérica, I: antecedentes paleobotánicos, geológicos y climáticos del Terciario del cono sur de América. Revista Chilena de Historia Natural, 70, ���-��9.

Hinojosa LF & C Villagrán (2005) Did South American Mixed Paleofloras evolve under thermal equability or in the absence of an effective Andean barrier during the Cenozoic? Palaeogeography, Palaeoclimatology, Palaeoecology, 217, �-��.

Hinojosa LF, JJ Armesto & C Villagrán (�00�). Are Chilean coastal forests pre-Pleistocene relicts? Evidence from foliar physiognomy, paleoclimate, and paleobiogeography. Journal of Biogeography, 33, ���-�4�.

Hodge J (�00�) Against “Revolution” and “Evolution”. Journal of the History of Biology, 38, �0�-���.

Hoffmann AJ & S Teillier (1991) La flora de la Isla San Félix (Archipiélago de las Desventuradas, Chile). Gayana Botánica, 48, 89-99.

Hofreiter A (�00�) Leontochir: a synonym of Bomarea (Alstroemeriaceae)? Harvard Papers in Botany, 11, ��–�0.

Holt-Jensen A (�999) Geography History & Concepts. A student’s Guide. �d edition. Sage Publications, London.

Hooghiemstra H & T van der Hammen (1998) Neogene and Quaternary development of the neotropical rain forest: the forest refugia hypothesis, and a literature overview. Earth-Science Reviews, 44, �47–�8�.

Hooker JD (�844-�8�0) The botany of the Antarctic voyage of H. M. Discovery ships Erebus and Terror, in the years 1839–1843. I Flora Antarctica, II Flora Nova-Zelandiae, III Flora Tasmaniae, L. Reeve, London.

�0�

Hoot SB & AW Douglas (�998) Phylogeny of the Proteaceae based on atpB and atpB-rbcL spacer region sequences. Australian Systematic Botany, 11, �0�-��0.

Houston J & AJ Hartley (�00�) The central andean west-slope rainshadow and its potential contribution to the origin of hyper-aridity in the Atacama desert. International Journal of Climatology, 23, �4��-�4�4.

Howard DJ (1998) Unanswered Questions and Future Directions in the Study of Speciation. Endless Forms: Species and Speciation (ed. by DJ Howard & SH Berlocher), pp. 4�9-448, Oxford University Press.

Howarth JM (1995) Ecology: modern hero or post-modern villain? From scientific trees to phenomenological wood. Biodiversity and Conservation, 4, 78�-797.

Hu HH (1950) A polyphyletic system of classification of angiosperms. Science Record (Peking), 3, ���-��0.

Hubbell (SP) (�00�) The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton.

Hueck K (�978) Los bosques de Sudamérica. GTZ, Eschborn.

Hughes C & R Eastwood (2006) Island radiation on a continental scale: exceptional rates of plant diversification after uplift of the Andes. Proceedings of the National Academy of Sciences of the USA, 103, �0��4-�0��9.

Hull DL (�988) Progress in ideas of progress. Evolutionary progress (ed. by M. Nitecki), pp. �7-48, University of Chicago Press.

Hull DL (�00�) The Role of Theories in Biological Systematics. Stud. Hist. Phil. Biol. & Biomed. Sci., 32, ���–��8.

Hull DL (�00�) Deconstructing Darwin: Evolutionary Theory in Context. Journal of the History of Biology, 38, ��7–���.

Hultén E (�9�0) Atlas of the Distribution of Vascular Plants in NW-Europe. Stockholm.

Hultén E (�9�4) The Circumpolar Plants I Vascular Cryptogams, Conifers, Monocotyledons. Stockholm.

Hultén E (�97�) The Circumpolar Plants II Dicotyledons. Stockholm.

Humboldt A von (�80�) Essai sur la géographie des plantes, accompagné d’un tableau physique des régions équinoxiales. Paris, Fr. Schoell.

Humphries CJ & LR Parenti (�999) Cladistic biogeography (�d edition). Oxford monographs on biogeography No. �. Clarendon Press, Oxford (�st edition �98�).

Huntington E (�9��). The Character of Races. As influenced by physical environment, natural selection and historical development. Charles Scribner’s Sons, New York, London.

Hunziker JH, RA Palacios, AG de Valesi & L Poggio (�97�) Species disjunctions in Larrea: evidence from morphology, cytogenetics, phenolic compounds, and seed albumins. Annals of the Missouri Botanical Garden, 59, ��4–���.

Hurr KA, PJ Lockhart, PB Heenan & D Penny (�999) Evidence for the recent dispersal of Sophora (Leguminosae) around the Southern Oceans: molecular data. Journal of Biogeography, 26, ���-�77.

Huxley CR, JM Lock & DF Cutler (eds) (�998) Chorology, taxonomy and ecology of the floras of Africa and Madagascar. Royal Botanic Gardens, Kew.

Illies H (1967) Randpazifische Tektonik und Vulkanismus im südlichen Chile. Geologische Rundschau, 57, 8�--�0�.

Irmscher E (1922) Pflanzenverbreitung und Entwicklung der Kontinente. I Teil. Studien zur genetische Pflanzengeographie. Mitteilungen aus dem Institut für allgemeine Botanik in Hamburg, 5, �7-���.

Irmscher E (1929) Pflanzenverbreitung und Entwicklung der Kontinente. II Teil. Weitere Beiträge zur genetische Pflanzengeographie unter besonderer Berücksichtigung der Laubmoose. Mitteilungen aus dem Institut für allgemeine Botanik in Hamburg, 8, ��9-�74.

�0�

Janssen T & K Bremer (�004) The age of major monocot groups inferred from 800+rbcL sequences. Botanical Journal of the Linnean Society, 146, �8�-�98.

Johnson LAS (�998) Proteaceae-Where Are We? Australian Systematic Botany, 11, ���-���.

Johnston IM (1929a) The coastal flora of the departments of Chañaral and Taltal. Contr. Gray Herb., 85, �-��8.

Johnston IM (1929b) The flora of the Nitrate Coast. Contr. Gray Herb., 85, ��8-���.

Johnston IM (1935) The flora of San Félix island. J. Arnold. Arbor., 16, 440-447.

Johnston IM (1940) The floristic significance of shrubs common to North and South American deserts. Journal of the Arnold Arboretum, 21, ���–���.

Johow F (�948). Flora de Zapallar. Revista Chilena de Historia Natural vol. IL: �-��4.

Johow F (�89�). Estudios sobre la flora de las Islas de Juan Fernández. Santiago, Chile.

Jokat W, T Boebel, M Könog & U Meyer (�00�) Timing and geometry of early Gondwana breakup. Journal of Geophysical Research, �08, N° B9, �4�8.

Jones EJW (�987) Fracture zones in the equatorial Atlantic and the breakup of western Pangea. Geology, 15, ���–���.

Jørgensen PM & S León-Yáñez (eds.) (�999) Catalogue of the vascular plants of Ecuador. Monographs in Systematic Botany, Missouri Botanical Garden, 75, i–viii, �–��8�.

Kamp PJJ (1980) Pacifica and New Zealand: proposed eastern elements in Gondwanaland’s history. Nature, 288, ��9– ��4.

Katinas L & JV Crisci (�000) Cladistic and Biogeographic Analyses of the Genera Moscharia and Polyachyrus (Asteraceae, Mutisieae). Systematic Botany, 25, ��-4�.

Katinas L, JJ Morrone & JV Crisci (�999) Track analysis reveals the composite nature of the Andean biota. Australian Journal of Botany, 47, ���-��0.

Katinas L, JV Crisci, W Wagner & PC Hoch (2004) Geographical diversification of tribes Epilobieae, Gongylocarpeae, and Onagreae (Onagraceae) in North America, based on parsimony analysis of endemicity and track compatibility analysis. Annals of the Missouri Botanical Garden, 91, ��9-�8�.

Kau A (�00�) Deconstruction and Science: How Post-Structuralist Literary Theory Appliesto Scientific Understanding. [http://www.physics.orst.edu/~stetza/COURSES/ph407h/Deconstruction.pdf]

Kennedy BA (�00�) Inventing the Earth: Ideas on Landscape Development Since 1740. Blackwell.

Kenrick P & PR Crane (�997) The origin and early diversification of land plants: a cladistic study. Smithsonian Institution Press, Washington DC.

Kim YD, SH Kim & LR Landrum (�004) Taxonomic and phytogeographic implications from ITS phylogeny in Berberis (Berberidaceae). Journal of Plant Research, 117, �7�-�8�.

Klicka J & RM Zink (�997) The Importance of Recent Ice Ages in Speciation: A Failed Paradigm. Science, 277, ����-���9.

Knapp M, K Stöckler, D Havell, F Delsuc, F Sebastiani & PJ Lockhart (�00�) Relaxed molecular clock provides evidence for long-distance dispersal of Nothofagus (southern beech). PLoS Biology, 3, �8-4�.

Knoll AH (�984) Patterns of extinction in the fossil record of vascular plants. Extinctions (ed. by MH Nitecki) pp. ��–�8. University of Chicago Press, Chicago.

Kohler RE (�00�) Landscapes and Labscapes: Exploring the Lab–Field Border in Biology. University of Chicago Press, Chicago.

�0�

Köppen W (�884) Die Wärmezonen der Erde, nach der Dauer der heissen, gemässigten and kaltern Zeit and nach der Wirkung der Wärme auf die organische Welt betrachtet. Meteorologische Zeitschrift, 1, ���-���.

Köppen W (1930) Die Klimagebiete nach Köppens Klassifikation. Klimakunde von Südamerika (ed. by K Knoch), vol. �, part G, pp. �4�-��9. Handbuch der Klimatologie (ed. by W Köppen & R Geiger), Berlin.

Kron KA (�99�) Phylogenetic relationships of Empetraceae, Epacridaceae, Ericaceae, Monotropaceae, and Pyrolaceae: Evidence from nuclear ribosomal �8s sequence data. Annals of Botany, 77, �9�-�0�.

Kron KA & JL Luteyn (�00�) Origins and biogeographic patterns in Ericaceae: New insights from recent phylogenetic analyses. Biol. Skr., 55, 479-�00.

Kropotkin PA (�90�) Mutual Aid: a Factor of Evolution. W Heinemann, London.

Kuschel G (�9��) Zur Naturgeschichte der Insel San Ambrosio (Islas Desventuradas, Chile). �. Reisebericht, geographische Verhältnisse und Pfalzenverbreitung. Ark. Bot., 4, 4��-4�9.

Kvasov DD & MY Verbitski (�98�) Causes of Antarctic glaciation in the Cenozoic. Quaternary Research, 15, �–�7.

Ladiges PY, J Kellermann, G Nelson, CJ Humphries & F Udovicic (�00�) Historical biogeography of Australian Rhamnaceae, tribe Pomaderreae. Journal of Biogeography, 32, �909–�9�9.

Lammers TG, TF Stuessy& M Silva O (�98�) Systematic relationships of the Lactoridaceae, an endemic family of the Juan Fernandez Islands, Chile. Plant Systematics and Evolution, 152, �4�-���.

Landrum L (�98�) The phylogeny and geography of Myrceugenia (Myrtaceae). Brittonia, 33, �0�-��9.

Lawrence GHM (�9��) Taxonomy of Vascular Plants. Macmillan, New York.

Lawver LA & LM Gahagan (�998) Opening of Drake Passage and its impact on Cenozoic ocean circulation. Tectonic boundary conditions for climate reconstructions (ed. by TJ Crowley & KC Burke) Oxford Monographs on Geology and Geophysics, 39, ���–���.

Leppe M & P Moisan (�00�) New records of Upper Triassic Cycadales and Cycadeoidales of Biobio river, Chile. Revista Chilena de Historia Natural, 76, 47�-484.

Leuenberger BE (�98�) Pereskia (Cactaceae). Memoirs of the New York Botanical Garden, 41, �–�4�.

Lewis LA & RM McCourt (�004) Green algae and the origin of land plants. American Journal of Botany, 91, ����–����.

Lidgard S & PR Crane (1990) Angiosperm diversification and Cretaceous floristic trends: a comparison of palynofloras and leaf macrofloras. Paleobiology, 16, 77–9�.

Lindemann B (�9�7) Kettengebirge, kontinentale Zerspaltung und Erdexpansion. Jena.

Linder CR & LH Rieseberg (�004) Reconstructing patterns of reticulate evolution in plants. American Journal of Botany, 91, �700–�708.

Linder HP (�00�) On areas of endemism, with an example from the African Restionaceae. Systematic Biology, 50, 89�–9��.

Linder HP & NP Barker (�000) Biogeography of the Danthonieae. Grasses: Systematics and Evolution (ed. by SWL Jacobs & J Everett), pp. ���–��8. CSIRO Publishing, Collingwood, Victoria, Australia.

Linder HP & MD Crisp (�99�) Nothofagus and Pacific biogeography. Cladistics, 11, �–��.

Linder HP & PJ Rudall (�00�) Evolutionary history of Poales. Annual Review of Ecology and Systematics, 36, �07-��4.

Linder HP, PK Eldenas & BG Briggs (�00�) Contrasting patterns of radiation in African and Australian Restionaceae. Evolution, 57, ��88–�70�.

�04

Livingstone DN (�99�) The Geographical Tradition: Episodes in the History of a Contested Enterprise. Blackwell, Oxford & Cambridge.

Livingstone DN (�994) Climate’s Moral Economy: Science, Race and Place in Post-Darwinian British and American Geography. Geography and Empire (ed. by A Godlewska & N Smith), pp. ���-��4, Blackwell.

Livingstone DN (�00�) Science, space and hermeneutics. Hettner-Lecture �00�, Dept. of Geography, University of Heidelberg.

Lomolino MV & LR Heaney (�004) Reticulations and Reintegration of Modern Biogeography. Frontiers in Biogeography: New Directions in the Geography of Nature. (ed. by MV Lomolino & LR Heaney), pp. �-�, Sinauer, Massachusetts.

Lomolino MV, BR Riddle & JH Brown (�00�) Biogeography. �d edition. Sinauer Associates, Sunderland, Massachusetts.

López RP (�00�) Phytogeographical relations of the Andean dry valleys of Bolivia. Journal of Biogeography, 30, ���9–���8.

Lu A-M & Y-C Tang (�00�) Consideration on some viewpoints in researches of the origin of angiosperms. Acta Phytotaxonomica Sinica, 43, 4�0-4�0.

Luebert F & R Gajardo (�000) Vegetación de los Andes áridos del norte de Chile. Lazaroa 21, ���-��0.

Luebert F & R Gajardo (�00�) Vegetación alto-andina de Parinacota (norte de Chile) y una sinopsis de la vegetación de la Puna meridional. Phytocoenologia 35, 79-��8.

Luebert F & P Pliscoff (�00�) Sinopsis bioclimática y vegetacional de Chile. Editorial Universitaria, Santiago.

Luebert F, M Muñoz-Schick & A Moreira-Muñoz (�00�) Vegetación y Flora de La Campana. Parque Nacional La Campana: Origen de una Reserva de la Biosfera en Chile Central. (ed. by S Elórtegui & A Moreira-Muñoz), pp. ��-�9, Taller La Era, Santiago.

Mabberley DJ (�997) The Plant-Book (�nd ed.). Cambridge University Press, Cambridge

MacArthur RH & EO Wilson (�9�7) The Theory of Island Biogeography. Princeton University Press, Princeton.

Macphail MK, AD Partridge & EM Truswell (�999) Fossil pollen records of the problematical primitive angiosperm family Lactoridaceae in Australia. Plant Systematics and Evolution, 214, �99-��0.

Magallón S & MJ Sanderson (�00�) Relationships among seed plants inferred from highly conserved genes: Sorting conflicting phylogenetic signals among ancient lineages. American Journal of Botany, 89,�99�–�00�.

Maldonado A, JL Betancourt, C Latorre & C Villagrán (�00�) Pollen analyses from a �0 000-yr rodent midden series in the southern Atacama Desert (�� degrees �0’S). Journal of Quaternary Science, 20, 49�-�07.

Mann G (�980) Ernst Haeckel und der Darwinismus: Popularisierung, Propaganda, und Ideologisierung. Medizin-historisches Journal, 15, ��9-�8�.

Marquet PA, M Fernández, SA Navarrete & C Valdovinos (�004) Diversity Emerging: Toward a Deconstruction of Biodiversity Patterns. Frontiers in Biogeography: New Directions in the Geography of Nature. (ed. by MV Lomolino & LR Heaney), pp. �9�-�09, Sinauer, Massachusetts.

Marticorena C (1990) Contribución a la estadística de la flora vascular de Chile. Gayana Botánica, 47, 8�-���.

Marticorena C (�99�) Bibliografía Botánica Taxonómica de Chile. Monographs in Systematic Botany vol. 4�, Missouri Botanical Garden.

Marticorena C (�99�) Historia de la exploración botánica en Chile. Flora de Chile, vol. 1 (ed. by C Marticorena & R Rodríguez), pp. �-��, Universidad de Concepción.

�0�

Marticorena C (�997) La presencia de Maireana brevifolia (R.Br.) Paul G. Wilson (Chenopodiaceae), en Chile continental. Gayana Botánica, 54, �9�-�94.

Marticorena C & R Rodríguez (eds.) (�99�-�00�) Flora de Chile. 2 vol. Universidad de Concepción, Concepción.

Marticorena C, O Matthei, R Rodríguez, MTK Arroyo, M Muñoz-Schick, FA Squeo & G Arancio (�998a) Catálogo de la Flora Vascular de la Segunda Región (Región de Antofagasta), Chile. Gayana Botánica, 55, ��-8�.

Marticorena C, TF Stuessy & CM Baeza (1998b) Catalogue of the vascular flora of the Robinson Crusoe or Juan Fernández Islands, Chile. Gayana Botánica, 55, �87-���.

Marticorena C, FA Squeo, G Arancio & M Muñoz-Schick (2001) Catálogo de la flora vascular de la IV Región de Coquimbo. Libro Rojo de la Flora Nativa y de los Sitios Prioritarios para su Conservación: Región de Coquimbo (ed. by FA Squeo, G Arancio & JR Gutiérrez), pp. �0�-�4�, Ediciones Universidad de La Serena, Chile.

Martin W, A Gieri & H Saedler (�989) Molecular evidence for pre-Cretaceous angiosperm origins. Nature, 339, 4�-48.

Martin W, D Lydiate, H Brinkmann, G Forkmann, H Saedler & R Cerff (�99�) Molecular phylogenies in angiosperm evolution. Mol. Biol. Evol., 10, �40 - ���.

Martínez-Pardo R (1990) Major Neogene events of Southeastern Pacific: the Chilean and Peruvian record. Palaeogeography, Palaeoclimatology, Palaeoecology, 77, ���-�78.

Mathias ME & L Constance (�9��) The genus Oreomyrrhis (Umbelliferae), a problem in South Pacific distribution. University of California Publications in Botany, �7, �47-4��.

Matthew WD (�9�9) Climate and Evolution. Arno Press, New York. Reprint �974. Originally published (�9��) as essay in the Annals of the New York Academy of Sciences.

Mattick F (�9�4) Übersicht über die Florenreiche und Florengebiete der Erde. A. Engler’s Syllabus der Pflanzenfamilien. 12 ed. vol. 2. Angiospermen. (ed. by H. Melchior), pp. ���-��9, Gerbrüder Borntraeger, Berlin.

Mauseth JD (�990) Continental drift, climate and the evolution of cacti. Cactus and Succulent Journal, ��, �0�-�08.

Maxwell R (�998) The Rhipsalis Riddle - or the day the cacti came down from the trees. [http://www.rhipsalis.com/maxwell.htm]

Mayr E (�9��) Animal Species and Evolution. Harvard University Press.

McCarthy D (2003) The trans-Pacific zipper effect: disjunct sister taxa and matching geological outlines that link the Pacific margins. Journal of Biogeography, 30, ��4�–����.

McCarthy D (2005a) Biogeographical and geological evidence for a smaller, completely-enclosed Pacific Basin in the Late Cretaceous. Journal of Biogeography, 32, ����–��77.

McCarthy D (2005b) Biogeography and Scientific Revolutions. The Systematist, 25, �-��.

McCarthy D (2007) Are Plate Tectonic Explanations for Trans-Pacific Disjunctions Plausible? Empirical Tests of Radical Dispersalist Theories. Biogeography in a Changing World (ed. by MC Ebach & RS Tangney), pp. �7�-�9�. Systematics Association Special Volumes, CRC Press.

McCourt RM, CF Delwiche & KG Karol (�004) Charophyte algae and land plant origins. Trends in Ecology and Evolution, 19, ���-���.

McGlone MS (�00�) Goodbye Gondwana. Journal of Biogeography, 32, 7�9–740.

McLaughlin SP (1992) Are floristic areas hierarchically arrange? Journal of Biogeography, 19, ��–��.

McLaughlin SP (1994) Floristic plant geography: the classification of floristic areas and floristic elements. Progress in Physical Geography, 18, �8�-�08.

McLoughlin S (2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany, 49, �7�–�00.

�0�

Meffe GK & CR Carroll (eds.) (�997) Principles of Conservation Biology, 2d edition, Sinauer, Massachusets (�st edition �994).

Melville R (1981) Vicarious plant distributions and paleogeography of the Pacific Region. Vicariance Biogeography: a critique (ed. by G Nelson & DE Rosen), pp. ��9-�74, Columbia U. Press, New York.

Mennema J (1985) The first plant distribution map. Taxon, 34, ���-��7.

Merleau-Ponty M (�9��) Phenomenology of perception, 1st ed. (C. Smith, trans.) London, Routledge.

Merriam CH (�89�) The geographical distribution of life in North America with special reference to the Mammalia. Proceedings of the Biological Society of Washington, 7, �–�4.

Meudt HM & BB Simpson (�00�) The biogeography of the austral, subalpine genus Ourisia (Plantaginaceae) based on molecular phylogenetic evidence South American origin and dispersal to New Zealand and Tasmania. Biological Journal of the Linnean Society, 87, 479–���.

Meusel H (�94�) Vergleichende Arealkunde. Einführung in die Lehre von der Verbreitung der Gewächse mit besonderer Berücksichtigung der mitteleuropäischen Flora. � vols. Verlag Borntraeger, Berlin-Zehlendorf.

Meusel H & E Jäger (eds.) (�99�) Vergleichende Chorologie der Zentraleuropäischen Flora, vol. 3, Jena.

Meusel H, E Jäger & E Weinert (eds.) (�9��) Vergleichende Chorologie der Zentraleuropäischen Flora, vol. 1, Jena.

Meusel H, E Jäger, S Rauschert & E Weinert (eds.) (�978) Vergleichende Chorologie der Zentraleuropäischen Flora, vol. 2, Jena.

Michaux B (2001) Dispersal versus vicariance, artifice rather than contest. Faunal and floral migrations and evolution in SE Asia-Australasia (ed. by I Metcalfe, JMB Smith, M Morwood & I Davidson), pp. ���–��8. A.A. Balkema Publ.

Michaux B & RAB Leschen (�00�) East meets west: biogeology of the Campbell Plateau. Biological Journal of the Linnean Society, 86, 9�-���.

Midgley M (�989) Wisdom, information and wonder. Routledge, New York.

Midgley M (�00�) Science and poetry. Routledge, New York.

Mihoc MAK, JJ Morrone, M Negritto & LA Cavieres (�00�) Evolución de la serie Microphyllae (Adesmia, Fabaceae) en la Cordillera de los Andes: una perspectiva biogeográfica. Revista Chilena de Historia Natural, 79, �89-404.

Milbradt J (�97�) Nordische Einstrahlungen in der Flora und Vegetation von Nordbayern, dargestellt an ausgewählten Beispielen. Hoppea, Denkschr. Regensb. Bot. Ges., 35, ���-��0.

Miller H (1970) Das Problem des hypothetischen „Pazifischen Kontinentes“ gesehen von der chilenischen Pazifikküste. International Journal of Earth Sciences, 59, 9�7-9�8.

Miller JM (�00�) The Origin of Angiosperms. Online, updated 08.0�.07. [http://www.gigantopteroid.org/html/angiosperm.htm]

Milne RI (�00�) Northern Hemisphere Plant Disjunctions: A Window on Tertiary Land Bridges and Climate Change? Annals of Botany, 98, 4��-47�.

Milton R (�00�) Alternative Science. Online [http://www.alternativescience.com/] Updated Dec. �00�.

Mitchell AD & SJ Wagstaff (�000) Phylogeny and biogeography of the Chilean Pseudopanax laetevirens. New Zealand Journal of Botany, 38, 409-4�4.

Molina JI (�77�) Compendio della storia geografica, naturale, e civile del Regno del Chile. Bologna.

Molina JI (�78�) Saggio sulla storia naturale del Chile. Bologna.

Molina JI (�8�0) Saggio sulla storia naturale del Chile. (�d revised edition), Bologna.

�07

Moline PM & HP Linder (�00�) Input data, analytical methods and biogeography of Elegia (Restionaceae). Journal of Biogeography, 33, 47-��.

Moore DM (�98�) Flora of Tierra del Fuego. Missouri Botanical Garden.

Moore J (�99�) Deconstructing Darwinism: The Politics of Evolution in the �8�0s. Journal of the History of Biology, 24, ���–408.

Moore MJ, A Tye & RK Jansen (�00�) Patterns of long-distance dispersal in Tiquilia subg. Tiquilia (Boraginaceae): implications for the origins of amphitropical disjuncts and Galapagos Islands endemics. American Journal of Botany, 93, ����-��77.

Morawetz W & M Röser (eds.) (�998) Eduard Friedrich Poeppig 1798-1868. Gelehrter und Naturforscher in Südamerika anlässlich seines 200. Geburtstages. Universität Leipzig.

Morley RJ (�00�) Why are there so many primitive angiosperms in the rain forests of Asia-Australasia? Faunal andfloral migrations and evolution in SE Asia-Australasia (ed. by I Metcalfe, JMB Smith, M Morwood & I Davidson), pp. �8�–�99. A.A. Balkema Publishers, Tokyo.

Morley RJ (�00�) Interplate dispersal paths for megathermal angiosperms. Persp. Plant Ecol. Evol. Syst., 6, �–�0.

Morrell PL, JM Porter & EA Friar (�000) Intercontinental dispersal: the origin of the widespread South American plant species Gilia laciniata (Polemoniaceae) from a rare California and Oregon coastal endemic. Plant Systematics and Evolution, 224,��-��.

Morrone JJ (1994) On the identification of areas of endemism. Systematic Biology, 43, 4�8-44�.

Morrone JJ (�00�) Biogeografía de América Latina y el Caribe. M&T–Manuales & TesisSEA, vol. �. Zaragoza, �48 pp.

Morrone JJ (�00�) Biogeographical regions under track and cladistic scrutiny. Journal of Biogeography, 29, �49-���.

Morrone JJ (2003) ¿Quién le teme al Darwinismo? Ciencia, 54, 78-88. http://www.sciencebuff.org/morrone_articles.php

Morrone JJ (�00�) Cladistic biogeography: identity and place. Journal of Biogeography, 32, ��8�–��84.

Morrone JJ (�00�) Biogeographic Areas and Transition Zones of Latin America and the Caribbean Islands based on Panbiogeographic and Cladistic Analyses of the Entomofauna. Annu. Rev. Entomol., 51, 4�7–494.

Morton AG (�98�) History of Botanical Science. Academic Press, London.

Müller GH (�99�) Friedrich Ratzel (1844-1904): Naturwissenschaftler, Geograph, Gelehrter: Neue Studien zu Leben und Werk und sein Konzept der „Allgemeinen Biogeographie“. Verlag für Geschichte der Naturwissenschaften und der Technik, Stuttgart.

Müller RD, WR Roest, J-Y Royer, LM Gahagan & JG Sclater (1997) Digital isochrones of the world’s ocean floor. J. Geophys. Res., 102, ����-���4.

Muellner AN, K Tremetsberger, TF Stuessy & CM Baeza (�00�) Pleistocene refugia and recolonization routes in the southern Andes: insights from Hypochaeris palustris (Asteraceae, Lactuceae). Molecular Ecology, 14, �0�–���.

Mummenhoff K, H Bruggemann & JL Bowman (�00�) Chloroplast DNA phylogeny and biogeography of Lepidium (Brassicaceae). American Journal of Botany, 88, �0��–�0��.

Muñoz J, AM Felicísimo, F Cabezas, AR Burgaz & I Martínez (�004) Wind as a long-distance dispersal vehiclein the Southern Hemisphere. Science, 304, ��44–��47.

Muñoz Cristi J (�94�) Rasgos generales de la constitución geológica de la Cordillera de la Costa, especialmente en la provincia de Coquimbo. Anales I. Congr. Panamericano Ing. Minas & Geol. (Santiago de Chile), 2, �8�-��8.

Muñoz Pizarro C (�944) El itinerario de don Claudio Gay. Boletín Museo Nacional de Historia Natural, 22, �7-44.

�08

Muñoz Pizarro C (�9�9) Sinopsis de la Flora Chilena. (�nd edition �9��), Ediciones de la Universidad de Chile.

Muñoz Pizarro C (�9��) Flores Silvestres de Chile. München.

Muñoz Pizarro C (�9�9) El Archipiélago de Juan Fernández y la conservación de sus recursos naturales renovables. Reprint �974, Museo Nacional de Historia Natural (Santiago) Serie Educativa, 9, �7-47.

Muñoz Pizarro C (�97�) Chile: plantas en extinción. Santiago.

Muñoz Pizarro C & E Pisano (1947) Estudio de la vegetación y flora de los Parques Nacionales Fray Jorge y Talinay. Agricultura Técnica (Santiago), 7, 7�-�90.

Muñoz-Schick M (�980) Flora del Parque Nacional Puyehue. Ed. Universitaria, Santiago.

Muñoz-Schick M & A Moreira-Muñoz (�000) Los Géneros Endémicos de Monocotiledóneas de Chile Continental. Museo Nacional de Historia Natural, Chile. Online version: http://www.mnhn.cl/apuntes/botanica/Principalbot.htm

Muñoz-Schick M & A Moreira-Muñoz (�00�) El Herbario Nacional y la Conservación de la Flora Chilena. Museo Nacional de Historia Natural, Chile. Online version: http://www.mnhn.cl/botanica/Herbario/index.html

Muñoz-Schick M, R Pinto, A Mesa & A Moreira-Muñoz (�00�) “Oasis de neblina” en los cerros costeros del sur de Iquique, región de Tarapacá, Chile, durante el evento El Niño �997–�998. Revista Chilena de Historia Natural, 74, �89–40�.

Mutke J & W Barthlott (�00�) Patterns of vascular plant diversity at continental to global scales. Biol. Skr., 55, ���-���.

Nair PKK (�979) The palynological basis for the triphyletic theory of angiosperms. Grana, 18, �4�-�44.

Near TJ & MJ Sanderson (�004) Assessing the quality of molecular divergence time estimates by fossil calibrations and fossil-based model selection. Philosophical Transactions of the Royal Society of London, Series B, 359, �477-�48�.

Nelson EC (�98�) Phytogeography of southern Australia. Ecological biogeography of Australia (ed. by A Keast), pp. 7��–7�9. Dr W. Junk, The Hague.

Nelson G (�978) From Candolle to Croizat: Comments on the History of Biogeography. Journal of the History of Biology, 11, �89-�0�.

Nelson G & N Platnick (�98�) Systematics and biogeography: Cladistics and vicariance. Columbia Univ. Press, New York.

Nelson G & PY Ladiges (�00�) Gondwana, vicariance biogeography and the New York School revisited. Australian Journal of Botany, 49, �89–409.

Nickrent DL, CL Parkinson, JD Palmer & RJ Duff (�000) Multigene phylogeny of land plants with special reference to bryophytes and the earliest land plants. Molecular Biology and Evolution, 17, �88�-�89�.

Nielsen SN (�00�) The Triassic Santa Juana Formation at the lower Biobio River, south central Chile. Journal of South American Earth Sciences, 19, �47-���

Nihei SS (�00�) Misconceptions about parsimony analysis of endemicity. Journal of Biogeography, 33, �099-��0�.

Nur A & Z Ben-Avraham (1977) Lost Pacifica continent. Nature, 270, 4�-4�.

Nur A & Z Ben-Avraham (1981a) The lost Pacifica continent: A mobilistic speculation. Vicariance Biogeography (ed. by G Nelson & DE Rosen), pp. �4�-��8, Columbia University Press.

Nur A & Z Ben-Avraham (�98�b) Volcanic gaps and the consumption of aseismic ridges in South America. Nazca Plate: Crustal Formation and Andean Convergence (ed. by LVD Kulm, J Dymond, EJ Dasch et al.) Geological Society of America Memoir, 154, 7�9-740.

Nyffeler R (�00�) Phylogenetic relationships in the cactus family (Cactaceae) based on evidence from trnK/matK and trnL-trnF sequences. American Journal of Botany, 89, ���-���.

�09

Ochsenius, C. (�999) The Neotropical Aridity. CCOS PCSS Geowissenschaftliche Beiträge, Baden-Württemberg, Santiago.

Olmstead RG, CW de Pamphilis, AD Wolfe, ND Young, WJ Elisens & PA Reeves (�00�) Disintegration of the Scrophulariaceae. American Journal of Botany, 88, �48-���.

Osterkamp WR & CR Hupp (�99�) The Evolution of Geomorphology, Ecology, and Other Composite Sciences. The Scientific Nature of Geomorphology: Proceedings of the 27th Binghamton Symposium in Geomorphology held 27-29 September 1996 (ed. by BL Rhoads & CE Thorn), pp. 4��-44�. John Wiley & Sons Ltd.

Page CN (�990) Cupressaceae. The Families and Genera of Vascular Plants (ed. by K Kubitzki), vol. I Pteridophytes and Gymnosperms (ed. by KU Kramer & PS Green), pp. �0�-���.

Palmer JD, DE Soltis & MW Chase (�004) The plant tree of life: An overview and some points of view. American Journal of Botany, 91, �4�7-�44�.

Parenti LR (�007) Common Cause and Historical Biogeography. Biogeography in a Changing World (ed. by MC Ebach & RS Tangney), pp. ��-8�. Systematics Association Special Volumes, CRC Press.

Parenti LR & MC Ebach (in prep.) Comparative Biogeography: Discovering and Classifying Biogeographical Patterns on a Dynamic Earth.

Pastorino MJ & LA Gallo (2002) Quaternary evolutionary history of Austrocedrus chilensis, a cypress native to the Andean–Patagonian forest. Journal of Biogeography, 29, ���7–��78.

Peña RC, L Iturriaga, G Montenegro & BK Cassels (�000) Phylogenetic and Biogeographic Aspects of Sophora Sect. Edwardsia (Papilionaceae). Pacific Science, 54, ��9-��7.

Perring FH & SM Walters (eds.) (�9��) Atlas of the British Flora. London-Edinburgh.

Philippi F (1875) La flora de las islas San Ambrosio i San Félix. Anales Univ. Chile, 47, �8�-�94.

Philippi F (�88�) Catalogus plantarum vascularium chilensium adhuc descriptarum. Anales Univ. de Chile, 59, 49-4��.

Philippi RA (�870) Vegetation der Inseln San Ambrosio und S. Felix. Bot. Zeitung (Berlin), 28, 49�-�0�.

Pisano E (�977) Fitogeografía de Fuego-Patagonia chilena I. Las comunidades vegetales entre las latitudes �� y �� S. Anales Instituto de la Patagonia, 8, ���-��0.

Pisano E (1981) Bosquejo fitogeográfico de Fuego-Patagonia. Anales Instituto de la Patagonia, 12, ��9-�7�.

Pisano E (�98�) Comunidades vegetales en el sector Norte de la Península Muñoz Gamero (Última Esperanza, Magallanes). Anales Instituto de la Patagonia, 14, 8�-�0�. Pliscoff P (�00�) Priorización de áreas para fortalecer la conservación de la flora arbórea nativa en la zona mediterránea de Chile. MScThesis. Universidad de Chile, Santiago.

Poole I, AMW Mennega & D Cantrill (�00�) Valdivian ecosystems in the Late Cretaceous and Early Tertiary ofAntartica: further evidence from myrtaceous and eucryphiaceous fossil wood. Review of Palaeobotany and Palynology, 124, 9–�7.

Posada D & Crandall KA (�00�) Intraspecific gene genealogies: trees grafting into networks. Trends in Ecology and Evolution, 16, �7–4�.

Potgieter K & VA Albert (�00�) Phylogenetic relationships within Apocynaceae s.l. based on trnL intron and trnL-F spacer sequences and propagule characters. Annals of the Missouri Botanical Garden, 88, ���-�49.

Pratt D (�000) Plate Tectonics: A Paradigm Under Threat. Journal of Scientific Exploration, 14, �07-���.

Premoli AC, T Kitzberger & TT Veblen (�000) Isozyme variation and recent biogeographical history of the long-lived conifer Fitzroya cupressoides. Journal of Biogeography, 27, ���-��0.

��0

Pryer KM, E Schuettpelz, PG Wolf, H Schneider, AR Smith & R Cranfill (2004) Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. American Journal of Botany, 91, ��8�-��98.

Pulquério MJF & RA Nichols (�007) Dates from the molecular clock: how wrong can we be? Trends in Ecology and Evolution, ��, �80-�84.

Qian H (1999) Floristic analysis of vascular plant genera of North America north of Mexico: characteristics of phytogeography. Journal of Biogeography, 26, ��07–����.

Qian H, J-S Song, P Krestov, Q Guo, Z Wu, X Shen & X Guo (2003) Large-scale phytogeographical patterns in East Asia in relation to latitudinal and climatic gradients. Journal of Biogeography, 30, ��9–�4�.

Quinn CJ & RA Price (2003) Phylogeny of the Southern Hemisphere Conifers. IV International Conifer Conference (ed. by RR Mill). ISHS Acta Horticulturae, 615, ��9-���.

Quintanilla V (1974) La carta bioclimática de Chile Central. Revista Geográfica de Valparaíso, 5, ��-�8.

Quintanilla V (1983) Biogeografía. Colección Geografía de Chile. Instituto Geográfico Militar, Santiago.

Quintanilla V (1985) Carta fitogeográfica de Chile mediterráneo. Contribuciones Científicas y Tecnológicas, Area Geociencias (U. de Santiago), 70, �-�0. Quintanilla V (1988) Fitogeografía y cartografía de la vegetación de Chile árido. Contribuciones Científicas y Tecnológicas, Area Geociencias (U. de Santiago), 82, �-�8.

Rahmanzadeh R, K Muller, E Fischer, D Bartels & T Borsch (�00�) The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales). Plant Biology, 7, �7-78.

Rapalini AE (�00�) The accretionary history of southern South America from the latest Proterozoic to the Lare Palaeozoic: some palaeomagnetic constraints. Terrane Processes at the Margins of Gondwana (ed. by APM Vaughan, PT Leat & RJ Pankhurst), �0�-��8. Geological Society Special Publications (London) n° �4�.

Rapoport EH (�98�) Areography: Geographical Strategies of Species. Pergamon Press.

Ratzel F (�90�) Der Lebensraum: eine biogeographische Studie. Laupp‘sche Buchhandlung, Tübingen.

Raup HM (�94�) Trends in the development of geographic botany. Annals of the Association of American Geographers, 32, ��9-��4.

Raven PH (1963) Amphitropical relationships in the floras of North and South America. Quarterly Review of Biology, 38, ���–�77.

Raven PH (�97�) Plant species disjunctions: a summary. Annals of the Missouri Botanical Garden, 59, ��4–�4�.

Raven PH & DI Axelrod (�974) Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden, 61, ��9–�7�.

Reason P & BC Goodwin (�999) Toward a science of qualities in organizations: lessons from Complexity theory and postmodern biology. Concepts and Transformations, 4, �8�-��7.

Reeves C & M de Wit (�000) Making ends meet in Gondwana: retracing the transforms of the Indian Ocean and reconnecting continental shear zones. Terra Nova, 12, �7�–�80.

Reiche K (�89�-�9��) Flora de Chile, 6 vols. Santiago.

Reiche K (1905) La distribución geográfica de las Compuestas de la flora de Chile. Anales del Museo Nacional de Chile, Sección Botánica, 17, �-4�.

Reiche K (�907) Grundzüge der Pflanzenverbreitung in Chile. Die Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, Vol. VIII. (ed. by A Engler & O Drude) Verlag von V. Engelmann, Leipzig.

���

Renner SS (�004) Variation in diversity among Laurales, Early Cretaceous to Present. Biol. Skr. 55, 44�-4�8.

Renner SS (�00�) Relaxed molecular clocks for dating historical plant dispersal events. Trends in Plant Science, 10, ��0-��8.

Renner SS, D Murray & D Foreman (�000) Timing transantarctic disjunctions in the Atherospermataceae (Laurales): Evidence from coding and noncoding chloroplast sequences. Systematic Biology, 49, �79-�9�.

Rhoads BL & CE Thorn (�99�) Toward a Philosophy of Geomorphology. The Scientific Nature of Geomorphology: Proceedings of the 27th Binghamton Symposium in Geomorphology held 27-29 September 1996. (ed. by BL Rhoads & CE Thorn), pp. ���-�4� John Wiley & Sons Ltd.

Ribichich AM (2002) El modelo clásico de la fitogeografía de Argentina: un análisis crítico. Interciencia, 27, ��9-�7�.

Riccardi AC (�988) The Cretaceous System of Southern South America. The Geological Society of America Memoir, 168, �-�4�.

Richardson JE, MF Fay, QCB Cronk, D Bowman & MW Chase (2000) A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences. American Journal of Botany, 87, ��09–���4.

Richter M (�99�) Klimaökologische Merkmale der Küstenkordillere in der Region Antofagasta (Nordchile). Geoökodynamik, 16, �8�-���.

Richter M (�997) Allgemeine Pflanzengeographie. Teubner Studienbücher der Geographie, Stuttgart.

Richter M, T Fickert & F Grüninger (2004) Pflanzen auf schuttbedeckten Gletschern – wandernde Kuriositäten. Geoöko, 25, ���-���.

Riddle BR (�00�) Is biogeography emerging from its identity crisis? Journal of Biogeography, 32, �8�–�8�.

Rivas-Martínez S & G Navarro (�000). Figure I.�4, in G Navarro & M Maldonado (�00�) Geografía Ecológica de Bolivia: Vegetación y Ambientes Acuáticos. Ed. Centro de Ecología Simón I. Patiño, Cochabamba, Bolivia.

Romero EJ (1993) South American Paleofloras. Biological Relationships between Africa and South America (ed. by P Goldblatt), pp. ��-8�, Yale University Press, New Haven/London.

Ronse De Craene LP (�004) Floral development of Berberidopsis corallina: a crucial link in the evolution of flowers in the Core Eudicots. Annals of Botany, 94, 74�–7��.

Rosen BR (�988) From fossils to earth history: applied historical biogeography. Analytical biogeography: an integrated approach to the study of animal and plant distributions (ed. by AA Myers & PS Giller), pp. 4�7–48�. Chapman & Hall, London.

Rosen BR (�99�) Empiricism and the biogeographical black box: concepts and methods in marine palaeobiogeography. Palaeogeography, Palaeoclimatology, Palaeoecology, 92, �7�–�0�.

Rothmaler W (�9��) Allgemeine Taxonomie und Chorologie der Pflanzen. Grundzüge der speziellen Botanik. Libri Botanici vol. �, IHW Verlang. Reprint �99�.

Rothwell GW (�987) Complex Paleozoic Filicales in the evolutionary radiation of ferns. American Journal of Botany, 74, 4�8–4��.

Rovito SM, MTK Arroyo & P Pliscoff (�004) Distributional modelling and parsimony analysis of endemicity of Senecio in the Mediterranean-type climate area of Central Chile. Journal of Biogeography, 31, ����-����.

Rundel PW, MO Dillon & B Palma (�99�) Flora y vegetación del Parque Nacional Pan de Azúcar en el Desierto de Atacama. Gayana Botánica, 53, �9�-���.

Rutschmann F (�00�) Molecular dating of phylogenetic trees: A brief review of current methods that estimate divergence times. Diversity and Distributions, 12, ��-48.

���

Salomon M (�00�). Evolutionary biogeography and speciation: essay on a synthesis. Journal of Biogeography, 28, ��-�7.

Sanders RW, TF Stuessy, C Marticorena & M Silva O (�987) Phytogeography and evolution of Dendroseris and Robinsonia, tree Compositae of the Juan Fernández Islands. Opera Botanica, 92, �9� ���.

Sanderson MJ (�997) A nonparametric approach to estimating divergence times in the absence of rate constancy. Molecular Biology and Evolution, 14, ���8–����.

Sanderson MJ (�00�) Estimating absolute rates of molecular evolution and divergence times: a penalized likelihood approach. Molecular Biology and Evolution, 19, �0�–�09.

Sanderson MJ (�00�) Molecular data from �7 proteins do not support a Precambrian origin of land plants. American Journal of Botany, 90, 9�4–9��.

Sanderson MJ & JA Doyle (2001) Sources of error and confidence intervals in estimating the age of angiosperms from rbcL and �8S rDNA data. American Journal of Botany, 88, �499–����.

Sanderson MJ, JL Thorne, N Wikström & K Bremer (�004) Molecular Evidence On Plant Divergence Times. American Journal of Botany, 91, ����–����.

Sanmartín I & F Ronquist (�004) Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology, 53, ���–�4�.

Sanmartín I, L Wanntorp & RC Winkworth (�007) West Wind Drift revisited: testing for directional dispersal in the Southern Hemisphere using event-based tree fitting. Journal of Biogeography, 34, �98–4��.

Santos CMD (�00�) Parsimony analysis of endemicity: time for an epitaph? Journal of Biogeography, 32, ��84–��8�.

Scher HD & EE Martin (�00�) Timing and climatic consequences of the opening of Drake Passage. Science, 312, 4�8-4�0.

Schmithüsen J (�9��) Die räumliche Ordnung der chilenischen Vegetation. Bonner Geographische Abhandlungen, 17, �–8�.

Schmithüsen J (�9�8) Allgemeine Vegetationsgeographie. �d ed., De Gruyter, Berlin.

Schneider H, E Schuettpelz, KM Pryer, R Cranfill, S Magallón & R Lupia (2004) Ferns diversified in the shadow of angiosperms. Nature, 428, ���–��7.

Schönfelder P & A Bresinsky (�990) Verbreitungsatlas der Farn- und Blütenpflanzen Bayerns. Ulmer Verlag.

Schouw JF (�8��) Grundzüge einer allgemeinen Pflanzengeographie (+ Atlas). Reimer, Berlin.

Schroeder F-G (�998) Lehrbuch der Pflanzengeographie. Quelle & Meyer, Wiesbaden.

Schröter C (1913) Genetische Pflanzengeographie (Epiontologie). Handwörterbuch der Naturwissenschaften, vol. 4, pp. 907-94�. Gustav Fischer Verlag, Jena.

Schuettpelz E & SB Hoot (�004) Phylogeny and biogeography of Caltha (Ranunculaceae) based on chloroplast and nuclear DNA sequences. American Journal of Botany, 91, �47-���.

Schwarz O (�9�8) Phytochorologie als Wissenschaft, am Beispiele der vorderasiatischen Flora. Feddes Rep. Beiheft �00, Dahlem, Berlin.

Sclater PL (�8�8) On the general geographical distribution of the members of the class Aves. Zoological Journal of Linnean Society, 2, ��0-�4�.

Scotese CR & WS McKerrow (�990) Revised world maps and introduction. Palaeozoic palaeogeography and biogeography (ed. by WS McKerrow & CR Scotese) Geological Society Memoir, 12, �–��.

Scotland RW & AH Wortley (�00�) How many species of seed plants are there? Taxon, 52,�0�–�04.

���

Sempere T, A Hartley & P Roperch (�00�) Comment on “Rapid Uplift of the Altiplano Revealed Through ��C-�8O Bonds in Paleosol Carbonates”. Science, 314, 7�0b.

Setoguchi H, TA Osawa, J-C Pintaud, T Jaffre & JM Veillon (�998) Phylogenetic relationships within Araucariaceae based on rbcL gene sequences. American Journal of Botany, 85, ��07-����.

Shields O (�997) Rapid earth expansion: An eclectic view. Gondwana Research, 1, 9�-94.

Shields O (1998) Upper Triassic Pacific vicariance as a test of geological theories. Journal of Biogeography, 25, �0�-���.

Simpson BB (�97�) Contrasting Modes of Evolution in Two Groups of Perezia (Mutisieae; Compositae) of Southern South America. Taxon, 22, ���-���.

Simpson BB & JL Neff (�98�) Plants, their pollinating bees, and the Great American Interchange. The Great American Biotic Interchange (ed. by FG Stehli & SD Webb), pp. 4�7–4��, Plenum, New York.

Simpson BB, JA Tate & A Weeks (�00�) The biogeography of Hoffmannseggia (Leguminosae, Caesalpinioideae, Caesalpinieae): a tale of many travels. Journal of Biogeography, 32, ��–�7.

Simpson GG (�9��) Evolution and Geography: An essay on historical biogeography with special reference to mammals. Condon lectures.

Simpson GG (�980) Splendid Isolation: The Curious History of South American Mammals. Yale University Press, New Haven, Connecticut.

Singh I (�944) The Gondwana and the Gonds. University Publishers, Lucknow.

Sklenar P & H Balslev (2007) Geographic flora elements in the Ecuadorian superpáramo. Flora, 202, �0–��.

Skottsberg C (�90�) Vegetationsbilder aus Feuerland, von den Falkland-Inseln und von Südgeorgien. Karsten und Schenck Vegetationsbilder, 4 (�-4), �� lám.

Skottsberg C (�9�0 a) Botanische Ergebnisse der schwedischen Expedition nach Patagonien unde dem Feuerlande �907-1909. I. Übersicht über die wichtigsten Pflanzenformationen Südamerikas S von 41°, ihre geographische Verbreitung und Beziehungen zum Klima. Kungl. Svenska Vetenskapakademiens Handlingar, 46, (�), �-�8 + � map.

Skottsberg C (�9�0 b) Vegetationsbilder von den Juan Fernández Insel. Karsten und Schenck Vegetationsbilder, 8 (�), 7-��.

Skottsberg C (�9��) Botanische Ergebnisse der schwedischen Expedition nach Patagonien und dem Feuerlande �907-�909. V. Die Vegetationsverhältnisse längs der Cordillera de los Andes südlich von 4� Grad südlicher Breite. Ein Beitrag zur Kenntnis der Vegetation in Chiloé, West-Patagonien, dem andinen Patagonien und Feuerland. Kungl. Svenska Vetenskapsakademien Handlingar, 56 (�) �-���, �� lám. Stockholm, Almqvist & Wiksell.

Skottsberg C (�9�0-�9��) (ed.) Natural History of Juan Fernandez and Easter Island. � vol. Almqvist & Wiksell, Uppsala.

Skottsberg C (�9��) Juan Fernandez and Hawaii. A phytogeographical discussion. Bernice P. Bishop Mus. Spec. Publ., ��, �–47.

Skottsberg C (�9��) La población botánica y zoológica de las islas chilenas del Pacífico. Imprenta Universitaria, Santiago. Translated by G. Looser from the original French: Le peuplement des îles pacifiques du Chili. Contribution à l’étude du peuplement zoologique et botanique des îles du Pacifique (ed. by L Berland, J Berlioz, EH Bryan et al.) (�9�4), pp. �7�–�80. Société de Biogéographie, Paris.

Skottsberg C (�9�7) Die Flora der Desventuradas-Inseln (San Félix und San Ambrosio) nach Sammlungen F. Johows. Göteborgs Kungl. Vetensk.-Vitterh.-Samh. Handl., ser, �, B, 5, �-88.

Skottsberg C (�9��) Weitere Beiträge zur Flora der Insel San Ambrosio. Arkiv för Botanik, 2, ��.

Skottsberg C (1956) Derivation of the flora and fauna of Juan Fernandez and Easter Island. The natural history of the Juan Fernandez and Easter Islands. Vol. 1. Botany (ed. by C. Skottsberg), pp. �9�–40�, Almqvist & Wiksell, Uppsala.

��4

Skottsberg C (1963) Zur Naturgeschichte der Inseln San Ambrosio (Islas Desventuradas, Chile) 2. Blütenpflanzen. Arkiv för Botanik, 4, 4�4-490 (+ tables �-��).

Sluys R (1994) Explanations for biogeographic tracks across the Pacific Ocean: a challenge for paleogeography and historical biogeography. Progress in Physical Geography, 18, 4�–�8.

Smith AD & Ch Lewis (�999) Differential rotation of lithosphere and mantle and the driving forces of plate tectonics. Geodynamics, 28, 97-���.

Smith CH (�989) Historical Biogeography: Geography as Evolution, Evolution as Geography. New Zealand Journal of Zoology, 16, 77�-78�. [http//www.wku.edu/~smithch/essays/SMITH89.htm]

Smith SDW & DA Baum (2006) Phylogenetics of the florally diverse Andean clade Iochrominae (Solanaceae). American Journal of Botany, 93, ��40-����.

Soja EW (�989) Postmodern Geographies: the reassertion of space in critical social theory. Verso, London.

Sokal AD & J Bricmont (�998) Fashionable Nonsense: Postmodern Intellectuals’ Abuse of Science. Picador USA, New York,

Sokal RR & PHA Sneath (�9��) The Principles of Numerical Taxonomy. W. H. Freeman, San Francisco.

Solbrig OT (�97� a) New approaches to the study of disjunctions with special emphasis on the American amphitropical desert disjunctions. Taxonomy, phytogeography and evolution (ed. by DH Valentine), pp. 8�-�00, Academic Press, London.

Solbrig OT (1972 b) The floristic disjunctions between the Monte in Argentina and the Sonoran Desert in Mexico and the United States. Annals of the Missouri Botanical Garden, 59, �0�-��4.

Squeo FA, JR Gutiérrez & IR Hernández (eds.) �004. Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena, Chile.

Stevens PF (�994) The development of Biological Systematics. Columbia Univ. Press, New York.

Stevens PF (�00� onwards) Angiosperm Phylogeny Website, version 7, may �00�, [http://www.mobot.org/MOBOT/research/APweb/]

Stewart WN & GW Rothwell (�99�) Paleobotany and the evolution of plants. �nd ed. Cambridge University Press, Cambridge, UK.

Stoddart DR (�98�) On Geography and its History. Basil Blackwell, Oxford, UK.

Storey BC (�99�) The role of mantle plumes in continental breakup: case histories from Gondwanaland, Nature, 377, �0� – �08.

Stuessy TF (�004) A transitional-combinational theory for the origin of angiosperms. Taxon, 53, �-��

Stuessy TF (�00�) Principles and practice of plant taxonomy. Taxonomy and Plant Conservation (ed. by E Leadlay & S Jury), pp. ��-44, Cambridge Univ. Press.

Stuessy TF & M Ono (eds) (�998) Evolution and Speciation of Island Plants. Cambridge University Press.

Stuessy TF & Ch Taylor (�99�) Evolución de la Flora Chilena. Flora de Chile vol. 1 (ed. by C Marticorena & R Rodríguez) pp. 8�-��8, Universidad de Concepción, Chile.

Stuessy TF, KA Foland, JF Sutter, RW Sanders & M Silva (1984a) Botanical and geological significance of Potassium-Argon dates from the Juan Fernandez Islands. Science, ���, 49–��.

Stuessy TF, RW Sanders & M Silva (1984b) Phytogeography and evolution of the flora of the Juan Fernández Islands: a progress report. Biogeography of the Tropical Pacific (ed . by FJ Redovsky, PH Raven & SH Sohmer), pp. ��-�9. Assoc. Syst. Collections and B.P. Bishop Museum, Lawrence.

���

Stuessy TF, DJ Crawford & C Marticorena C (1990) Patterns of phylogeny in the endemic vascular flora of the Juan Fernandez Islands, Chile. Systematic Botany, 15, ��8–�4�.

Stuessy TF, DJ Crawford, C Marticorena & M Silva O (�998a) Isolating mechanisms and modes of speciation in endemic angiosperms of the Juan Fernández Islands. Evolution and Speciation of Island Plants (ed. by TF Stuessy & M Ono), pp. 79-9�, Cambridge University Press.

Stuessy TF, U Swenson, C Marticorena, O Matthei & DJ Crawford (�998b) Loss of plant diversity and extinction on Robinson Crusoe Islands, Chile. Rare, threatened and endangered floras of Asia and the Pacific Rim (ed. by CI Peng & PP Lowry). Academia Sinica monographs, 16, �4�-��7.

Stuessy TF, K Tremetsberger, AN Müllner, J Jankowicz, Y-P Guo, CM Baeza & RM Samuel (�00�) The melding of systematics and biogeography through investigations at the populational level: examples from the genus Hypochaeris (Asteraceae). Basic and Applied Ecology, 4, �87–�9�.

Stuessy TF, G Jakubowsky, R Salguero G, M Pfosser, PM Schlüter, TFer, B-Y Sun & H Kato (�00�) Anagenetic evolution in island plants. Journal of Biogeography, 33, ���9-����.

Suessenguth K (�9�8) Neue Ziele der Botanik. Lehmanns Verlag, München/Berlin.

Sun G, DL Dilcher, S Zheng & Z Zhou (�998) In Search of the First Flower: A Jurassic Angiosperm, Archaefructus, from Northeast China. Science, 282, ��9�-��9�.

Sun G, Q Ji, DL Dilcher, S Zheng, KC Nixon & X Wang (2002) Archaefructaceae, a New Basal Angiosperm Family. Science, 296, 899-904.

Swenson U & K Bremer (1997) Pacific biogeography of the Asteraceae genus Abrotanella (Senecioneae, Blennospermatinae). Systematic Botany, 22, 49�-�08.

Swenson U, TF Stuessy, M Baeza & DJ Crawford (�997) New and historical plant introductions, and potential pests in the Juan Fernández Islands, Chile. Pacific Science, 51, ���-���.

Swenson U, RS Hill & S McLoughlin (�00�) Biogeography of Nothofagus supports the sequence of Gondwana break-up. Taxon, 50, �0��–�04�.

Szumik CA, F Cuezzo, P Goloboff & AE Chalup (�00�) An optimality criterion to determine areas of endemism. Systematic Biology, 51, 80�–8��.

Szumik C & P Goloboff (�004) Areas of endemism: an improved optimality criterion. Systematic Biology, 53, 9�8–977.

Takhtajan A (�9��) The Origin of Angiospermous Plants, 2nd ed. (original Russian edition, Moscow).

Takhtajan A (�978) The Floristic Regions of the World (original Russian edition, Leningrad). Translated as Floristic Regions of the World by T.J. Crovello, ed. by A. Cronquist (�98�), U. of California. Reprint �988, B. Singh, M. P. Singh, Dehra Dun., India.

Takhtajan A (�98�) Flowering plants, origin and dispersal. From the Russian by C. Jeffrey, Kew (�9�9). Reprint �98�, Otto Koeltz Science Publishers, Koenigstein, Germany.

Taylor DW, H Li, J Dahl, FJ Fago, D Zinniker JM & Moldowan (�00�) Biogeochemical evidence for the presence of the angiosperm molecular fossil oleanane in Paleozoic and Mesozoic non-angiospermous fossils. Paleobiology, 32, �79-�90.

Taylor EL, TN Taylor & NR Cuneo (�99�) The present is not the key to the past: a polar forest from the Permian ofAntarctica. Science, 257, ��7�–��77.

Taylor, T Griffith (1921) The Evolution and Distribution of Race, Culture, and Language. Geographical Review, 11, �4-��9.

Teillier S & Ch Taylor (�997) Maireana Moq., un nuevo género para Chile (Islas Desventuradas). Gayana Botánica, 54, ��-�7.

���

Teillier S, A Hoffmann, F Saavedra & L Pauchard (�994) Flora del Parque Nacional El Morado (Región Metropolitana, Chile). Gayana Botánica, 51, ��-47.

Teillier S, G Aldunate, P Ridemann & H Niemayer (�00�) Flora de la Reserva Nacional Clarillo. CONAF, U. de Chile.

Templeton AR (�998) Species and Speciation: Geography, Population Structure, Ecology, and Gene Trees. Endless Forms: Species and Speciation (ed. by DJ Howard & SH Berlocher), pp. ��-4�, Oxford University Press.

Thiselton-Dyer WT (�909) Geographical distribution of plants. Darwin and modern science. (ed. by AC Seward), pp. �98-��8. Cambridge University Press, Cambridge.

Thomas HH (�947) The history of plant form. Advanc. Science, 4, �4�-��4.

Thorne JL & H Kishino (�00�) Divergence time and evolutionary rate estimation with multilocus data. Systematic Biology, 51, �89–70�.

Thorne RF (�00�) How many species of seed plants are there? Taxon, 51, ���-���.

Torres T & M Rallo (�98�) Anatomía de troncos fósiles del Cretácico Superior de Pichasca en el norte de Chile. Anais II Congreso Latinoaméricano Paleontología. Porto Alegre, Brasil, vol. 2, �8�-�98.

Torres T & M Philippe (�00�) New species of Coniferales and Ginkgoales from the Lias of La Ligua (Chile) with an evaluation of paleoxylological record in the Jurassic of South America. Revista Geológica de Chile, 29, ���-���.

Tremetsberger K, H Weiss-Schneeweiss, TF Stuessy, R Samuel, G Kadlec, MA Ortiz & S Talavera (�00�) Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South American Hypochaeris (Asteraceae, Cichorieae). Molecular Phylogenetics and Evolution, 35, �0�–���.

Treviranus GR (�80�) Biologie, oder Philosophie der lebenden Natur. Röwer, Göttingen.

Troncoso A & R Herbst (�999) Triassic Ginkgoales from northern Chile. Revista Geológica de Chile, 26, ���-�7�

Troncoso A & EJ Romero (1998) Evolución de las comunidades florísticas en el extremo sur de Sudamérica durante el Cenofítico. Proceedings of the Congreso Latinoamericano de Botánica, No. � (ed. by R Fortunato & N Bacigalupo), pp. �49-�7�, Monographs in Systematic Botany from the Missouri Botanical Garden.

Troncoso A, C Villagrán & M Muñoz-Schick (�980) Una nueva hipótesis acerca del origen y edad del bosque de Fray Jorge (Coquimbo, Chile). Boletín Museo Nacional de Historia Natural (Chile), 37, ��7-���.

Turner S, M Regelous, S Kelley, C Hawkesworth & M Mantovani (�994) Magmatism and continental break-up in the South Atlantic: high precision 40Ar–�9Ar geochronology. Earth and Planetary Science Letters, 121, ���–�48.

Turrill WB (�9��) Pioneer Plant Geography: the Phytogeographical Researches of Sir Joseph Dalton Hooker. Contributions from the Int. Biohistorical Comission, N°�. The Hague, Netherlands.

Upchurch P (�00�) What’s wrong with palaeobiogeography? Abstracts. Palaeogeography and Palaeobiogeography: Biodiversity in Space and Time, University of Cambridge, �0-��.04.0�.

Upchurch P, CA Hunn & DB Norman (�00�) An analysis of dinosaurian biogeography: evidence for the existence of vicariance and dispersal patterns caused by geological events. Proceedings of the Royal Society of London Series B-Biological Sciences, 269, ���-���.

van Balgooy MMJ (1971) Plant-geography of the Pacific. Blumea Supplement, 6, �–���.

van Balgooy MMJ (�97�) Pacific Plant Areas, vol. 3. Leiden, The Netherlands.

van Balgooy MMJ (�984) Pacific Plant Areas, vol. 4. Leiden, The Netherlands.

van Balgooy MMJ (ed.) (�99�) Pacific Plant Areas, vol. 5. Rijksherbarium/Hortus Botanicus, Leiden University, The Netherlands.

��7

van der Hammen T & AM Cleef (1983) Datos para la historia de la flora andina. Revista Chilena de Historia Natural, 56, 97-�07.

van Steenis CGGJ (�9��) The theory of Land Bridges in Botany, with particular references to tropical plants. Blumea (Nederland), 11, ���-�4�.

van Steenis CGGJ (�9��) Pacific Plant Areas, vol. 1. Manila.

van Steenis CGGJ & MMJ van Balgooy (�9��) Pacific Plant Areas, vol. 2. Blumea Supplement vol. �.

Van Veller MGP, DR Brooks & M Zandee (�00�) Cladistic and phylogenetic biogeography: the art and the science of discovery. Journal of Biogeography, 30, ��9–��9.

Varela F (�000) El Fenómeno de la Vida. Editorial Dolmen, Santiago de Chile.

Veit H (�994) Holocene landscape and climate evolution of the Central Andes. Zentralblatt Geologie und Paläontologie, 7/8, 887-89�.

Verdú M, P Dávila, P García-Fayos, N Flores-Hernández & A Valiente-Banuet (�00�) ‘Convergent’ traits of mediterranean woody plants belong to pre-mediterranean lineages. Biological Journal of the Linnean Society, 78, 4��–4�7.

Vester H (�940) Die Areale und Arealtypen der Angiospermen-Familien. Doctoral thesis, Naturwissenschaftlichen Fakultät der Ludwig-Maximilians Universität zu München. Akad. Verlagsgesellschaft Leipzig.

Villa-Martínez R, C Villagrán & B Jenny (�00�) The last 7�00 cal yr B.P. of westerly rainfall in Central Chile inferred from a high-resolution pollen record from Laguna Aculeo (�4°S). Quaternary Research, 60, �84–�9�

Vijverberg K, THM Mes & K Bachmann (�999). Chloroplast DNA evidence for the evolution of Microseris (Asteraceae) in Australia and New Zealand after long-distance dispersal from western North America. American Journal of Botany, 86, �448–�4��.

Villagrán C (�00�) Flora y vegetación del Parque Nacional Chiloé: guía de excursion botánica por la Cordillera de Piuchué. Conaf, Chiloé.

Villagrán C (�00�) Botánica Indígena de Chile. Revista Chagual (Santiago), 1, �0-��.

Villagrán C & LF Hinojosa (1997) Historia de los bosques del sur de Sudamérica, II: Análisis fitogeográfico. Revista Chilena de Historia Natural, 70, �4�-��7.

Villagrán C, A León & FA Roig (�004) Paleodistribución del alerce y ciprés de las Guaitecas durante períodos interestadiales de la Glaciación Llanquihue: provincias de Llanquihue y Chiloé, Región de Los Lagos, Chile. Revista Geológica de Chile, 31, ���-���.

Vinnersten A & K Bremer (�00�) Age and Biogeography of Major Clades in Liliales. American Journal of Botany, 88, ��9�–�70�.

Vogel K (�990) The expansion of the earth – an alternative model to plate tectonics theory. Critical aspects of the plate tectonics theory vol. II. (ed. by V Belousov), pp. �9-�4, Athen.

von Hagen KB & JW Kadereit (�00�) The phylogeny of Gentianella (Gentianaceae) and its colonization of the southern hemisphere as revealed by nuclear and chloroplast DNA sequence variation. Organisms Diversity and Evolution, 1, ��–79.

von Ihering H (�89�) Das neotropische Florengebiet und seine Geschichte. Engler’s Botanische Jahrbücher, 17, �-�4.

von Ihering H (�907) Archhelenis und Archinotis: Gesammelte Beiträge zur Geschichte der Neotropischen Region. W. Engelmann, Leipzig.

von Wettstein R (�898) Grundzüge der geographisch-morphologischen Methode der Pflanzensystematik. Jena, Verlag Gustav Fischer.

��8

Wagner M (�8�8) Die Darwinsche Theorie und das Migrationsgesetz der Organismen. Leipzig.

Wagstaff SJ & PJ Garnock-Jones (�998) Evolution and biogeography of the Hebe complex (Scrophulariaceae) inferred from ITS sequences. New Zealand Journal of Botany, 36, 4��–4�7.

Wagstaff SJ, K Martinsson & U Swenson (�000) Divergence estimates of Tetrachondra hamiltonii and T. patagonica (Tetrachondraceae) and their implications for austral biogeography. New Zealand Journal of Botany, 38, �87-�9�.

Wagstaff SJ, I Breitwieser & U Swenson (�00�) Origin and relationships of the austral genus Abrotanella (Asteraceae) inferred from DNA sequences. Taxon, 55, 9�-�0�.

Wallace AR (�87�) The Geographical Distribution of Animals. Macmillan & Co., London.

Wallace AR (�880) Island Life: or, The Phenomena and Causes of Insular Faunas and Floras, Including a Revision and Attempted Solution of the Problem of Geological Climates. Macmillan & Co., London, �st edition.

Wallace RS (�99�) Molecular systematic study of the Cactaceae: using chloroplast DNA variation to elucidate cactus phylogeny. Bradleya, ��, �–��.

Wallace RS & AC Gibson (�00�) Evolution and systematics. Cacti: biology and uses (ed. by PS Nobel), pp. �–��. University of California Press, Berkeley, California.

Walter H (�9�4) Grundlagen der Pflanzenverbreitung. Einführung in die Pflanzengeographie f. Studierende der Hochschulen, 2.Teil Arealkunde.

Walter H & H Straka H (�970) Arealkunde: Floristisch-historische Geobotanik. �d. edition. Series Einführung in die Phytologie, Band III, �. Teil. Ulmer Verlag.

Walter HS (�004) The mismeasure of islands: implications for biogeographical theory and the conservation of nature. Journal of Biogeography, 31, �77-�97.

Walters SM (�9��) The shaping of angiosperm taxonomy. New Phytologist, 60, 74-84.

Walters SM (1986) The name of the rose: a review of ideas on the European bias in angiosperm classification. New Phytologist, 104, ��7–�4�.

Wangerin W (�9��) Florenelemente und Arealtypen. Beiträge zur Arealgeographie der deutschen Flora. Beih. Z. Centralbl., vol. 49, Drude Festschrift.

Wanntorp L & H-E Wanntorp (�00�) The biogeography of Gunnera L.: vicariance and dispersal. Journal of Biogeography, 30, 979–987.

Waters JM & D Craw (�00�) Goodbye Gondwana? New Zealand biogeography, geology, and the problem of circularity. Systematic Biology, 55, ���–���.

Weber A (�004) Gesneriaceae and Scrophulariaceae: Robert Brown and now. Telopea, 10, �4�–�7�.

Webster GL (�99�) The Panorama of Neotropical Cloud Forests. Biodiversity and Conservation of Neotropical Montane Forests (ed. by SP Churchill, H Balslev, E Forero & JL Luteyn), pp. ��-77. New York Botanical Garden.

Wegener A (�9��) Die Entstehung der Kontinente und Ozeane. Sammlung Vieweg Nr. ��, Braunschweig.

Welch JJ & L Bromham (�00�) Molecular dating when rates vary. Trends in Ecology and Evolution, 20, ��0–��7.

Wellman CH & J Gray (�000) The microfossil record of early land plants. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 355, 7�7-7��.

Wellman CH, PL Osterloff & U Mohiuddin (�00�) Fragments of the earliest land plants. Nature, 425, �8�-�8�.

Welß W & R Lindacher R (�994) Beiträge zur Chorologie und Florenstatistik der Kanarischen Inseln. Hoppea, Denkschr. Regensb. Bot. Ges., 55, 84�-8�7.

��9

Whalen MA (�987) Systematics of Frankenia (Frankeniaceae) in North and South America. Systematic Botany Monographs, ��, �-9�.

White F (�97�) The taxonomic and ecological basis of chorology. Mitt. Bot. Staatssamml. München, 10, 9�-���.

White F (1993) The AETFAT chorological classification of Africa history. Methods and applications. Bull. Jard. Bot. Belg., 62, ���-�8�.

Wielgorskaya T (�99�) Dictionary of Generic Names of Seed Plants. Columbia Univ. Press.

Wikström N & P Kenrick (�00�) Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by use of non-parametric rate smoothing. Molecular Phylogenetics and Evolution, 19, �77–�8�.

Wikström N, V Savolainen & MW Chase (�00�) Evolution of the angiosperms: calibrating the family tree. Proc. R. Soc. Lond. B, 268, ����-���0.

Wiley EO (�978) The evolutionary species reconsidered. Systematic Zoology, 27, �7-��.

Wiley EO (�98�) Phylogenetics: The Theory and Practice of Phylogenetic Systematics. John Wiley, New York.

Wilkins AS (�00�) The evolution of developmental pathways. Sinauer Assoc., Sunderland, Mass.

Wilkinson DM (�00�). Dispersal, cladistics and the nature of biogeography. Journal of Biogeography, 30, �779-�780.

Wilkinson M (�990) A commentary on Ridley’s cladistic solution to the species problem. Biology and Philosophy, 5, 4��-44�.

Williams DL (�97�) Piptochaetium (Gramineae) and associated taxa: evidence for the Tertiary migration of plants between North and South America. Journal of Biogeography, 2, 7�–8�.

Williams DM (�007) Ernst Haeckel and Louis Agassiz: Trees That Bite and Their Geographical Dimension. Biogeography in a Changing World (ed. by MC Ebach & RS Tangney), pp. �-�9. Systematics Association Special Volumes, CRC Press.

Willis KJ & JC McElwain (�00�) The evolution of plants.Oxford University Press.

Winkworth R, D Bryant, P Lockhart, D Havell & V Moulton (�00�) Biogeographic Interpretation of Splits Graphs: Least Squares Optimization of Branch Lengths. Systematic Biology, 54, ��-��.

Wolf PG, H Schneider & TA Ranker (�00�) Geographic distributions of homosporous ferns:does dispersal obscure evidence of vicariance? Journal of Biogeography, 28, ���-�70.

Wolfe KH, M Gouy, Y-W Yang, PM Sharp & W-H Li (�989) Date of the monocot-dicot divergence estimated from chloroplast DNA sequence data. Proc. Natl. Acad. Sci. U.S.A., 86, ��0�-��0�.

Wood CE Jr (�97�) Morphology and Phytogeography: The Classical Approach to the Study of Disjunctions. Annals of the Missouri Botanical Garden, 59, �07-��4.

Wu, Z-Y, A-M Lu, Y-C Tang, Z-D Chen & W-H Li (�00�) Synopsis of a new “polyphyletic-polychromic-polytopic” system of the angiosperms. Acta Phytotaxonomica Sinica, 40, �89-���.

Wulff EV (�9�0) An introduction to historical plant geography. The Chronica Botanica Company. Waltham, Massachusets.

Yáñez GA, CR Ranero, R von Huene & J Díaz (�00�) Magnetic anomaly interpretation across the southern central Andes (��°-�4°S): The role of the Juan Fernández Ridge in the late Tertiary evolution of the margin. Journal of Geophysical Research-Solid Earth, 106, ����-��4�.

��0

Zachos J, M Pagani, L Sloan, E Thomas & K Billups (�00�) Trends, Rhythms, and Aberrations in Global Climate �� Ma to Present. Science, 292, �8�-�9�.

Zhou ZK, XF Yang & QS Yang (2006) Land bridge and long-distance dispersal - Old views, new evidence. Chinese Science Bulletin, 51, �0�0-�0�8.

Zimmermann W (�9�9) Die Phylogenie der Pflanzen (�d edition). Stuttgart.

Zizka G (�99�) Flowering Plants of Easter Island. Palmarum Hortus Francofurtensis, 3, �-�08.

Zuloaga FO, O Morrone & D Rodríguez (�999) Análisis de la biodiversidad en plantas vasculares de la Argentina. Kurtziana, 27, �7-��7.

Zunino M & A Zullini (�00�) Biogeografía: la dimensión espacial de la evolución. Fondo de Cultura Económica, México. Transl. from the Italian edition, �99�.

���

Appendix A: List of Chilean genera, geographic distributi-on, floristic elements

���

GEN

US

CLA

SE

AO

RD

EN

FAM

ILIA

N°

SP

DIS

TRIB

TRA

CK

ELE

MS

P C

HIL

EN

_MA

XS

_MA

XM

it-te

lWR

ange

Aa

Mon

ocot

sA

spar

agal

esO

rchi

dace

ae27

And

es C

olom

bia

to A

rgen

tina

AN

D3

1-1

7,58

-24,

50-2

1,04

6,92

Abr

otan

ella

Ast

erid

sA

ster

ales

Ast

erac

eae

20N

ew G

uine

a, A

us, T

asm

ania

, NZ,

S A

m (T

Fueg

o), J

F,

Falk

land

Is.

SP

T2

4-4

3,00

-55,

75-4

9,38

12,7

5

Aca

cia

Ros

ids

Faba

les

Faba

ceae

1200

Trop

ics,

Sub

trop,

esp

Afr

y A

us1

1-2

7,25

-37,

90-3

2,58

10,6

5

Aca

ena

Ros

ids

Ros

ales

Ros

acea

e10

0S

Afr

(1),

New

Gui

nea,

Aus

, Tas

man

ia, N

Z, s

uban

t-ar

ctic

Is, P

olyn

esia

, Haw

aii,

Cal

iforn

ia to

SA

, JF

419

-22,

00-5

6,00

-39,

0034

,00

Aca

ntho

lippi

aA

ster

ids

Lam

iale

sVe

rben

acea

e6

Arid

S A

m, A

rg, C

hile

, Bol

AN

D3

3-1

8,42

-30,

00-2

4,21

11,5

8

Ach

yroc

line

Ast

erid

sA

ster

ales

Ast

erac

eae

30Tr

opic

s A

fr, A

m, M

adag

asca

r, 1

1-1

8,23

-18,

23-1

8,23

0,00

Acr

isio

neA

ster

ids

Ast

eral

esA

ster

acea

e2

Cen

tral C

hile

62

-31,

62-4

5,50

-38,

5613

,88

Ade

noca

ulon

Ast

erid

sA

ster

ales

Ast

erac

eae

5W

NA

, Mex

ico,

Haw

aii,

Gua

tem

ala,

Chi

le, A

rg,

Him

alay

as, E

Asi

aW

-AN

T4

1-3

6,83

-55,

00-4

5,92

18,1

7

Ade

nope

ltis

Ros

ids

Mal

pigh

iale

sE

upho

rbia

ceae

1C

hile

61

-30,

50-3

6,83

-33,

676,

33

Ade

smia

Ros

ids

Faba

les

Faba

ceae

230

Per

u, C

hile

, Arg

, S B

rasi

l, TF

uego

S-A

MZ

313

2-1

8,20

-54,

83-3

6,52

36,6

3

Adi

antu

mFi

licop

sida

Pte

ridal

esP

terid

acea

e15

0C

osm

op, e

sp N

eotro

ps, M

adag

76

-21,

27-4

5,40

-33,

3324

,13

Aex

toxi

con

Eud

icot

sB

erbe

ridop

sida

les

Aex

toxi

cace

ae1

Chi

le, A

rg5

1-3

0,67

-43,

67-3

7,17

13,0

0

Aga

linis

Ast

erid

sLa

mia

les

Oro

banc

hace

ae40

Trop

, war

m C

& S

Am

, US

AA

NT-

P4

1-2

9,00

-30,

00-2

9,50

1,00

Age

ratin

aA

ster

ids

Ast

eral

esA

ster

acea

e29

0E

US

A, C

& W

S A

m, W

est I

ndie

sW

-NT

31

-25,

00-3

6,67

-30,

8311

,67

Ago

seris

Ast

erid

sA

ster

ales

Ast

erac

eae

119

sp W

N A

m, M

exic

o, 2

sp

Chi

le, A

rgA

NT-

P4

2-3

0,95

-56,

00-4

3,48

25,0

5

Agr

ostis

Mon

ocot

sP

oale

sP

oace

ae22

0te

mp

& w

arm

regi

ons,

mon

tane

trop

ics,

JF,

IPW

-AN

T4

24-3

1,00

-56,

00-4

3,50

25,0

0

Alc

hem

illa

Ros

ids

Ros

ales

Ros

acea

e25

0te

mp

N h

emis

, Mid

dle

Asi

a, m

onta

ne tr

opic

s, S

Afr

W-A

NT

43

-18,

20-1

8,23

-18,

220,

03

Alis

ma

Mon

ocot

sA

lism

atal

esA

lism

atac

eae

10te

mp

N H

emis

hp, E

uro,

Mid

dle

asia

, Aus

tralia

, S A

m7

1-3

3,43

-40,

00-3

6,72

6,57

Alli

onia

Eud

icot

sC

aryo

phyl

lale

sN

ycta

gina

ceae

3S

W &

C U

S, M

exic

o to

Chi

le &

Arg

AN

T-P

41

-20,

17-2

3,50

-21,

833,

33

Alo

naA

ster

ids

Sol

anal

esS

olan

acea

e6

Chi

le6

6-1

8,83

-32,

00-2

5,42

13,1

7

Alo

nsoa

Ast

erid

sLa

mia

les

Scr

ophu

laria

ceae

16Tr

op A

m M

exic

o, C

Am

, Chi

le, B

ol, 2

sp

S A

frW

-NT

31

-29,

50-3

6,67

-33,

087,

17

Alo

pecu

rus

Mon

ocot

sP

oale

sP

oace

ae36

war

m te

mp

& s

ubtro

p re

gion

s &

mon

tane

trop

ics

W-A

NT

44

-31,

50-5

3,67

-42,

5822

,17

Alo

ysia

Ast

erid

sLa

mia

les

Verb

enac

eae

30A

m: S

W U

S to

Chi

le &

Arg

W-N

T3

3-2

0,50

-30,

50-2

5,50

10,0

0

Als

troe

mer

iaM

onoc

ots

Lilia

les

Als

troem

eria

ceae

75C

hile

, S B

rasi

l, P

erú,

Arg

S-A

MZ

333

-20,

33-5

4,00

-37,

1733

,67

Alte

rnan

ther

aE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

100

Trop

ics

& s

ubtro

p, S

Am

, N A

m, A

us, 1

sp

Cau

casu

s1

2-1

8,50

-28,

50-2

3,50

10,0

0

Am

aran

thus

Eud

icot

sC

aryo

phyl

lale

sA

mar

anth

acea

e60

Sub

trop

& te

mp,

Eur

o, S

Am

, N A

m, 1

6 sp

Rus

sia,

7

2-3

3,23

-40,

35-3

6,79

7,12

Am

blyo

papp

usA

ster

ids

Ast

eral

esA

ster

acea

e1

1 sp

. Cal

if, N

W M

exic

o, C

hile

AN

T-P

41

-22,

08-3

0,65

-26,

378,

57

Am

bros

iaA

ster

ids

Ast

eral

esA

ster

acea

e43

Can

ada,

Mex

ico,

SA

, Wes

t Ind

ies

W-N

T3

2-1

7,58

-42,

72-3

0,15

25,1

3

Am

omyr

tus

Ros

ids

Myr

tale

sM

yrta

ceae

2Te

mp

S A

m: A

rg, C

hile

52

-35,

33-4

6,83

-41,

0811

,50

Am

phib

rom

usM

onoc

ots

Poa

les

Poa

ceae

12A

us, N

Z, S

Am

AU

S2

1-3

5,40

-43,

00-3

9,20

7,60

Am

phis

cirp

usM

onoc

ots

Poa

les

Cyp

erac

eae

1C

anad

a, U

SA

, Arg

, Chi

leA

NT-

P4

1-2

7,00

-55,

00-4

1,00

28,0

0

���

Am

sinc

kia

Ast

erid

sU

npla

ced

Bor

agin

acea

e15

W U

S, W

tem

p S

Am

AN

T-P

42

-22,

25-5

3,00

-37,

6330

,75

Ana

galli

sA

ster

ids

Eric

ales

Myr

sina

ceae

20E

uro,

Rus

sia,

N &

S A

fr, M

adag

, W A

sia,

Him

alay

as, N

A

us, 2

sp.

S A

m7

2-2

5,00

-55,

00-4

0,00

30,0

0

Ana

rthr

ophy

llum

Ros

ids

Faba

les

Faba

ceae

15A

ndes

Chi

le A

rg5

5-3

0,47

-52,

67-4

1,57

22,2

0

Ana

ther

ostip

aM

onoc

ots

Poa

les

Poa

ceae

11A

rg, B

ol, C

hile

, Ecu

ador

, Per

uA

ND

34

-19,

83-2

5,50

-22,

675,

67

And

eim

alva

Ros

ids

Mal

vale

sM

alva

ceae

4C

hile

, Per

u, B

olA

ND

31

-28,

58-3

6,90

-32,

748,

32

And

rosa

ceA

ster

ids

Eric

ales

Prim

ulac

eae

150

N H

emis

pher

e, C

hina

, few

sp

SA

(Tfu

ego)

W-A

NT

41

-53,

17-5

3,17

-53,

170,

00

Ane

mon

eE

udic

ots

Ran

uncu

lale

sR

anun

cula

ceae

90E

uras

, Sum

atra

, S &

E A

fr, N

Z, A

mul

tifida

dis

j N A

m

to C

hile

W-A

NT

46

-30,

83-5

3,15

-41,

9922

,32

Ani

som

eria

Eud

icot

sC

aryo

phyl

lale

sP

hyto

lacc

acea

e2

Chi

le6

2-2

4,83

-35,

97-3

0,40

11,1

3

Ant

enna

riaA

ster

ids

Ast

eral

esA

ster

acea

e50

Arc

tic &

tem

p E

uras

, And

es S

Am

W

-AN

T4

1-3

1,00

-53,

13-4

2,07

22,1

3

Ant

hoch

loa

Mon

ocot

sP

oale

sP

oace

ae1

And

es C

hile

Per

uA

ND

31

-18,

38-2

1,50

-19,

943,

12

Ant

hoxa

nthu

mM

onoc

ots

Poa

les

Poa

ceae

20te

mp

& w

arm

Eur

asia

, N A

fr, m

onta

ne tr

opic

s7

7-3

6,50

-56,

00-4

6,25

19,5

0

Ant

idap

hne

Eud

icot

sS

anta

lale

sS

anta

lace

ae8

W tr

op S

A: S

Mex

ico,

Gua

tem

, Cos

ta R

ica,

Col

om, V

e-ne

z, E

cuad

, Per

u, B

ol, B

ras,

Chi

le, C

uba,

Hai

ti, P

.Ric

oW

-NT

31

-37,

67-4

2,37

-40,

024,

70

Aph

anes

Ros

ids

Ros

ales

Ros

acea

e20

Am

, Eur

o, M

edit,

Eth

iop,

Aus

tralia

, Cau

casu

s7

3-3

3,40

-37,

17-3

5,28

3,77

Aph

yllo

clad

usA

ster

ids

Ast

eral

esA

ster

acea

e5

And

es S

Bol

ivia

, N C

hile

, NW

Arg

AN

D3

1-1

8,82

-27,

78-2

3,30

8,97

Api

umA

ster

ids

Api

ales

Api

acea

e27

Eur

o, M

edit,

N &

S A

fr, M

adag

, W &

E A

sia,

Mal

esia

, A

us, N

, C &

S A

m, J

F, IP

78

-19,

80-5

4,93

-37,

3735

,13

Apo

dasm

iaM

onoc

ots

Poa

les

Res

tiona

ceae

10S

E A

sia,

Mal

esia

, New

Gui

nea,

Aus

, Tas

m, N

Z, C

hile

, A

rgS

PT

21

-36,

83-5

1,00

-43,

9214

,17

Ara

chni

tisM

onoc

ots

Lilia

les

Cor

siac

eae

1C

hile

, Arg

51

-33,

00-5

0,00

-41,

5017

,00

Ara

eoan

dra

Ros

ids

Ger

ania

les

Vivi

ania

ceae

1C

hile

61

-30,

17-3

0,83

-30,

500,

67

Ara

ucar

iaP

inop

sida

Pin

ales

Ara

ucar

iace

ae18

New

Gui

nea,

Aus

, New

Cal

edon

ia, S

Chi

le, S

E B

rasi

lS

PT

21

-37,

50-4

0,05

-38,

782,

55

Are

naria

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae16

0A

mer

ica,

tem

p E

uras

ia, N

Afr

W-A

NT

47

-18,

00-5

5,00

-36,

5037

,00

Arg

emon

eE

udic

ots

Ran

uncu

lale

sP

apav

erac

eae

28N

& S

AM

, WI,

Haw

aii

W-N

T3

4-1

8,48

-35,

35-2

6,92

16,8

7

Arg

ylia

Ast

erid

sLa

mia

les

Big

noni

acea

e12

S P

eru,

N C

Chi

le &

Arg

, A. u

spal

late

nsis

S B

olA

ND

311

-20,

82-3

4,47

-27,

6413

,65

Aris

tegu

ietia

Ast

erid

sA

ster

ales

Ast

erac

eae

20C

olom

bia,

Ecu

ador

, Per

u, C

hile

AN

D3

1-2

9,50

-36,

45-3

2,98

6,95

Aris

tida

Mon

ocot

sP

oale

sP

oace

ae29

0N

Am

, Med

it, te

mp

Asi

a, tr

opic

s &

sub

tropi

cs7

3-1

8,83

-37,

83-2

8,33

19,0

0

Aris

tolo

chia

Mag

nolii

dsP

iper

ales

Aris

tolo

chia

ceae

300

trop,

sub

trop,

som

e in

tem

p re

gion

s1

2-2

7,50

-33,

48-3

0,49

5,98

Aris

tote

liaR

osid

sO

xalid

ales

Ela

eoca

rpac

eae

5E

Aus

, Tas

man

, NZ,

Arg

, Chi

leA

US

21

-30,

83-4

4,00

-37,

4213

,17

Arjo

naE

udic

ots

San

tala

les

San

tala

ceae

10Te

mp

S A

m, B

razi

lS

-AM

Z3

3-3

1,00

-54,

50-4

2,75

23,5

0

Arm

eria

Eud

icot

sC

aryo

phyl

lale

sP

lum

bagi

nace

ae90

SA

m A

ndes

to T

fueg

o, E

uro,

Rus

sia

W-A

NT

41

-30,

67-5

6,00

-43,

3325

,33

Art

emis

iaA

ster

ids

Ast

eral

esA

ster

acea

e39

0te

mp

Eur

asia

& W

N A

m, N

+ S

Afr,

S A

mW

-AN

T4

2-1

7,75

-53,

00-3

5,38

35,2

5

Art

hrop

teris

Filic

opsi

daD

aval

liale

sO

lean

drac

eae

15N

ew G

uine

a, N

Z, J

uan

Fern

ande

z, M

adag

SP

T2

10,

00

Asp

leni

umFi

licop

sida

Asp

leni

ales

Asp

leni

acea

e65

0S

ubco

smop

, Mad

ag, J

F7

7-1

8,00

-55,

92-3

6,96

37,9

2

Ast

elia

Mon

ocot

sA

spar

agal

esA

stel

iace

ae25

Mas

c, N

ew G

uine

a, A

us, N

Z, P

olin

esia

, Chi

leS

PT

21

-40,

17-5

6,00

-48,

0815

,83

Ast

erA

ster

ids

Ast

eral

esA

ster

acea

e25

0te

mp

Eur

asia

, S A

fr, M

adag

, Am

, Haw

aii

W-A

NT

44

-30,

65-5

6,00

-43,

3325

,35

��4

Ast

eran

ther

aA

ster

ids

Lam

iale

sG

esne

riace

ae1

Chi

le, A

rg5

1-3

5,60

-47,

00-4

1,30

11,4

0

Ast

eris

cium

Ast

erid

sA

pial

esA

piac

eae

8M

exic

o to

Pat

agon

iaW

-NT

34

-25,

20-3

7,83

-31,

5212

,63

Ast

raga

lus

Ros

ids

Faba

les

Faba

ceae

2000

N A

m, A

ndes

, tem

p E

uras

ia, m

onta

ne tr

op A

frW

-AN

T4

46-1

7,58

-53,

50-3

5,54

35,9

2

Atr

iple

xE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

250

cosm

op, E

uras

ia, A

us, I

D7

20-1

8,00

-53,

33-3

5,67

35,3

3

Aus

troc

actu

sE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae5

Arg

, Chi

le5

2-3

3,78

-46,

50-4

0,14

12,7

2

Aus

troc

edru

sP

inop

sida

Pin

ales

Cup

ress

acea

e1

Arg

, Chi

le5

1-3

2,65

-40,

22-3

6,43

7,57

Avel

lani

taR

osid

sM

alpi

ghia

les

Eup

horb

iace

ae1

Chi

le6

1-3

3,83

-34,

10-3

3,97

0,27

Axo

nopu

sM

onoc

ots

Poa

les

Poa

ceae

110

pant

rops

, IP

11

0,00

Aza

raR

osid

sM

alpi

ghia

les

Sal

icac

eae

108

tem

p C

hile

-Arg

, s s

ubtro

p N

W A

rg, B

ol, S

E B

ra,

Uru

guay

, JF

S-A

MZ

38

-30,

00-4

6,00

-38,

0016

,00

Azo

llaFi

licop

sida

Sal

vini

ales

Azo

llace

ae6

trop

& w

arm

, Eur

o, A

sia,

Mad

ag, T

rista

n da

Cun

ha,

Haw

aii

71

-18,

18-4

2,55

-30,

3724

,37

Azo

rella

Ast

erid

sA

pial

esA

piac

eae

70A

ndes

to te

mp

S A

m, M

alvi

nas,

Ant

arct

ic Is

land

sA

US

214

-18,

00-5

5,50

-36,

7537

,50

Bac

char

isA

ster

ids

Ast

eral

esA

ster

acea

e40

0A

M, e

sp S

AM

W-N

T3

43-1

7,58

-56,

00-3

6,79

38,4

2

Bac

opa

Ast

erid

sLa

mia

les

Gra

tiola

ceae

60Tr

op &

Sub

trop,

Haw

aii,

Mad

ag, A

us1

1-2

0,00

-30,

50-2

5,25

10,5

0

Bah

iaA

ster

ids

Ast

eral

esA

ster

acea

e10

SW

US

, Mex

, Chi

leA

NT-

P4

1-2

1,27

-36,

62-2

8,94

15,3

5

Bak

erol

imon

Eud

icot

sC

aryo

phyl

lale

sP

lum

bagi

nace

ae1

Chi

le6

1-2

3,50

-30,

00-2

6,75

6,50

Bal

bisi

aR

osid

sG

eran

iale

sLe

doca

rpac

eae

8P

eru,

Bol

, Chi

le, A

rgA

ND

33

-18,

00-3

0,00

-24,

0012

,00

Bal

sam

ocar

pon

Ros

ids

Faba

les

Faba

ceae

1C

hile

61

-27,

37-2

9,83

-28,

602,

47

Bar

neou

dia

Eud

icot

sR

anun

cula

les

Ran

uncu

lace

ae3

Arg

, Chi

leA

ND

33

-29,

82-3

4,25

-32,

034,

43

Bar

tsia

Ast

erid

sLa

mia

les

Oro

banc

hace

ae60

Circ

umbo

real

, Eur

, Med

it, A

fr, A

ndes

W-A

NT

43

-18,

00-3

6,50

-27,

2518

,50

Bei

lsch

mie

dia

Mag

nolii

dsLa

ural

esLa

urac

eae

230

pant

ropi

cs, M

ex to

S B

ras,

Chi

le, A

us, N

Z, A

fr, M

adag

12

-32,

55-3

6,67

-34,

614,

12

Bel

loa

Ast

erid

sA

ster

ales

Ast

erac

eae

1A

ndes

Arg

, Chi

le5

1-3

4,83

-40,

75-3

7,79

5,92

Ben

tham

iella

Ast

erid

sS

olan

ales

Sol

anac

eae

12S

Chi

le-A

rg, P

atag

onia

54

-46,

58-5

3,67

-50,

137,

08

Ber

berid

opsi

sE

udic

ots

Ber

berid

opsi

-da

les

Ber

berid

opsi

-da

ceae

2B

. bec

kler

i en

Aus

: Que

ensl

and

& N

ew S

Wal

es, B

. co

ralli

na: C

hile

AU

S2

1-3

7,13

-37,

67-3

7,40

0,53

Ber

beris

Eud

icot

sR

anun

cula

les

Ber

berid

acea

e47

0E

uras

, N A

fr, N

& S

AM

W-A

NT

447

-25,

02-5

5,97

-40,

4930

,95

Bid

ens

Ast

erid

sA

ster

ales

Ast

erac

eae

240

cosm

op7

7-1

7,58

-37,

17-2

7,38

19,5

8

Bip

innu

laM

onoc

ots

Asp

arag

ales

Orc

hida

ceae

8Te

mp

S A

m: C

hile

, Arg

54

-25,

00-4

0,50

-32,

7515

,50

Ble

chnu

mFi

licop

sida

Ble

chna

les

Ble

chna

ceae

175

Sub

cosm

op, M

adag

, JF

710

-30,

67-5

5,97

-43,

3225

,30

Ble

nnos

perm

aA

ster

ids

Ast

eral

esA

ster

acea

e3

2 C

alifo

rnia

, 1 C

hile

AN

T-P

41

-33,

42-3

6,60

-35,

013,

18

Ble

phar

ocal

yxR

osid

sM

yrta

les

Myr

tace

ae17

trop

& s

ubtro

S A

m, W

IA

ND

31

-32,

50-4

1,30

-36,

908,

80

Blu

men

bach

iaA

ster

ids

Cor

nale

sLo

asac

eae

12B

rasi

l, U

rugu

ay, P

arag

uay,

Arg

entin

a, C

hile

S-A

MZ

33

-33,

75-4

5,50

-39,

6311

,75

Boe

hmer

iaR

osid

sR

osal

esU

rtica

ceae

50P

antro

p, J

F1

10,

00

Boe

rhav

iaE

udic

ots

Car

yoph

ylla

les

Nyc

tagi

nace

ae35

tropi

cs &

sub

tropi

cs1

2-2

0,50

-37,

00-2

8,75

16,5

0

Bol

axA

ster

ids

Api

ales

Api

acea

e5

Tem

p S

A5

2-4

9,15

-55,

52-5

2,33

6,37

Bom

area

Mon

ocot

sLi

liale

sA

lstro

emer

iace

ae10

0M

ex to

trop

Am

W-N

T3

3-1

8,00

-39,

00-2

8,50

21,0

0

Boo

pis

Ast

erid

sA

ster

ales

Cal

ycer

acea

e13

And

es, A

rg, S

Bra

sil

S-A

MZ

38

-36,

00-5

4,00

-45,

0018

,00

Boq

uila

Eud

icot

sR

anun

cula

les

Lard

izab

alac

eae

1A

rg, C

hile

51

-35,

83-4

2,70

-39,

276,

87

Bot

rych

ium

Filic

opsi

daO

phio

glos

sale

sO

phig

loss

acea

e35

tem

p, p

olar

, tro

p m

onta

neW

-AN

T4

1-5

0,50

-55,

67-5

3,08

5,17

���

Bou

telo

uaM

onoc

ots

Poa

les

Poa

ceae

40A

m fr

om C

anad

a to

Arg

entin

aW

-NT

31

-18,

37-2

4,00

-21,

185,

63

Bow

lesi

aA

ster

ids

Api

ales

Api

acea

e15

N &

S A

m, B

. inc

ana

S U

SW

-NT

36

-18,

17-5

2,00

-35,

0833

,83

Bra

chyc

lado

sA

ster

ids

Ast

eral

esA

ster

acea

e3

Tem

p S

A, C

hile

-Arg

51

-30,

83-3

0,95

-30,

890,

12

Bra

chys

tele

Mon

ocot

sA

spar

agal

esO

rchi

dace

ae22

trop

Am

esp

. Bra

sil

S-A

MZ

31

-30,

67-4

3,62

-37,

1412

,95

Brid

gesi

aR

osid

sS

apin

dale

sS

apin

dace

ae1

Chi

le6

1-2

8,57

-32,

88-3

0,73

4,32

Bro

mid

ium

Mon

ocot

sP

oale

sP

oace

ae5

S U

S &

sub

trop

S A

mA

NT-

P4

2-3

1,67

-40,

00-3

5,83

8,33

Bro

mus

Mon

ocot

sP

oale

sP

oace

ae15

0E

uras

ia, N

& S

Afr,

mon

tane

trop

Asi

a, A

m, J

FW

-AN

T4

11-1

8,20

-55,

00-3

6,60

36,8

0

Bro

wni

ngia

Eud

icot

sC

aryo

phyl

lale

sC

acta

ceae

11P

eru,

Bol

, Chi

leA

ND

31

-18,

67-1

9,17

-18,

920,

50

Bud

dlej

aA

ster

ids

Lam

iale

sS

crop

hula

riace

ae90

E A

sia,

tro

& s

ubtro

p A

M, A

fr1

3-2

7,28

-42,

10-3

4,69

14,8

2

Bul

bost

ylis

Mon

ocot

sP

oale

sC

yper

acea

e90

tropi

cs &

sub

tropi

cs1

1-3

7,33

-39,

67-3

8,50

2,33

Bul

nesi

aR

osid

sZy

goph

ylla

les

Zygo

phyl

lace

ae8

S A

m: V

enez

uela

Col

ombi

a di

sjun

ct to

Chi

le-A

rg, B

razi

lA

ND

31

-27,

33-3

0,60

-28,

973,

27

Cae

salp

inia

Ros

ids

Faba

les

Faba

ceae

150

Pan

tropi

cs, s

ubtro

pics

, Arg

, Mad

ag, S

Afr,

Ara

bia

12

-19,

47-3

0,67

-25,

0711

,20

Cai

opho

raA

ster

ids

Cor

nale

sLo

asac

eae

56P

eru,

Bol

, Arg

, Chi

le, 1

sp

Uru

g, B

ras,

Ecu

adS

-AM

Z3

13-1

7,58

-38,

40-2

7,99

20,8

2

Cal

amag

rost

isM

onoc

ots

Poa

les

Poa

ceae

230

tem

p &

sub

trop

reg

& m

onta

ne tr

opic

s1

24-1

8,33

-55,

00-3

6,67

36,6

7

Cal

andr

inia

Eud

icot

sC

aryo

phyl

lale

sP

ortu

laca

ceae

13A

m W

US

A to

Chi

leW

-NT

311

-18,

00-5

4,50

-36,

2536

,50

Cal

ceol

aria

Ast

erid

sLa

mia

les

Cal

ceol

aria

ceae

300

AM

from

Mex

to T

fueg

oW

-NT

350

-18,

17-5

5,00

-36,

5836

,83

Cal

dclu

via

Ros

ids

Oxa

lidal

esC

unon

iace

ae11

Phi

lippi

nes,

Sul

awes

i, M

oluc

cas,

New

Gui

nea,

trp

Aus

, S

Chi

le A

rgS

PT

21

-35,

33-4

9,42

-42,

3814

,08

Cal

liand

raR

osid

sFa

bale

sFa

bace

ae20

0C

& S

Am

, Afr,

Mad

ag, t

rop

Asi

a1

1-2

9,00

-30,

50-2

9,75

1,50

Cal

litric

heA

ster

ids

Lam

iale

sP

lant

agin

acea

e17

Cos

mop

75

-25,

00-5

6,00

-40,

5031

,00

Cal

opap

pus

Ast

erid

sA

ster

ales

Ast

erac

eae

1C

hile

61

-32,

50-3

5,50

-34,

003,

00

Cal

othe

caM

onoc

ots

Poa

les

Poa

ceae

1A

rg, B

ras,

Chi

le, U

rugu

ayS

-AM

Z3

1-3

4,40

-37,

83-3

6,12

3,43

Cal

tha

Eud

icot

sR

anun

cula

les

Ran

uncu

lace

ae10

Arc

tic &

tem

p N

Hem

is, N

Z, te

mp

S A

MW

-AN

T4

3-2

9,88

-55,

97-4

2,93

26,0

8

Cal

ycer

aA

ster

ids

Ast

eral

esC

alyc

erac

eae

17Te

mp

S A

m, B

razi

lS

-AM

Z3

11-2

2,00

-40,

00-3

1,00

18,0

0

Cal

ydor

eaM

onoc

ots

Asp

arag

ales

Irida

ceae

15Tr

op &

Sub

trop

Am

W-N

T3

1-3

1,00

-36,

67-3

3,83

5,67

Cal

yste

gia

Ast

erid

sS

olan

ales

Con

volv

ulac

eae

25C

osm

op, t

emp

& tr

op h

emis

N &

S, J

F7

2-3

2,88

-40,

00-3

6,44

7,12

Cam

isso

nia

Ros

ids

Myr

tale

sO

nagr

acea

e62

W N

Am

, Mex

, S A

m te

mp

AN

T-P

41

-27,

37-4

3,17

-35,

2715

,80

Cam

psid

ium

Ast

erid

sLa

mia

les

Big

noni

acea

e1

Arg

, Chi

le: C

. val

divi

anum

51

-35,

83-5

2,25

-44,

0416

,42

Car

dam

ine

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e20

0C

osm

op, t

emp

N h

emis

& m

onta

ne tr

opic

s, J

FW

-AN

T4

20-3

0,00

-56,

00-4

3,00

26,0

0

Car

dion

ema

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae6

W N

Am

to C

hile

AN

T-P

43

-18,

00-3

7,83

-27,

9219

,83

Car

exM

onoc

ots

Poa

les

Cyp

erac

eae

2000

cosm

op, J

F7

55-2

1,00

-56,

00-3

8,50

35,0

0

Car

pha

Mon

ocot

sP

oale

sC

yper

acea

e15

trop

& S

Afr,

Mad

ag, j

apan

, New

Gui

nea,

Aus

, Tas

m, N

Z,

tem

p S

Am

11

-40,

50-5

6,00

-48,

2515

,50

Car

pobr

otus

Eud

icot

sC

aryo

phyl

lale

sA

izoa

ceae

30S

Afr,

Aus

, Tas

man

, NZ,

Cal

if, C

hile

41

-23,

50-4

0,00

-31,

7516

,50

Cas

tille

jaA

ster

ids

Lam

iale

sO

roba

ncha

ceae

200

Eur

as, E

N A

m, C

& S

Am

W-A

NT

44

-31,

50-3

9,27

-35,

387,

77

Cat

abro

saM

onoc

ots

Poa

les

Poa

ceae

4te

mp

Eur

asia

, N A

m, C

hile

W-A

NT

42

-18,

33-5

5,63

-36,

9837

,30

Cen

taur

eaA

ster

ids

Ast

eral

esA

ster

acea

e50

0E

uras

ia, N

Chi

na, N

& tr

o A

fr, A

us, N

Z, N

Am

77

-22,

00-3

7,80

-29,

9015

,80

���

Cen

taur

ium

Ast

erid

sG

entia

nale

sG

entia

nace

ae60

Cos

mop

exc

trop

& S

Afr,

N h

emis

p 1

ext.

to A

us, 1

to

Chi

le7

1-2

5,00

-41,

50-3

3,25

16,5

0

Cen

taur

oden

dron

Ast

erid

sA

ster

ales

Ast

erac

eae

2JF

62

0,00

Cen

tella

Ast

erid

sA

pial

esA

piac

eae

40P

antro

p to

Chi

le, N

Z, J

F1

1-3

5,12

-43,

00-3

9,06

7,88

Cen

tiped

aA

ster

ids

Ast

eral

esA

ster

acea

e6

Mad

ag, M

asca

rene

Is, A

sia,

Aus

, NZ,

New

Cal

e, T

ahiti

, C

hile

11

-30,

00-4

1,83

-35,

9211

,83

Cer

astiu

mE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

100

cosm

op7

2-2

3,00

-33,

33-2

8,17

10,3

3

Cer

atop

hyllu

mC

erat

ophy

llale

sC

erat

ophy

llace

ae20

cosm

op7

1-3

3,42

-36,

82-3

5,12

3,40

Ces

trum

Ast

erid

sS

olan

ales

Sol

anac

eae

175

trop

Am

, esp

Bra

sil &

And

esW

-NT

31

-18,

75-4

0,33

-29,

5421

,58

Cha

etan

ther

aA

ster

ids

Ast

eral

esA

ster

acea

e44

S P

eru,

Bol

, Chi

le, A

rgA

ND

337

-17,

58-4

0,25

-28,

9222

,67

Cha

ptal

iaA

ster

ids

Ast

eral

esA

ster

acea

e55

trop

& s

ubtro

p A

m, S

US

AW

-NT

31

-33,

03-3

6,67

-34,

853,

63

Cha

scol

ytru

mM

onoc

ots

Poa

les

Poa

ceae

6C

entra

l & S

Am

, US

AA

NT-

P4

1-3

2,13

-43,

83-3

7,98

11,7

0

Che

ilant

hes

Filic

opsi

daP

terid

ales

Pte

ridac

eae

150

Cos

mop

, esp

Neo

trops

& S

Afr,

Mad

ag7

9-1

8,00

-46,

55-3

2,28

28,5

5

Che

nopo

dium

Eud

icot

sC

aryo

phyl

lale

sA

mar

anth

acea

e12

5co

smop

, JF,

ID7

12-1

8,00

-54,

00-3

6,00

36,0

0

Che

rsod

oma

Ast

erid

sA

ster

ales

Ast

erac

eae

9S

Per

u, B

ol, C

hile

, Arg

AN

D3

3-1

7,72

-23,

58-2

0,65

5,87

Che

vreu

liaA

ster

ids

Ast

eral

esA

ster

acea

e5

S B

rasi

l, B

ol, P

arag

uay,

Uru

g, C

hile

, N C

Arg

, Fal

klan

dS

-AM

Z3

3-3

3,03

-43,

00-3

8,02

9,97

Chi

lioph

yllu

mA

ster

ids

Ast

eral

esA

ster

acea

e3

S A

ndes

Chi

le, A

rgen

tina

52

-52,

62-5

3,25

-52,

930,

63

Chi

liotr

ichu

mA

ster

ids

Ast

eral

esA

ster

acea

e7

S A

ndes

Chi

le, A

rgen

tina

52

-36,

00-5

6,00

-46,

0020

,00

Chi

rope

talu

mR

osid

sM

alpi

ghia

les

Eup

horb

iace

ae20

Mex

ico

2, P

eru,

S B

ras,

Arg

, Chi

leA

NT-

P4

4-2

2,50

-38,

00-3

0,25

15,5

0

Chl

orae

aM

onoc

ots

Asp

arag

ales

Orc

hida

ceae

50A

ndes

Chi

le P

erú,

Arg

, Bra

zil

S-A

MZ

329

-30,

50-5

5,33

-42,

9224

,83

Chl

oris

Mon

ocot

sP

oale

sP

oace

ae55

Cos

mop

71

-18,

47-1

8,47

-18,

470,

00

Cho

rizan

the

Eud

icot

sC

aryo

phyl

lale

sP

olyg

onac

eae

50ar

id &

sem

iarid

W A

m, C

hile

AN

T-P

424

-18,

00-3

7,22

-27,

6119

,22

Chr

ysos

plen

ium

Eud

icot

sS

axifr

agal

esS

axifr

agac

eae

64N

& te

mp

S A

m, E

uro,

N A

fr, A

sia,

W

-AN

T4

2-5

2,00

-56,

00-5

4,00

4,00

Chu

quira

gaA

ster

ids

Ast

eral

esA

ster

acea

e23

And

es S

Am

, Pat

agon

iaA

ND

35

-17,

72-3

5,75

-26,

7318

,03

Chu

sque

aM

onoc

ots

Poa

les

Poa

ceae

110

C &

S A

m, W

I, JF

W-N

T3

13-3

0,67

-46,

67-3

8,67

16,0

0

Cic

endi

aA

ster

ids

Gen

tiana

les

Gen

tiana

ceae

21

Eur

o &

Med

it, 1

Cal

if, W

S A

mW

-AN

T4

1-2

5,00

-38,

00-3

1,50

13,0

0

Cic

losp

erm

umA

ster

ids

Api

ales

Api

acea

e3

Arg

, Bol

, Bra

sil,

Chi

le, P

arag

uay,

Uru

guay

S-A

MZ

31

-20,

82-4

0,00

-30,

4119

,18

Cis

saro

bryo

nR

osid

sG

eran

iale

sVi

vian

iace

ae1

Chi

le6

1-3

3,00

-37,

80-3

5,40

4,80

Cis

sus

Eud

icot

sVi

tale

sVi

tace

ae20

0tro

pics

incl

Aus

tralia

11

-30,

00-4

3,00

-36,

5013

,00

Cis

tant

heE

udic

ots

Car

yoph

ylla

les

Por

tula

cace

ae49

N A

m, C

alif,

Ariz

ona,

N M

éxic

o &

S A

m, P

eru,

Bol

, Arg

, C

hile

AN

T-P

418

-18,

58-3

7,93

-28,

2619

,35

Citr

onel

laA

ster

ids

Unp

lace

dIc

acin

acea

e20

Mal

esia

, E A

us, N

ew C

ale,

Fiji

, Sam

oa, C

& tr

op S

Am

SP

T2

1-3

0,67

-38,

75-3

4,71

8,08

Cla

rkia

Ros

ids

Myr

tale

sO

nagr

acea

e42

N A

m s

pec.

Cal

iforn

ia, 1

sp

Chi

le/A

rgA

NT-

P4

1-2

9,67

-40,

33-3

5,00

10,6

7

Cle

ome

Ros

ids

Bra

ssic

ales

Cle

omac

eae

275

tropi

cs &

sub

tropi

cs, s

pec.

Am

11

-20,

22-2

6,33

-23,

286,

12

Clio

cocc

aR

osid

sM

alpi

ghia

les

Lina

ceae

1te

mp

S A

m5

1-3

4,88

-40,

32-3

7,60

5,43

Cod

onor

chis

Mon

ocot

sA

spar

agal

esO

rchi

dace

ae3

S tr

op &

tem

p S

Am

, Bra

zil

S-A

MZ

31

-35,

00-5

6,00

-45,

5021

,00

Col

letia

Ros

ids

Ros

ales

Rha

mna

ceae

5su

btro

p &

tem

p S

Am

, Bra

zil,

JFS

-AM

Z3

2-3

0,25

-46,

50-3

8,38

16,2

5

Col

ligua

jaR

osid

sM

alpi

ghia

les

Eup

horb

iace

ae5

tem

p S

Am

, S B

rasi

lS

-AM

Z3

4-2

3,50

-46,

67-3

5,08

23,1

7

Col

lom

iaA

ster

ids

Eric

ales

Pol

emon

iace

ae15

W N

Am

, Bol

to P

atag

onia

AN

T-P

42

-32,

90-5

3,00

-42,

9520

,10

��7

Col

oban

thus

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae20

S P

acifi

c, A

us, N

Z, te

mp

S A

m, E

cuad

orS

PT

23

-17,

58-5

6,00

-36,

7938

,42

Com

bera

Ast

erid

sS

olan

ales

Sol

anac

eae

2A

ndes

Chi

le A

rg5

2-3

7,00

-44,

00-4

0,50

7,00

Con

anth

era

Mon

ocot

sA

spar

agal

esTe

coph

ilaea

ceae

5C

hile

65

-22,

08-3

7,95

-30,

0215

,87

Con

volv

ulus

Ast

erid

sS

olan

ales

Con

volv

ulac

eae

150

cosm

op e

sp. t

emp,

trop

& s

ubtro

ps7

5-2

5,00

-40,

00-3

2,50

15,0

0

Con

yza

Ast

erid

sA

ster

ales

Ast

erac

eae

60tro

pics

& s

ubtro

pics

127

-17,

58-5

5,00

-36,

2937

,42

Cop

iapo

aE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae25

N C

hile

625

-22,

00-3

1,00

-26,

509,

00

Cop

rosm

aA

ster

ids

Gen

tiana

les

Rub

iace

ae90

E M

ales

ia, A

us, N

Z, P

acifi

c, J

F, 1

2 sp

Haw

aii

SP

T2

20,

00

Cor

dia

Ast

erid

sU

npla

ced

Bor

agin

acea

e32

0tro

p &

sub

tropi

cs1

1-2

7,82

-31,

83-2

9,83

4,02

Cor

eops

isA

ster

ids

Ast

eral

esA

ster

acea

e11

4N

& S

Am

W-N

T3

1-1

8,17

-18,

17-1

8,17

0,00

Cor

iaria

Ros

ids

Cuc

urbi

tale

sC

oria

riace

ae15

Mex

to C

hile

, W M

edit,

Him

al to

Jap

an, N

ew G

uine

a,

NZ.

C ru

scifo

lia d

isj N

Z7

1-3

5,02

-43,

83-3

9,43

8,82

Cor

rigio

laE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

13S

Am

And

es C

olom

to C

hile

, Eur

o, M

edit,

Afr,

Mad

ag7

3-3

0,00

-37,

00-3

3,50

7,00

Cor

ryoc

actu

sE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae12

Bol

, S P

eru,

N C

hile

AN

D3

1-1

9,75

-20,

00-1

9,88

0,25

Cor

tade

riaM

onoc

ots

Poa

les

Poa

ceae

25S

Am

, 4 N

Z, 1

New

Gui

nea

AU

S2

6-1

8,47

-56,

00-3

7,23

37,5

3

Cor

ynab

utilo

nR

osid

sM

alva

les

Mal

vace

ae7

Chi

le, A

rg5

7-3

0,67

-42,

53-3

6,60

11,8

7

Cot

ula

Ast

erid

sA

ster

ales

Ast

erac

eae

55S

Afr,

NZ,

New

Gui

nea,

Aus

, S A

m1

1-2

0,25

-22,

00-2

1,13

1,75

Cra

ssul

aE

udic

ots

Sax

ifrag

ales

Cra

ssul

acea

e20

0co

smop

, Eur

o, A

& S

Afr,

Mad

ag, A

us, A

m7

7-1

8,00

-56,

00-3

7,00

38,0

0

Cre

mol

obus

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e7

And

es C

olom

bia

to C

hile

AN

D3

1-1

7,58

-18,

50-1

8,04

0,92

Cre

ssa

Ast

erid

sS

olan

ales

Con

volv

ulac

eae

5tro

p &

sub

trop

11

-20,

45-3

5,33

-27,

8914

,88

Crin

oden

dron

Ros

ids

Oxa

lidal

esE

laeo

carp

acea

e5

Bol

, N A

rg, C

Chi

le, B

rasi

lS

-AM

Z3

2-3

2,23

-42,

77-3

7,50

10,5

3

Cris

taria

Ros

ids

Mal

vale

sM

alva

ceae

75C

hile

, Arg

, Bol

, Per

ú, ID

AN

D3

19-1

8,17

-35,

63-2

6,90

17,4

7

Cro

ton

Ros

ids

Mal

pigh

iale

sE

upho

rbia

ceae

1250

tropi

cs &

sub

tropi

cs1

1-2

4,90

-25,

33-2

5,12

0,43

Cru

cksh

anks

iaA

ster

ids

Gen

tiana

les

Rub

iace

ae7

Chi

le, A

rgA

ND

37

-22,

08-3

3,83

-27,

9611

,75

Cry

ptan

tha

Ast

erid

sU

npla

ced

Bor

agin

acea

e10

0W

US

, W te

mp

S A

mA

NT-

P4

39-1

8,83

-44,

50-3

1,67

25,6

7

Cry

ptoc

arya

Mag

nolii

dsLa

ural

esLa

urac

eae

230

tropi

cs &

sub

tropi

cs, M

adag

, S A

fr, S

Indi

a, C

eylo

n,

Mal

aysi

a, P

hilip

pine

s, N

Aus

tralia

11

-30,

33-4

0,25

-35,

299,

92

Cry

ptog

ram

ma

Filic

opsi

daP

terid

ales

Pte

ridac

eae

2te

mpe

rate

N &

SW

-AN

T4

1-3

1,30

-37,

40-3

4,35

6,10

Cua

trec

asas

iella

Ast

erid

sA

ster

ales

Ast

erac

eae

2E

cuad

or, P

eru,

Chi

le, A

rgA

ND

31

-17,

72-2

4,50

-21,

116,

78

Cum

inia

Ast

erid

sLa

mia

les

Lam

iace

ae1

JF6

10,

00

Cum

ulop

untia

Eud

icot

sC

aryo

phyl

lale

sC

acta

ceae

20B

ol, S

Per

u, N

Chi

leA

ND

32

-18,

00-3

3,00

-25,

5015

,00

Cus

cuta

Ast

erid

sS

olan

ales

Con

volv

ulac

eae

155

cosm

op7

10-1

9,25

-40,

35-2

9,80

21,1

0

Cyn

odon

Mon

ocot

sP

oale

sP

oace

ae9

tropi

cs &

sub

tropi

cs1

5-1

8,47

-33,

43-2

5,95

14,9

7

Cyn

oglo

ssum

Ast

erid

sU

npla

ced

Bor

agin

acea

e75

tem

p &

sub

trop

reg

71

-32,

95-3

9,98

-36,

477,

03

Cyp

erus

Mon

ocot

sP

oale

sC

yper

acea

e40

0tro

p, s

ubtro

p, te

mp

regi

ons,

JF

726

-20,

00-4

3,00

-31,

5023

,00

Cyp

hoca

rpus

Ast

erid

sA

ster

ales

Cam

panu

lace

ae3

Chi

le6

3-2

7,18

-32,

35-2

9,77

5,17

Cys

topt

eris

Filic

opsi

daD

ryop

terid

ales

Dry

opte

ridac

eae

12te

mp

& w

arm

regi

ons,

mon

tane

trop

ics

W-A

NT

41

-18,

00-5

5,00

-36,

5037

,00

Dal

eaR

osid

sFa

bale

sFa

bace

ae16

5A

m, C

anad

a to

Arg

W-N

T3

3-1

8,00

-25,

40-2

1,70

7,40

��8

Dan

thon

iaM

onoc

ots

Poa

les

Poa

ceae

20N

& S

Am

, Eur

o, C

auca

sus

W-A

NT

48

-32,

00-4

2,62

-37,

3110

,62

Das

yphy

llum

Ast

erid

sA

ster

ales

Ast

erac

eae

40C

hile

, And

es, S

E B

razi

lS

-AM

Z3

2-3

2,88

-42,

47-3

7,68

9,58

Dau

cus

Ast

erid

sA

pial

esA

piac

eae

22E

uro,

Med

it, S

W &

S A

sia,

trop

Afr.

Aus

, NZ,

& A

m7

2-2

5,38

-46,

30-3

5,84

20,9

2

Dav

allia

Filic

opsi

daD

aval

liale

sD

aval

liace

ae90

W M

edit,

Him

al, J

apan

, Aus

, Tah

iti, A

fr, M

adag

, IP

11

0,00

Den

dros

eris

Ast

erid

sA

ster

ales

Ast

erac

eae

11JF

611

0,00

Den

nsta

edtia

Filic

opsi

daD

enns

taed

tiale

sD

enns

taed

tiace

ae45

trop

to w

arm

tem

p, M

adag

11

-32,

45-3

5,85

-34,

153,

40

Des

cham

psia

Mon

ocot

sP

oale

sP

oace

ae35

tem

p &

coo

l, es

pec

N E

uro,

mon

tane

trop

ics

W-A

NT

416

-28,

80-5

6,00

-42,

4027

,20

Des

cura

inia

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e40

S A

m, N

Am

, Can

ary

Is.

W-A

NT

414

-18,

20-5

3,15

-35,

6834

,95

Des

font

aini

aA

ster

ids

Unp

lace

dD

esfo

ntai

niac

eae

1C

osta

Ric

a to

Cap

e H

orn

AN

D3

1-3

5,33

-53,

00-4

4,17

17,6

7

Des

mar

iaE

udic

ots

San

tala

les

Lora

ntha

ceae

1C

hile

61

-35,

60-4

0,18

-37,

894,

58

Deu

tero

cohn

iaM

onoc

ots

Poa

les

Bro

mel

iace

ae13

Per

u, B

ol, S

W B

ra, N

Chi

le, N

Arg

S-A

MZ

31

-25,

00-2

6,17

-25,

581,

17

Dic

hant

heliu

mM

onoc

ots

Poa

les

Poa

ceae

58To

da A

mer

ica

W-N

T3

1-3

6,00

-38,

00-3

7,00

0,00

Dic

hond

raA

ster

ids

Sol

anal

esC

onvo

lvul

acea

e7

tropi

cs &

sub

tropi

cs, J

F1

1-2

5,00

-40,

83-3

2,92

15,8

3

Dic

kson

iaFi

licop

sida

Den

nsta

edtia

les

Dic

kson

iace

ae20

trop

Am

, Mal

esia

, New

Gui

nea,

Aus

, New

Cal

ed, N

ZS

PT

22

0,00

Dic

lipte

raA

ster

ids

Lam

iale

sA

cant

hace

ae20

0tro

pics

& s

ubtro

pics

11

-24,

50-2

5,50

-25,

001,

00

Die

lsio

chlo

aM

onoc

ots

Poa

les

Poa

ceae

1P

eru,

Bol

, Arg

, Chi

leA

ND

31

-18,

00-1

8,20

-18,

100,

20

Din

emag

onum

Ros

ids

Mal

pigh

iale

sM

alpi

ghia

ceae

1C

hile

61

-27,

50-3

1,50

-29,

504,

00

Din

eman

dra

Ros

ids

Mal

pigh

iale

sM

alpi

ghia

ceae

1C

hile

61

-21,

50-2

7,42

-24,

465,

92

Dio

scor

eaM

onoc

ots

Dio

scor

eale

sD

iosc

orea

ceae

400

trop,

sub

trop,

war

m-te

mp

regi

ons

141

-23,

50-4

2,00

-32,

7518

,50

Dio

stea

Ast

erid

sLa

mia

les

Verb

enac

eae

4te

mpe

rate

S A

m, A

rg, C

hile

51

-31,

00-4

3,83

-37,

4212

,83

Dip

lazi

umFi

licop

sida

Dry

opte

ridal

esD

ryop

terid

acea

e30

0co

smop

71

0,00

Dip

lole

pis

Ast

erid

sG

entia

nale

sA

pocy

nace

ae4

Chi

le, A

rg5

4-3

3,35

-47,

00-4

0,18

13,6

5

Dip

lost

ephi

umA

ster

ids

Ast

eral

esA

ster

acea

e90

trop

And

es fr

om C

olom

bia

to B

ol &

N C

hile

, 1 s

p C

osta

R

ica

AN

D3

3-1

7,58

-23,

00-2

0,29

5,42

Dip

osis

Ast

erid

sA

pial

esA

piac

eae

3te

mp

S A

m5

1-3

1,97

-34,

47-3

3,22

2,50

Dis

caria

Ros

ids

Ros

ales

Rha

mna

ceae

8A

us, N

Z, S

Am

AU

S2

4-2

8,72

-55,

00-4

1,86

26,2

8

Dis

sant

heliu

mM

onoc

ots

Poa

les

Poa

ceae

16P

eru,

Chi

le, B

ol to

Cal

iforn

iaA

NT-

P4

1-1

7,58

-18,

20-1

7,89

0,62

Dis

tichi

aM

onoc

ots

Poa

les

Junc

acea

e3

And

es S

Am

AN

D3

2-1

8,00

-21,

00-1

9,50

3,00

Dis

tichl

isM

onoc

ots

Poa

les

Poa

ceae

9A

m fr

om C

anad

a to

Arg

entin

a, 1

sp

Aus

tralia

C-P

AC

43

-18,

47-4

6,50

-32,

4828

,03

Dod

onae

aR

osid

sS

apin

dale

sS

apin

dace

ae68

tropi

cs &

sub

trop,

esp

ec A

ustra

lia1

1-3

1,00

-31,

17-3

1,08

0,17

Dom

eyko

aA

ster

ids

Api

ales

Api

acea

e4

Per

u C

hile

AN

D3

2-2

2,00

-25,

43-2

3,72

3,43

Don

atia

Ast

erid

sA

ster

ales

Sty

lidia

ceae

2Ta

sman

, NZ,

sub

anta

rctic

S A

mA

US

21

-37,

80-5

6,00

-46,

9018

,20

Don

ioph

yton

Ast

erid

sA

ster

ales

Ast

erac

eae

2A

ndes

Chi

le A

rgA

ND

32

-21,

50-3

0,67

-26,

089,

17

Doo

dia

Filic

opsi

daB

lech

nale

sB

lech

nace

ae12

Aus

tralia

, NZ,

Pac

ific

isla

nds,

Haw

aii,

Eas

ter I

slan

d, N

G

uine

aS

PT

21

0,00

Dow

ning

iaA

ster

ids

Ast

eral

esC

ampa

nula

ceae

11W

N A

m, 1

Chi

leA

NT-

P4

1-3

1,50

-38,

00-3

4,75

6,50

Dra

baR

osid

sB

rass

ical

esB

rass

icac

eae

350

Arc

tic &

tem

p n

Hem

is, A

sia,

mon

tane

C &

S A

mW

-AN

T4

7-1

8,00

-55,

00-3

6,50

37,0

0

Dra

pete

sR

osid

sM

alva

les

Thym

elae

acea

e1

tem

p S

Am

51

-46,

33-5

6,00

-51,

179,

67

��9

Drim

ysM

agno

liids

Can

ella

les

Win

tera

ceae

11M

exic

o, C

& S

Am

, JF

W-N

T3

1-3

0,33

-55,

97-4

3,15

25,6

3

Dro

sera

Eud

icot

sC

aryo

phyl

lale

sD

rose

race

ae11

5co

smop

, esp

ec A

us, T

asm

& N

Z7

1-3

7,83

-56,

00-4

6,92

18,1

7

Dry

mar

iaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

50pa

ntro

p1

3-2

2,00

-25,

40-2

3,70

3,40

Dry

opte

risFi

licop

sida

Dry

opte

ridal

esD

ryop

terid

acea

e15

0co

smop

71

0,00

Dun

alia

Ast

erid

sS

olan

ales

Sol

anac

eae

5A

ndes

Col

ombi

a to

Arg

entin

aA

ND

31

-18,

20-1

8,83

-18,

520,

63

Dys

opsi

sR

osid

sM

alpi

ghia

les

Eup

horb

iace

ae1

And

es C

osta

Ric

a, E

cuad

or, C

hile

, Arg

, JF

AN

D3

1-3

0,67

-55,

00-4

2,83

24,3

3

Eccr

emoc

arpu

sA

ster

ids

Lam

iale

sB

igno

niac

eae

5P

eru,

Chi

le, A

rgA

ND

31

-32,

93-4

0,25

-36,

597,

32

Echi

nops

isE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae10

0S

Am

AN

D3

9-1

8,50

-35,

00-2

6,75

16,5

0

Elap

hogl

ossu

mFi

licop

sida

Ble

chna

les

Lom

ario

psid

acea

e40

0pa

ntro

ps &

sub

trop,

Mad

ag, I

P1

3-3

9,83

-43,

83-4

1,83

4,00

Elat

ine

Ros

ids

Mal

pigh

iale

sE

latin

acea

e12

tem

p, tr

op &

sub

trop

regi

ons

W-A

NT

41

-34,

80-3

9,63

-37,

224,

83

Eleo

char

isM

onoc

ots

Poa

les

Cyp

erac

eae

200

cosm

op, J

F7

19-1

7,67

-55,

00-3

6,33

37,3

3

Elod

eaM

onoc

ots

Alis

mat

ales

Hyd

roch

arita

ceae

9A

M te

mp

W-N

T3

1-1

9,25

-36,

50-2

7,88

17,2

5

Elym

usM

onoc

ots

Poa

les

Poa

ceae

150

tem

p &

sub

trop,

mon

tane

trop

ics

76

-22,

00-5

3,00

-37,

5031

,00

Elyt

ropu

sA

ster

ids

Gen

tiana

les

Apo

cyna

ceae

1C

hile

, Arg

51

-33,

45-4

3,83

-38,

6410

,38

Embo

thriu

mE

udic

ots

Pro

teal

esP

rote

acea

e1

Chi

le y

Arg

entin

a, 1

spp

.5

1-3

5,25

-55,

67-4

5,46

20,4

2

Empe

trum

Ast

erid

sE

rical

esE

ricac

eae

3N

tem

p &

Arc

tic, E

urop

a, S

And

es, F

alkl

and,

JF

W-A

NT

41

-32,

83-5

6,00

-44,

4223

,17

Ence

liaA

ster

ids

Ast

eral

esA

ster

acea

e15

SW

US

A, M

exic

o, C

hile

, Per

u, G

alap

agos

AN

T-P

41

-19,

28-3

0,67

-24,

9811

,38

Enne

apog

onM

onoc

ots

Poa

les

Poa

ceae

35te

mp

Eur

asia

, Am

, tro

p &

sub

trop

71

-19,

87-2

3,00

-21,

433,

13

Ephe

dra

Gne

tops

ida

Eph

edra

les

Eph

edra

ceae

60E

uras

ia, A

MW

-AN

T4

7-1

8,17

-53,

00-3

5,58

34,8

3

Epilo

bium

Ros

ids

Myr

tale

sO

nagr

acea

e16

4te

mp

Eur

o, N

Am

, arc

tic, m

onta

ne tr

opW

-AN

T4

12-1

8,00

-55,

50-3

6,75

37,5

0

Epip

etru

mM

onoc

ots

Dio

scor

eale

sD

iosc

orea

ceae

3C

hile

63

-25,

00-3

7,67

-31,

3312

,67

Equi

setu

mS

phen

op-

sida

Equ

iset

ales

Equ

iset

acea

e30

alm

ost c

osm

op, M

adag

72

-18,

40-4

4,35

-31,

3825

,95

Erag

rost

isM

onoc

ots

Poa

les

Poa

ceae

300

tem

p &

trop

, ID

76

-18,

17-3

8,00

-28,

0819

,83

Erci

llaE

udic

ots

Car

yoph

ylla

les

Phy

tola

ccac

eae

2C

hile

62

-33,

50-4

4,58

-39,

0411

,08

Erec

htite

sA

ster

ids

Ast

eral

esA

ster

acea

e10

N &

S A

mW

-NT

31

-23,

53-2

5,92

-24,

732,

38

Erem

ocha

risA

ster

ids

Api

ales

Api

acea

e9

And

es C

hile

Per

uA

ND

31

-21,

50-2

7,07

-24,

285,

57

Erem

odra

baR

osid

sB

rass

ical

esB

rass

icac

eae

2A

ndes

Chi

le P

eru

AN

D3

1-1

9,80

-19,

80-1

9,80

0,00

Eria

chae

nium

Ast

erid

sA

ster

ales

Ast

erac

eae

1Ti

erra

del

Fue

go5

1-4

8,83

-53,

50-5

1,17

4,67

Erig

eron

Ast

erid

sA

ster

ales

Ast

erac

eae

200

Eur

asia

, N &

S A

m, A

us, J

FW

-AN

T4

12-2

9,00

-56,

00-4

2,50

27,0

0

Erio

syce

Eud

icot

sC

aryo

phyl

lale

sC

acta

ceae

35C

hile

, Arg

, S P

eru

AN

D3

28-1

8,42

-36,

72-2

7,57

18,3

0

Erod

ium

Ros

ids

Ger

ania

les

Ger

ania

ceae

75C

osm

op7

1-3

2,75

-33,

38-3

3,07

0,63

Erra

zuriz

iaR

osid

sFa

bale

sFa

bace

ae4

SW

US

, Mex

, 1 s

p C

hile

AN

T-P

41

-25,

00-3

0,50

-27,

755,

50

Eryn

gium

Ast

erid

sA

pial

esA

piac

eae

240

trop

& te

mp,

Eur

asia

, Am

, Afr,

Mal

esia

, Aus

, JF

710

-25,

20-4

0,60

-32,

9015

,40

Esca

lloni

aA

ster

ids

Unp

lace

dE

scal

loni

acea

e40

And

es S

Am

, SE

Bra

sil,

JFA

ND

313

-18,

50-5

3,00

-35,

7534

,50

Eucr

yphi

aR

osid

sO

xalid

ales

Cun

onia

ceae

6S

E A

us, T

asm

, Chi

leA

US

22

-36,

08-4

5,50

-40,

799,

42

Eude

ma

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e6

And

es E

cuad

or to

Arg

AN

D3

2-1

8,23

-22,

95-2

0,59

4,72

��0

Euly

chni

aE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae7

Per

u, C

hile

AN

D3

6-1

8,50

-32,

25-2

5,38

13,7

5

Euph

orbi

aR

osid

sM

alpi

ghia

les

Eup

horb

iace

ae20

00su

bcos

mop

719

-18,

47-5

2,67

-35,

5734

,20

Euph

rasi

aA

ster

ids

Lam

iale

sO

roba

ncha

ceae

300

tem

p N

Hem

ishp

, Eur

o, M

iddl

e as

ia, A

ustra

lia, N

Z, te

mp

S A

m, J

FW

-AN

T4

15-3

0,00

-55,

00-4

2,50

25,0

0

Evol

vulu

sA

ster

ids

Sol

anal

esC

onvo

lvul

acea

e10

0tro

ps &

sub

trops

11

-35,

33-3

7,83

-36,

582,

50

Exod

econ

usA

ster

ids

Sol

anal

esS

olan

acea

e6

S A

m, P

eru,

Ecu

ador

, Gal

apag

os, N

Chi

leA

ND

32

-18,

38-2

4,00

-21,

195,

62

Fabi

ana

Ast

erid

sS

olan

ales

Sol

anac

eae

15S

Per

u, B

ol, C

hile

, Arg

AN

D3

7-1

8,20

-45,

77-3

1,98

27,5

7

Face

lisA

ster

ids

Ast

eral

esA

ster

acea

e3

S B

rasi

l, P

eru,

Bol

, Par

agua

y, U

rug,

Chi

le, A

rgS

-AM

Z3

2-2

5,40

-39,

82-3

2,61

14,4

2

Fago

nia

Ros

ids

Zygo

phyl

lale

sZy

goph

ylla

ceae

30M

edit,

Eur

o, S

W A

sia,

NE

Afr,

SW

N A

m, P

eru,

Chi

leW

-AN

T4

2-1

8,83

-31,

67-2

5,25

12,8

3

Fasc

icul

aria

Mon

ocot

sP

oale

sB

rom

elia

ceae

1C

hile

61

-33,

58-4

4,00

-38,

7910

,42

Fest

uca

Mon

ocot

sP

oale

sP

oace

ae50

0co

smop

724

-17,

67-5

6,00

-36,

8338

,33

Fitz

roya

Pin

opsi

daP

inal

esC

upre

ssac

eae

1C

hile

Arg

51

-40,

17-4

2,58

-41,

382,

42

Flav

eria

Ast

erid

sA

ster

ales

Ast

erac

eae

21W

N A

m, M

exic

o, S

Am

, 1 s

p A

usC

-PA

C4

1-1

9,00

-36,

60-2

7,80

17,6

0

Flou

rens

iaA

ster

ids

Ast

eral

esA

ster

acea

e32

SW

US

A, M

exic

o, P

eru,

Chi

le, A

rgA

NT-

P4

1-2

9,45

-33,

83-3

1,64

4,38

Fonk

iaA

ster

ids

Lam

iale

sG

ratio

lace

ae1

Chi

le, A

rg5

1-4

0,75

-40,

75-4

0,75

0,00

Frag

aria

Ros

ids

Ros

ales

Ros

acea

e15

N &

S A

m, t

emp

Eur

asW

-AN

T4

1-3

5,00

-46,

43-4

0,72

11,4

3

Fran

coa

Ros

ids

Ger

ania

les

Fran

coac

eae

1C

hile

61

-32,

23-4

3,50

-37,

8711

,27

Fran

keni

aE

udic

ots

Car

yoph

ylla

les

Fran

keni

acea

e50

tem

p E

uro,

Afr,

Aus

, Am

, ID

74

-20,

63-5

3,00

-36,

8232

,37

Fuch

sia

Ros

ids

Myr

tale

sO

nagr

acea

e10

5C

& S

Am

, NZ,

Tah

itiA

US

22

-28,

50-5

5,00

-41,

7526

,50

Fuer

tesi

mal

vaR

osid

sM

alva

les

Mal

vace

ae14

Mex

ico,

C A

ndes

, Gal

apag

os, I

DW

-NT

33

-18,

30-3

0,90

-24,

6012

,60

Gai

mar

dia

Mon

ocot

sP

oale

sC

entro

lepi

dace

ae3

New

Gui

nea,

NZ,

Tas

man

ia, T

Fueg

o, F

alkl

and

SP

T2

1-3

9,88

-56,

00-4

7,94

16,1

2

Gal

inso

gaA

ster

ids

Ast

eral

esA

ster

acea

e15

tem

p &

sub

trop

C &

S A

mW

-NT

32

-17,

58-4

2,62

-30,

1025

,03

Gal

ium

Ast

erid

sG

entia

nale

sR

ubia

ceae

350

cosm

op, J

F7

17-1

8,00

-56,

00-3

7,00

38,0

0

Gam

ocar

pha

Ast

erid

sA

ster

ales

Cal

ycer

acea

e6

tem

p S

Am

56

-36,

83-4

6,00

-41,

429,

17

Gam

ocha

eta

Ast

erid

sA

ster

ales

Ast

erac

eae

40N

& S

Am

, Bra

zil,

JFW

-NT

322

-17,

58-5

5,25

-36,

4237

,67

Gam

ocha

etop

sis

Ast

erid

sA

ster

ales

Ast

erac

eae

1S

Arg

, Chi

le5

1-3

7,48

-37,

48-3

7,48

0,00

Gau

lther

iaA

ster

ids

Eric

ales

Eric

acea

e14

0E

& S

E A

sia,

Mal

esia

, Aus

, NZ,

N, C

, S A

m, W

I, JF

C-P

AC

411

-30,

67-5

6,00

-43,

3325

,33

Gav

ilea

Mon

ocot

sA

spar

agal

esO

rchi

dace

ae15

Tem

p S

Am

, JF

512

-30,

67-5

5,33

-43,

0024

,67

Gay

ophy

tum

Ros

ids

Myr

tale

sO

nagr

acea

e9

tem

p W

N A

m &

W S

Am

AN

T-P

42

-28,

50-3

7,00

-32,

758,

50

Gen

tiana

Ast

erid

sG

entia

nale

sG

entia

nace

ae36

0E

uro,

NW

Afr,

Asi

a, E

Aus

, Am

W-A

NT

46

-18,

00-5

5,00

-36,

5037

,00

Gen

tiane

llaA

ster

ids

Gen

tiana

les

Gen

tiana

ceae

256

Eur

asia

, NW

Afr,

N, C

& S

Am

, Aus

, NZ

73

-21,

00-5

6,00

-38,

5035

,00

Geo

ffroe

aR

osid

sFa

bale

sFa

bace

ae3

S A

m fr

om C

olom

bia

& V

enez

to A

rg, B

razi

l A

ND

31

-18,

47-3

1,50

-24,

9813

,03

Ger

aniu

mR

osid

sG

eran

iale

sG

eran

iace

ae40

0co

smop

, esp

ec te

m re

gs &

trop

ic m

onta

ne7

5-1

7,67

-55,

50-3

6,58

37,8

3

Get

hyum

Mon

ocot

sA

spar

agal

esA

lliac

eae

2C

hile

62

-29,

50-3

4,25

-31,

884,

75

Geu

mR

osid

sR

osal

esR

osac

eae

50A

rctic

& te

mp

regi

ons

74

-33,

95-5

5,00

-44,

4821

,05

Gev

uina

Eud

icot

sP

rote

ales

Pro

teac

eae

3N

ew G

uine

a, A

us, C

hile

, Arg

SP

T2

1-3

5,40

-45,

42-4

0,41

10,0

2

Gili

aA

ster

ids

Eric

ales

Pol

emon

iace

ae25

W N

Am

& S

Am

AN

T-P

44

-18,

00-5

1,00

-34,

5033

,00

���

Gill

iesi

aM

onoc

ots

Asp

arag

ales

Alli

acea

e3

Chi

le, A

rg5

2-3

2,37

-38,

25-3

5,31

5,88

Gla

ndul

aria

Ast

erid

sLa

mia

les

Verb

enac

eae

100

disj

unct

N A

m S

Am

AN

T-P

46

-21,

27-3

8,00

-29,

6316

,73

Gle

iche

nia

Filic

opsi

daG

leic

heni

ales

Gle

iche

niac

eae

110

pant

rop,

Mad

ag, J

F1

4-3

7,23

-55,

83-4

6,53

18,6

0

Glin

usE

udic

ots

Car

yoph

ylla

les

Mol

lugi

nace

ae9

pant

rop

11

-31,

50-3

7,17

-34,

335,

67

Gly

ceria

Mon

ocot

sP

oale

sP

oace

ae45

tem

p &

sub

trop,

mon

tane

trop

ics

71

-33,

25-4

1,67

-37,

468,

42

Gly

cyrr

hiza

Ros

ids

Faba

les

Faba

ceae

20E

uras

ia, A

us, N

Am

& te

mp

S A

mW

-AN

T4

1-3

4,08

-37,

58-3

5,83

3,50

Gna

phal

ium

Ast

erid

sA

ster

ales

Ast

erac

eae

100

cosm

op7

30-1

7,58

-54,

50-3

6,04

36,9

2

Goc

hnat

iaA

ster

ids

Ast

eral

esA

ster

acea

e66

Trop

& S

ubtro

p A

m, S

US

A, S

E A

sia

C-P

AC

41

-30,

67-3

7,80

-34,

237,

13

Gom

orte

gaM

agno

liids

Laur

ales

Gom

orte

gace

ae1

Chi

le6

1-3

5,62

-37,

67-3

6,64

2,05

Gom

phre

naE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

100

trop

& s

ubtro

p A

m, A

fr, In

doch

ina,

Aus

12

-18,

00-2

2,50

-20,

254,

50

Gra

bow

skia

Ast

erid

sS

olan

ales

Sol

anac

eae

4M

exic

o, G

alap

agos

, tem

p S

Am

W-N

T3

1-2

5,40

-25,

40-2

5,40

0,00

Gra

mm

itis

Filic

opsi

daP

olyp

odia

les

Gra

mm

itida

ceae

400

New

Gui

nea,

Asi

a, A

m, A

us, M

adag

, war

m &

tem

p, J

F1

3-3

7,42

-55,

87-4

6,64

18,4

5

Gra

mm

ospe

rma

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e1

Pat

agon

ia5

1-5

1,00

-51,

00-5

1,00

0,00

Gra

tiola

Ast

erid

sLa

mia

les

Gra

tiola

ceae

25te

mp

& s

ubtro

p, m

onta

ne tr

opic

s7

1-3

4,77

-42,

50-3

8,63

7,73

Gre

igia

Mon

ocot

sP

oale

sB

rom

elia

ceae

27C

& S

Am

, JF

W-N

T3

2-3

5,00

-43,

50-3

9,25

8,50

Grin

delia

Ast

erid

sA

ster

ales

Ast

erac

eae

60W

N A

m &

S A

m, B

razi

lW

-NT

34

-18,

42-3

8,78

-28,

6020

,37

Gris

elin

iaA

ster

ids

Api

ales

Gris

elin

iace

ae6

NZ,

Chi

le, P

arag

uay,

S B

rasi

lA

US

24

-24,

93-4

9,20

-37,

0724

,27

Gui

ndili

aR

osid

sS

apin

dale

sS

apin

dace

ae3

Chi

le A

rgA

ND

31

-31,

00-3

5,97

-33,

484,

97

Gun

nera

Eud

icot

sG

unne

rale

sG

unne

race

ae40

Am

from

Mex

ic to

Chi

le, t

rop

& S

Afr,

Mad

ag, M

ales

ia,

Tasm

, NZ,

Haw

aii,

JF1

7-2

9,83

-56,

00-4

2,92

26,1

7

Gut

ierr

ezia

Ast

erid

sA

ster

ales

Ast

erac

eae

27W

N A

m &

Sou

th S

Am

AN

T-P

46

-21,

00-5

2,50

-36,

7531

,50

Guy

neso

mia

A

ster

ids

Ast

eral

esA

ster

acea

e1

Chi

le6

1-2

9,75

-31,

50-3

0,63

1,75

Gym

nach

neM

onoc

ots

Poa

les

Poa

ceae

1C

hile

61

-32,

88-4

0,00

-36,

447,

12

Gym

noph

yton

Ast

erid

sA

pial

esA

piac

eae

6A

ndes

Chi

le A

rgA

ND

35

-23,

50-3

4,55

-29,

0311

,05

Gyp

otha

mni

umA

ster

ids

Ast

eral

esA

ster

acea

e1

N C

hile

61

-25,

00-2

6,13

-25,

571,

13

Haa

geoc

ereu

sE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae20

Per

u, C

hile

AN

D3

2-1

8,33

-20,

00-1

9,17

1,67

Hab

enar

iaM

onoc

ots

Asp

arag

ales

Orc

hida

ceae

600

pant

rop

& s

ubtro

p1

1-2

8,50

-52,

00-4

0,25

23,5

0

Hal

orag

isE

udic

ots

Sax

ifrag

ales

Hal

orag

acea

e27

Aus

, N. C

aled

, NZ,

Rap

a, J

FA

US

22

0,00

Ham

adry

asE

udic

ots

Ran

uncu

lale

sR

anun

cula

ceae

5A

ntar

ctic

S A

m5

4-5

0,62

-54,

92-5

2,77

4,30

Hap

lopa

ppus

Ast

erid

sA

ster

ales

Ast

erac

eae

75A

m e

spec

Chi

leW

-NT

364

-21,

18-4

5,50

-33,

3424

,32

Hap

lorh

usR

osid

sS

apin

dale

sA

naca

rdia

ceae

1C

hile

, Per

uA

ND

31

-18,

75-1

9,25

-19,

000,

50

Heb

eA

ster

ids

Lam

iale

sP

lant

agin

acea

e80

New

Gui

nea,

Aus

, Tas

m, N

Z, te

mp

S A

m, F

alkl

and

SP

T2

2-4

6,67

-56,

00-5

1,33

9,33

Hed

yotis

Ast

erid

sG

entia

nale

sR

ubia

ceae

150

tropi

cs e

spec

Asi

a &

Mal

esia

, JF

11

-34,

77-4

2,50

-38,

637,

73

Hel

eniu

mA

ster

ids

Ast

eral

esA

ster

acea

e32

W N

Am

, Mex

ico,

C A

mer

ica,

And

esW

-NT

37

-22,

00-4

0,33

-31,

1718

,33

Hel

icto

tric

hon

Mon

ocot

sP

oale

sP

oace

ae50

tem

p N

Hem

is, m

onta

ne tr

ops,

S A

fr7

1-3

6,83

-36,

83-3

6,83

0,00

Hel

iotr

opiu

mA

ster

ids

Unp

lace

dB

orag

inac

eae

260

tem

p, s

ubtro

p, tr

op re

gs, a

rid7

24-1

8,20

-38,

63-2

8,42

20,4

3

Hel

ogyn

eA

ster

ids

Ast

eral

esA

ster

acea

e8

Per

u, C

hile

, Bol

, Arg

AN

D3

2-1

7,58

-23,

87-2

0,73

6,28

���

Her

bert

iaM

onoc

ots

Asp

arag

ales

Irida

ceae

5te

mp

S A

m, C

hile

, Bra

zil

S-A

MZ

31

-33,

00-4

0,00

-36,

507,

00

Her

reria

Mon

ocot

sA

spar

agal

esA

gava

ceae

8B

rasi

l, U

ru, P

arag

, Arg

, Chi

leS

-AM

Z3

1-3

5,33

-37,

00-3

6,17

1,67

Het

eros

perm

aA

ster

ids

Ast

eral

esA

ster

acea

e5

SW

US

A, M

exic

o, C

Am

, Per

u, A

rg, C

hile

W-N

T3

2-1

7,58

-23,

00-2

0,29

5,42

Het

eroz

oste

raM

onoc

ots

Alis

mat

ales

Zost

erac

eae

1co

asta

l tem

p A

us, T

asm

, N C

hile

AU

S2

1-3

0,20

-30,

30-3

0,25

0,10

Hie

raci

umA

ster

ids

Ast

eral

esA

ster

acea

e20

0te

mp

regi

ons

& m

onta

ne tr

opic

s, e

xcl A

ustra

lasi

aW

-AN

T4

6-3

4,65

-55,

00-4

4,83

20,3

5

Hip

puris

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

1ci

rcum

bore

al, S

Am

, Aus

tralia

W-A

NT

41

-37,

00-5

5,00

-46,

0018

,00

His

tiopt

eris

Filic

opsi

daD

enns

taed

tiale

sD

enns

taed

tiace

ae5

pant

rop,

Mad

ag1

1-4

2,05

-54,

07-4

8,06

12,0

2

Hof

fman

nseg

gia

Ros

ids

Faba

les

Faba

ceae

2825

SW

US

, Chi

le, A

rgA

NT-

P4

7-1

8,00

-33,

45-2

5,73

15,4

5

Hol

lerm

ayer

aR

osid

sB

rass

ical

esB

rass

icac

eae

1C

hile

61

-38,

67-4

1,00

-39,

832,

33

Hom

aloc

arpu

sA

ster

ids

Api

ales

Api

acea

e6

Chi

le6

6-2

5,20

-33,

75-2

9,48

8,55

Hor

deum

Mon

ocot

sP

oale

sP

oace

ae30

tem

p &

sub

trop

regs

, mon

tane

trop

ics

W-A

NT

411

-18,

18-5

5,00

-36,

5936

,82

Hua

naca

Ast

erid

sA

pial

esA

piac

eae

4P

atag

onia

54

-36,

60-5

2,95

-44,

7816

,35

Hui

dobr

iaA

ster

ids

Cor

nale

sLo

asac

eae

2C

hile

62

-19,

00-2

9,50

-24,

2510

,50

Hup

erzi

aLy

cops

ida

Lyco

podi

ales

Lyco

podi

acea

e35

0S

ubco

smop

, Mad

ag7

1-5

3,67

-54,

67-5

4,17

1,00

Hyb

anth

usR

osid

sM

alpi

ghia

les

Viol

acea

e13

0tro

ps &

sub

trops

11

-36,

50-3

7,08

-36,

790,

58

Hyd

rang

eaA

ster

ids

Cor

nale

sH

ydra

ngea

ceae

29H

imal

ayas

, Asi

a, P

hilip

pine

s, N

, C A

m, A

ndes

to C

hile

W-A

NT

41

-33,

50-4

6,67

-40,

0813

,17

Hyd

roco

tyle

Ast

erid

sA

pial

esA

piac

eae

130

cosm

op7

8-1

8,83

-51,

27-3

5,05

32,4

3

Hym

enog

loss

umFi

licop

sida

Hym

enop

hylla

-le

sH

ymen

ophy

llace

ae1

Chi

le, J

F6

1-3

7,60

-49,

15-4

3,38

11,5

5

Hym

enop

hyllu

mFi

licop

sida

Hym

enop

hylla

-le

sH

ymen

ophy

llace

ae25

0tro

p &

S te

mp,

Eur

o, J

apan

, Mad

ag, J

F7

19-3

0,67

-56,

00-4

3,33

25,3

3

Hyp

eric

umR

osid

sM

alpi

ghia

les

Hyp

eric

acea

e37

0te

mp

regs

, mon

tane

trop

ics

72

-34,

77-4

2,50

-38,

637,

73

Hyp

ocha

eris

Ast

erid

sA

ster

ales

Ast

erac

eae

55te

mp

S A

m, E

uras

ia M

edit,

N A

frW

-AN

T4

33-1

7,58

-54,

00-3

5,79

36,4

2

Hyp

olep

isFi

licop

sida

Den

nsta

edtia

les

Den

nsta

edtia

ceae

40tro

p &

sub

trop,

Mad

ag1

1-3

0,67

-49,

15-3

9,91

18,4

8

Hyp

sela

Ast

erid

sA

ster

ales

Cam

panu

lace

ae5

E A

us, N

Z, A

ndes

S A

mA

US

21

-18,

00-5

4,00

-36,

0036

,00

Impe

rata

Mon

ocot

sP

oale

sP

oace

ae9

trop,

sub

trop

& te

mp

regi

ons

74

-30,

60-4

0,33

-35,

479,

73

Ipom

oea

Ast

erid

sS

olan

ales

Con

volv

ulac

eae

650

pant

rop

11

0,00

Ipom

opsi

sA

ster

ids

Eric

ales

Pol

emon

iace

ae26

W N

Am

& F

lorid

a, 1

Chi

le A

rgA

NT-

P4

1-2

1,00

-26,

93-2

3,97

5,93

Isoe

tes

Lyco

psid

aIs

oeta

les

Isoe

tace

ae15

0co

smop

, Mad

ag7

2-3

7,80

-53,

53-4

5,67

15,7

3

Ivan

iaR

osid

sB

rass

ical

esB

rass

icac

eae

1N

Chi

le6

1-2

7,40

-27,

40-2

7,40

0,00

Jabo

rosa

Ast

erid

sS

olan

ales

Sol

anac

eae

23te

mp

S A

m. S

Per

u, B

ol, B

ras,

Par

ag, C

hile

, Arg

, Uru

S-A

MZ

39

-18,

18-5

3,83

-36,

0135

,65

Jara

vaM

onoc

ots

Poa

les

Poa

ceae

59A

rg, B

ol, C

hile

, Col

ombi

a, C

osR

ica,

Ecu

ador

, El S

alva

dor,

Gua

tem

ala,

Mex

ico,

Per

u, U

SA

, Uru

guay

, Ven

ezue

laW

-NT

322

-17,

67-5

3,50

-35,

5835

,83

Jove

llana

Ast

erid

sLa

mia

les

Cal

ceol

aria

ceae

5N

Z, C

hile

AU

S2

3-3

5,75

-40,

00-3

7,88

4,25

Juan

iaM

onoc

ots

Are

cale

sA

reca

ceae

1JF

61

0,00

Juba

eaM

onoc

ots

Are

cale

sA

reca

ceae

1C

hile

61

-30,

50-3

5,50

-33,

005,

00

Junc

usM

onoc

ots

Poa

les

Junc

acea

e27

5C

osm

op, J

F, IP

728

-17,

58-5

6,00

-36,

7938

,42

���

June

llia

Ast

erid

sLa

mia

les

Verb

enac

eae

47P

eru,

Bol

, Chi

le, A

rgA

ND

331

-18,

00-5

2,50

-35,

2534

,50

Kag

enec

kia

Ros

ids

Ros

ales

Ros

acea

e4

Chi

le, P

eru,

Bol

ivia

, Arg

AN

D3

2-3

0,33

-38,

67-3

4,50

8,33

Koe

leria

Mon

ocot

sP

oale

sP

oace

ae45

tem

p &

sub

trops

, mon

tane

trop

s7

2-3

1,50

-54,

83-4

3,17

23,3

3

Kra

mer

iaR

osid

sZy

goph

ylla

les

Kra

mer

iace

ae18

SW

US

, Mex

ico,

Ecu

ador

, Chi

le, A

rg, B

razi

lW

-NT

32

-17,

58-3

3,50

-25,

5415

,92

Kur

zam

raA

ster

ids

Lam

iale

sLa

mia

ceae

1C

hile

/ A

rgA

ND

31

-28,

72-3

0,50

-29,

611,

78

Kyl

linga

Mon

ocot

sP

oale

sC

yper

acea

e40

pant

rops

11

0,00

Lact

oris

Mag

nolii

dsP

iper

ales

Lact

orid

acea

e1

JF6

10,

00

Lage

noph

ora

Ast

erid

sA

ster

ales

Ast

erac

eae

15S

E A

sia,

Mal

esia

, Aus

tralia

, Tas

man

ia, N

Z, C

entra

l and

S

outh

Am

eric

a, J

F, F

alkl

and,

Tris

tan

da C

unha

. S

PT

23

-33,

67-5

6,00

-44,

8322

,33

Lam

paya

Ast

erid

sLa

mia

les

Verb

enac

eae

3B

ol, C

hile

, Arg

AN

D3

1-1

8,17

-23,

50-2

0,83

5,33

Land

oltia

Mon

ocot

sA

lism

atal

esLe

mna

ceae

1co

smop

71

-55,

50-5

5,50

-55,

500,

00

Lapa

geria

Mon

ocot

sLi

liale

sP

hile

siac

eae

1C

hile

6

1-3

0,67

-40,

83-3

5,75

10,1

7

Lard

izab

ala

Eud

icot

sR

anun

cula

les

Lard

izab

alac

eae

1C

hile

and

adj

acen

t Arg

entin

a5

1-3

2,50

-39,

87-3

6,18

7,37

Lare

tiaA

ster

ids

Api

ales

Api

acea

e1

Chi

le, A

rgen

tina

AN

D3

1-3

0,17

-35,

50-3

2,83

5,33

Larr

eaR

osid

sZy

goph

ylla

les

Zygo

phyl

lace

ae5

SW

N A

m, S

Am

AN

T-P

43

-28,

80-3

6,75

-32,

787,

95

Last

arria

eaE

udic

ots

Car

yoph

ylla

les

Pol

ygon

acea

e3

Cal

iforn

ia, C

hile

AN

T-P

41

-27,

83-3

5,97

-31,

908,

13

Last

heni

aA

ster

ids

Ast

eral

esA

ster

acea

e18

SW

US

, Chi

le-A

rgA

NT-

P4

1-3

1,25

-51,

00-4

1,13

19,7

5

Lath

yrus

Ros

ids

Faba

les

Faba

ceae

155

tem

p N

hem

is, M

edit,

mon

tane

trop

ics,

E A

fr, S

Am

W-A

NT

411

-29,

47-5

3,00

-41,

2323

,53

Latu

aA

ster

ids

Sol

anal

esS

olan

acea

e1

Chi

le6

1-4

0,17

-43,

00-4

1,58

2,83

Laur

elia

Mag

nolii

dsLa

ural

esA

ther

ospe

rmat

a-ce

ae2

NZ

& C

hile

AU

S2

1-3

4,73

-41,

17-3

7,95

6,43

Laur

elio

psis

Mag

nolii

dsLa

ural

esA

ther

ospe

rmat

a-ce

ae1

Chi

le A

rg5

1-3

8,37

-46,

67-4

2,52

8,30

Lebe

tant

hus

Ast

erid

sE

rical

esE

ricac

eae

1P

atag

onia

51

-43,

42-5

6,00

-49,

7112

,58

Leca

noph

ora

Ros

ids

Mal

vale

sM

alva

ceae

7A

rg, C

hile

51

-52,

10-5

2,10

-52,

100,

00

Lege

nere

Ast

erid

sA

ster

ales

Cam

panu

lace

ae2

1 C

alif,

1 C

hile

AN

T-P

41

-40,

00-4

0,00

-40,

000,

00

Legr

andi

aR

osid

sM

yrta

les

Myr

tace

ae1

Chi

le6

1-3

5,27

-36,

68-3

5,98

1,42

Lem

naM

onoc

ots

Alis

mat

ales

Lem

nace

ae14

Cos

mop

73

-18,

00-4

6,50

-32,

2528

,50

Lenz

iaE

udic

ots

Car

yoph

ylla

les

Por

tula

cace

ae1

Chi

le A

rgA

ND

31

-23,

00-3

0,25

-26,

637,

25

Leon

toch

irM

onoc

ots

Lilia

les

Als

troem

eria

ceae

1C

hile

61

-27,

90-2

8,08

-27,

990,

18

Lepe

chin

iaA

ster

ids

Lam

iale

sLa

mia

ceae

40C

alifo

rnia

, And

es, M

exic

o to

S B

rasi

l, A

rg, C

hile

W

-NT

33

-29,

83-4

0,00

-34,

9210

,17

Lepi

dium

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e18

0C

osm

op, I

D7

22-1

8,00

-55,

00-3

6,50

37,0

0

Lepi

doce

ras

Eud

icot

sS

anta

lale

sS

anta

lace

ae2

S P

eru,

Chi

leA

ND

31

-36,

33-4

3,00

-39,

676,

67

Lepi

doph

yllu

mA

ster

ids

Ast

eral

esA

ster

acea

e1

Pat

agon

ia C

hile

-Arg

.5

1-5

2,33

-53,

00-5

2,67

0,67

Lepi

doth

amnu

sP

inop

sida

Pin

ales

Pod

ocar

pace

ae3

2 N

Z, 1

S C

hile

/Arg

AU

S2

1-4

0,17

-55,

00-4

7,58

14,8

3

Lept

inel

laA

ster

ids

Ast

eral

esA

ster

acea

e33

New

Gui

nea,

Aus

, NZ,

Sub

anta

rctic

Is, S

Am

SP

T2

1-4

0,00

-56,

00-4

8,00

16,0

0

Lept

ocar

pha

Ast

erid

sA

ster

ales

Ast

erac

eae

1C

hile

61

-35,

78-4

0,82

-38,

305,

03

��4

Lept

ochl

oaM

onoc

ots

Poa

les

Poa

ceae

17tro

pics

& s

ubtro

p1

3-1

8,42

-24,

00-2

1,21

5,58

Lept

ophy

lloch

loa

Mon

ocot

sP

oale

sP

oace

ae1

Arg

Chi

le, J

F5

1-3

8,00

-40,

00-3

9,00

2,00

Lept

ostig

ma

Ast

erid

sG

entia

nale

sR

ubia

ceae

6S

E A

sia,

NZ,

W S

Am

SP

T2

1-3

6,75

-42,

47-3

9,61

5,72

Lepu

rope

talo

nE

udic

ots

Sax

ifrag

ales

Sax

ifrag

acea

e1

SE

US

A, M

exic

o, C

hile

, Arg

Uru

guay

AN

T-P

41

-32,

57-4

0,00

-36,

287,

43

Leuc

heria

Ast

erid

sA

ster

ales

Ast

erac

eae

46P

eru,

Bol

, Arg

, Chi

le, F

alkl

and

AN

D3

43-1

7,58

-55,

00-3

6,29

37,4

2

Leuc

ocor

yne

Mon

ocot

sA

spar

agal

esA

lliac

eae

11C

hile

611

-20,

83-4

0,00

-30,

4219

,17

Leun

isia

Ast

erid

sA

ster

ales

Ast

erac

eae

1C

Chi

le6

1-3

0,80

-32,

75-3

1,78

1,95

Libe

rtia

Mon

ocot

sA

spar

agal

esIri

dace

ae10

New

Gui

nea,

E A

us, N

Z, A

ndes

S A

m, J

FS

PT

24

-32,

23-4

9,15

-40,

6916

,92

Liga

riaE

udic

ots

San

tala

les

Lora

ntha

ceae

2C

Per

u, E

Bol

, C C

hile

, Uru

g, n

Arg

, S B

rasi

lS

-AM

Z3

1-3

0,50

-37,

00-3

3,75

6,50

Lila

eaM

onoc

ots

Alis

mat

ales

Junc

agin

acea

e1

Pac

ific

Am

Can

ada

to C

hile

AN

T-P

41

-33,

22-4

2,05

-37,

638,

83

Lila

eops

isA

ster

ids

Api

ales

Api

acea

e13

Am

Can

ada

to T

Fueg

o, A

us, N

ZC

-PA

C4

1-1

8,25

-53,

67-3

5,96

35,4

2

Lim

oniu

mE

udic

ots

Car

yoph

ylla

les

Plu

mba

gina

ceae

250

N te

mpe

rate

, circ

umbo

real

, aus

tral

W-A

NT

42

-27,

07-3

2,55

-29,

815,

48

Lim

osel

laA

ster

ids

Lam

iale

sG

ratio

lace

ae15

Cos

mop

, Mad

ag7

1-1

8,00

-55,

00-3

6,50

37,0

0

Lina

nthu

sA

ster

ids

Eric

ales

Pol

emon

iace

ae35

W N

Am

, Chi

leA

NT-

P4

1-3

1,00

-37,

67-3

4,33

6,67

Lind

erni

aA

ster

ids

Lam

iale

sLi

nder

niac

eae

100

trops

, sub

trops

, tem

p, M

adag

71

-38,

60-3

8,60

-38,

600,

00

Linu

mR

osid

sM

alpi

ghia

les

Lina

ceae

200

tem

p &

sub

trops

75

-23,

00-3

7,58

-30,

2914

,58

Lipp

iaA

ster

ids

Lam

iale

sVe

rben

acea

e20

0A

m tr

op, m

any

in A

fr tro

p1

2-2

3,50

-30,

50-2

7,00

7,00

Lith

rea

Ros

ids

Sap

inda

les

Ana

card

iace

ae4

S A

m: A

rg, B

ol, P

arag

uay,

SE

Bra

zil,

Chi

le.

S-A

MZ

31

-30,

00-3

8,25

-34,

138,

25

Litto

rella

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

31

N A

m, 1

S A

m, 1

Eur

oW

-AN

T4

1-3

9,67

-55,

00-4

7,33

15,3

3

Llag

unoa

Ros

ids

Sap

inda

les

Sap

inda

ceae

3A

ndea

n tro

p S

Am

AN

D3

1-2

9,73

-34,

13-3

1,93

4,40

Loas

aA

ster

ids

Cor

nale

sLo

asac

eae

36C

hile

, Arg

, 1 s

p co

asta

l Per

uA

ND

336

-21,

03-5

0,00

-35,

5228

,97

Lobe

liaA

ster

ids

Ast

eral

esC

ampa

nula

ceae

330

trops

& s

ubtro

ps1

8-2

5,22

-43,

83-3

4,53

18,6

2

Lom

atia

Eud

icot

sP

rote

ales

Pro

teac

eae

12A

us, T

as, S

Am

AU

S2

3-3

0,28

-50,

00-4

0,14

19,7

2

Loph

opap

pus

Ast

erid

sA

ster

ales

Ast

erac

eae

6P

eru,

Bol

, Chi

le, A

rgA

ND

32

-18,

00-2

0,08

-19,

042,

08

Loph

osor

iaFi

licop

sida

Dic

kson

iale

sLo

phos

oria

ceae

1tro

p A

m

W-N

T3

1-3

5,22

-46,

42-4

0,82

11,2

0

Lotu

sR

osid

sFa

bale

sFa

bace

ae10

0M

acar

ones

ia, t

emp

eura

sia,

Afr,

Aus

, S A

m7

1-2

4,83

-40,

00-3

2,42

15,1

7

Luci

liaA

ster

ids

Ast

eral

esA

ster

acea

e12

And

es S

Am

, Chi

le, A

rg, P

eru

AN

D3

3-3

3,00

-38,

67-3

5,83

5,67

Luci

liocl

ine

Ast

erid

sA

ster

ales

Ast

erac

eae

12A

ndes

S A

m, C

hile

, Arg

, Per

u A

ND

32

-17,

58-2

3,50

-20,

545,

92

Ludw

igia

Ros

ids

Myr

tale

sO

nagr

acea

e81

Cos

mop

74

-28,

50-4

0,60

-34,

5512

,10

Lum

aR

osid

sM

yrta

les

Myr

tace

ae4

Chi

le A

rg5

2-2

9,50

-45,

42-3

7,46

15,9

2

Lupi

nus

Ros

ids

Faba

les

Faba

ceae

220

N &

S A

m, M

edit,

mon

tane

trop

s, E

Afr

W-A

NT

46

-18,

00-4

0,00

-29,

0022

,00

Luzu

laM

onoc

ots

Poa

les

Junc

acea

e75

cosm

op, J

F7

6-3

0,00

-56,

00-4

3,00

26,0

0

Luzu

riaga

Mon

ocot

sLi

liale

sLu

zuria

gace

ae3

1 N

Z, 3

Chi

le, A

rg to

T F

uego

AU

S2

3-3

4,77

-56,

00-4

5,38

21,2

3

Lyca

psus

Ast

erid

sA

ster

ales

Ast

erac

eae

1D

esve

ntur

adas

Isla

nds

(Chi

le)

61

0,00

Lyci

umA

ster

ids

Sol

anal

esS

olan

acea

e80

N A

m, t

em S

Am

, Eur

asia

, S A

fr, A

us, I

P7

9-1

8,00

-35,

33-2

6,67

17,3

3

Lyco

pers

icon

Ast

erid

sS

olan

ales

Sol

anac

eae

10S

Am

Col

ombi

a to

Chi

le, B

razi

lA

ND

31

-18,

27-2

8,88

-23,

5810

,62

Lyco

podi

umLy

cops

ida

Lyco

podi

ales

Lyco

podi

acea

e55

trop

& te

mp

75

-37,

00-5

6,00

-46,

5019

,00

���

Lysi

mac

hia

Ast

erid

sE

rical

esM

yrsi

nace

ae15

0C

osm

op7

1-3

6,33

-41,

67-3

9,00

5,33

Mac

haer

ina

Mon

ocot

sP

oale

sC

yper

acea

e45

trops

esp

Aus

tralia

, Haw

aii,

JF1

10,

00

Mac

rach

aeni

umA

ster

ids

Ast

eral

esA

ster

acea

e1

Pat

agon

ia &

Tfu

ego

51

-38,

27-5

5,00

-46,

6316

,73

Mad

iaA

ster

ids

Ast

eral

esA

ster

acea

e10

W N

Am

, Chi

le-A

rgA

NT-

P4

2-3

0,00

-55,

00-4

2,50

25,0

0

Mag

alla

naR

osid

sB

rass

ical

esTr

opae

olac

eae

2C

hile

, Arg

51

-46,

53-4

6,53

-46,

530,

00

Mai

huen

iaE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae2

Chi

le A

rg5

1-3

5,55

-37,

50-3

6,53

1,95

Mai

huen

iops

isE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae18

And

es P

eru,

Bol

, Chi

le, A

rgA

ND

311

-19,

42-3

0,00

-24,

7110

,58

Mai

rean

aE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

58A

ustra

lia, C

hile

AU

S2

1-2

7,68

-27,

68-2

7,68

0,00

Mal

acot

hrix

Ast

erid

sA

ster

ales

Ast

erac

eae

16W

N A

m, C

hile

AN

T-P

42

-22,

00-3

0,67

-26,

338,

67

Mal

eshe

rbia

Ros

ids

Mal

pigh

iale

sM

ales

herb

iace

ae24

And

es P

eru,

Chi

le, A

rgA

ND

318

-18,

50-3

5,00

-26,

7516

,50

Mal

vella

Ros

ids

Mal

vale

sM

alva

ceae

43

sp W

US

A, M

exic

o, 1

sp

Per

u to

Uru

guay

, 1 s

p. M

edit

W-A

NT

41

-25,

00-3

3,68

-29,

348,

68

Man

coa

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e7

Mex

ico,

And

es P

eru,

Bol

, Arg

, Chi

leA

NT-

P4

2-1

8,00

-25,

00-2

1,50

7,00

Mar

gyric

arpu

sR

osid

sR

osal

esR

osac

eae

3S

And

es to

Chi

le, J

F, B

razi

lS

-AM

Z3

1-2

5,00

-43,

67-3

4,33

18,6

7

Mar

sile

aFi

licop

sida

Mar

sile

ales

Mar

sile

acea

e70

trop

& te

mp,

Mad

ag7

1-3

0,63

-32,

15-3

1,39

1,52

Mar

sipp

ospe

r-m

umM

onoc

ots

Poa

les

Junc

acea

e3

NZ,

S S

Am

, Tfu

ego,

Fal

klan

dA

US

23

-37,

00-5

6,00

-46,

5019

,00

Mar

ticor

enia

Ast

erid

sA

ster

ales

Ast

erac

eae

1C

hile

61

-32,

72-3

3,60

-33,

160,

88

Mas

tigos

tyla

Mon

ocot

sA

spar

agal

esIri

dace

ae16

Per

u, A

rgA

ND

31

-18,

17-2

3,00

-20,

584,

83

Mat

hew

sia

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e7

Per

u, C

hile

AN

D3

6-2

2,00

-42,

00-3

2,00

20,0

0

May

tenu

sE

udic

ots

Cel

astra

les

Cel

astra

ceae

200

trops

& s

ubtro

ps1

4-2

8,50

-55,

00-4

1,75

26,5

0

Mec

ardo

nia

Ast

erid

sLa

mia

les

Gra

tiola

ceae

10A

m S

US

to A

rgW

-NT

31

-35,

33-3

5,47

-35,

400,

13

Meg

alac

hne

Mon

ocot

sP

oale

sP

oace

ae2

JF6

20,

00

Meg

alas

trum

Filic

opsi

daD

ryop

terid

ales

Dry

opte

ridac

eae

40Tr

op A

m, 1

Afr

to M

asca

rene

s (n

o M

adag

), JF

11

-30,

67-4

6,68

-38,

6816

,02

Mel

ica

Mon

ocot

sP

oale

sP

oace

ae80

tem

p re

gs &

mon

tane

trop

sW

-AN

T4

7-2

5,10

-39,

88-3

2,49

14,7

8

Mel

ospe

rma

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

2C

hile

, Arg

52

-31,

00-3

6,50

-33,

755,

50

Men

odor

aA

ster

ids

Lam

iale

sO

leac

eae

25tro

p &

sub

trop

Am

, S A

fr4

1-3

2,05

-32,

63-3

2,34

0,58

Men

onvi

llea

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e26

Chi

le, A

rg5

19-2

2,08

-51,

17-3

6,63

29,0

8

Men

tzel

iaA

ster

ids

Cor

nale

sLo

asac

eae

80C

hile

, Arg

to C

anad

a, W

IW

-NT

37

-22,

08-3

3,33

-27,

7111

,25

Met

harm

eR

osid

sZy

goph

ylla

les

Zygo

phyl

lace

ae1

N C

hile

61

-20,

08-2

0,63

-20,

360,

55

Mic

role

pia

Filic

opsi

daD

enns

taed

tiale

sD

enns

taed

tiace

ae45

pale

otro

p, 1

Afr,

Mad

ag, J

apan

, NZ

11

0,00

Mic

roph

yes

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae3

Chi

le6

3-2

4,67

-37,

50-3

1,08

12,8

3

Mic

rops

isA

ster

ids

Ast

eral

esA

ster

acea

e8

Bra

zil,

Arg

, Uru

g, C

hile

S-A

MZ

31

-31,

33-3

6,50

-33,

925,

17

Mic

rose

risA

ster

ids

Ast

eral

esA

ster

acea

e15

W N

Am

, 1 C

hile

, 1 A

us, N

Z C

-PA

C4

1-2

5,00

-37,

83-3

1,42

12,8

3

Mic

roso

rum

Filic

opsi

daP

olyp

odia

les

Pol

ypod

iace

ae40

trop

& w

arm

, Mad

ag1

10,

00

Mic

rost

eris

Ast

erid

sE

rical

esP

olem

onia

ceae

1W

N A

m &

S A

mA

NT-

P4

1-1

8,00

-30,

50-2

4,25

12,5

0

Mie

rsia

Mon

ocot

sA

spar

agal

esA

lliac

eae

1C

hile

61

-31,

00-3

5,33

-33,

174,

33

Mik

ania

Ast

erid

sA

ster

ales

Ast

erac

eae

430

pant

rops

11

-34,

50-3

7,92

-36,

213,

42

���

Mim

ulus

Ast

erid

sLa

mia

les

Phr

ymac

eae

150

Am

, S A

fr, M

adag

, E A

sia,

JF

78

-22,

00-4

5,50

-33,

7523

,50

Min

uart

iaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

120

Arc

tic &

tem

p re

gion

s, M

exic

o, E

thio

pia,

Him

alay

as,

Chi

leW

-AN

T4

1-3

3,00

-36,

75-3

4,88

3,75

Miq

uelio

punt

iaE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae1

Chi

le6

1-2

7,50

-30,

00-2

8,75

2,50

Mira

bilis

Eud

icot

sC

aryo

phyl

lale

sN

ycta

gina

ceae

54A

m tr

op, H

imal

, Chi

naW

-AN

T4

4-1

8,33

-36,

00-2

7,17

17,6

7

Mis

oden

drum

Eud

icot

sS

anta

lale

sM

isod

endr

acea

e8

S A

m, s

outh

of 3

35

8-3

3,00

-56,

00-4

4,50

23,0

0

Mitr

aria

Ast

erid

sLa

mia

les

Ges

neria

ceae

1C

hile

, Arg

.5

1-3

0,67

-49,

88-4

0,28

19,2

2

Mni

odes

Ast

erid

sA

ster

ales

Ast

erac

eae

4P

eru,

Chi

leA

ND

31

-17,

58-1

8,17

-17,

880,

58

Mon

tiaE

udic

ots

Car

yoph

ylla

les

Por

tula

cace

ae20

tem

p E

uras

ia, M

edit,

mon

tane

trop

ics,

Afr,

A A

us, A

& S

A

m,

71

-29,

50-5

5,00

-42,

2525

,50

Mon

tiops

isE

udic

ots

Car

yoph

ylla

les

Por

tula

cace

ae18

Chi

le, A

rgen

tina,

Bol

ivia

y P

erú

AN

D3

18-1

9,25

-40,

75-3

0,00

21,5

0

Mon

ttea

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

3C

hile

1, A

rg. 2

AN

D3

1-2

5,00

-31,

67-2

8,33

6,67

Mor

ella

Ros

ids

Faga

les

Myr

icac

eae

40S

udam

And

es P

eru

Ecu

ador

, Afri

ca, E

Asi

a, F

ilipi

nas

y M

alas

ia1

1-1

8,83

-19,

12-1

8,98

0,28

Mos

char

iaA

ster

ids

Ast

eral

esA

ster

acea

e2

Chi

le6

2-2

9,72

-36,

50-3

3,11

6,78

Mos

chop

sis

Ast

erid

sA

ster

ales

Cal

ycer

acea

e8

Per

ú, C

hile

, Arg

, Pat

agon

iaA

ND

34

-17,

67-5

5,00

-36,

3337

,33

Mue

hlen

beck

iaE

udic

ots

Car

yoph

ylla

les

Pol

ygon

acea

e25

New

Gui

nea,

Aus

, Tas

m, N

Z, W

S A

mS

PT

21

-19,

65-4

5,50

-32,

5825

,85

Muh

lenb

ergi

aM

onoc

ots

Poa

les

Poa

ceae

150

tem

p &

sub

trop

Am

, Asi

aW

-AN

T4

3-1

9,50

-41,

00-3

0,25

21,5

0

Mul

inum

Ast

erid

sA

pial

esA

piac

eae

20S

And

es5

7-2

0,98

-45,

48-3

3,23

24,5

0

Mun

roa

Mon

ocot

sP

oale

sP

oace

ae5

1 W

US

, 4 S

W S

Am

AN

T-P

42

-18,

18-2

4,00

-21,

095,

82

Mut

isia

Ast

erid

sA

ster

ales

Ast

erac

eae

60A

ndes

Col

ombi

a to

Chi

le, A

rg, P

arag

, SE

Bra

sil,

Uru

gA

ND

323

-17,

58-4

8,92

-33,

2531

,33

Myo

schi

los

Eud

icot

sS

anta

lale

sS

anta

lace

ae1

Chi

le, A

rg5

1-3

1,62

-55,

00-4

3,31

23,3

8

Myo

sotis

Ast

erid

sU

npla

ced

Bor

agin

acea

e10

0E

uras

ia, m

onta

ne tr

ops,

S A

fr, N

Gui

nea,

Aus

, NZ

W-A

NT

42

-48,

47-5

4,88

-51,

686,

42

Myo

suru

sE

udic

ots

Ran

uncu

lale

sR

anun

cula

ceae

15te

mp

regs

hem

is N

& S

, M a

peta

lus

disj

N A

mW

-AN

T4

2-2

9,80

-53,

38-4

1,59

23,5

8

Myr

ceug

enia

Ros

ids

Myr

tale

sM

yrta

ceae

40C

hile

, Arg

, SE

Bra

sil,

JFS

-AM

Z3

12-3

0,67

-46,

43-3

8,55

15,7

7

Myr

cian

thes

Ros

ids

Myr

tale

sM

yrta

ceae

50tro

p A

m, A

ndes

, Bra

zil,

WI

AN

D3

1-2

9,43

-30,

00-2

9,72

0,57

Myr

ioph

yllu

mE

udic

ots

Sax

ifrag

ales

Hal

orag

acea

e60

subc

osm

op7

2-1

9,50

-55,

00-3

7,25

35,5

0

Myr

teol

aR

osid

sM

yrta

les

Myr

tace

ae3

Col

ombi

a to

Chi

le, J

F, B

razi

lA

ND

31

-36,

23-5

6,00

-46,

1219

,77

Nam

aA

ster

ids

Unp

lace

dB

orag

inac

eae

45N

Am

from

Cal

iforn

ia,M

ex, C

Am

to B

rasi

l, W

I, H

awai

iW

-NT

32

-21,

50-2

5,50

-23,

504,

00

Nan

odea

Eud

icot

sS

anta

lale

sS

anta

lace

ae1

tem

p S

Am

51

-40,

33-5

6,00

-48,

1715

,67

Nar

doph

yllu

m

Ast

erid

sA

ster

ales

Ast

erac

eae

7B

ol, A

rg, C

hile

AN

D3

5-2

3,00

-53,

00-3

8,00

30,0

0

Nas

aA

ster

ids

Cor

nale

sLo

asac

eae

100

Col

ombi

a, E

cuad

or, P

eru,

few

in C

hile

, Bol

ivia

, Ven

ezu-

ela

and

C A

m (t

o S

Mex

ico)

.W

-NT

31

-20,

00-2

5,00

-22,

505,

00

Nas

sauv

iaA

ster

ids

Ast

eral

esA

ster

acea

e39

Bol

, Arg

, Chi

leA

ND

325

-30,

00-5

5,67

-42,

8325

,67

Nas

sella

Mon

ocot

sP

oale

sP

oace

ae79

C &

S A

mW

-NT

326

-17,

67-5

0,00

-33,

8332

,33

Nas

tant

hus

Ast

erid

sA

ster

ales

Cal

ycer

acea

e9

Chi

le, A

rg, F

alkl

and

Is.

55

-25,

00-4

6,00

-35,

5021

,00

Nav

arre

tiaA

ster

ids

Eric

ales

Pol

emon

iace

ae30

W N

Am

, Chi

le, A

rgA

NT-

P4

1-3

3,25

-40,

53-3

6,89

7,28

Neo

wer

derm

anni

aE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae2

Per

u, B

ol, C

hile

, Arg

AN

D3

1-1

8,50

-18,

50-1

8,50

0,00

Ner

tera

Ast

erid

sG

entia

nale

sR

ubia

ceae

13M

adag

, SE

Asi

a, M

ales

ia, A

us, N

Z, H

awai

i, S

Am

, JF

C-A

US

21

-30,

67-5

6,00

-43,

3325

,33

��7

Nes

ocar

yum

Ast

erid

sU

npla

ced

Bor

agin

acea

e1

Des

vent

urad

as Is

land

s (C

hile

)6

10,

00

Neu

onto

both

rys

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e2

Chi

le6

2-2

0,13

-20,

13-2

0,13

0,00

Nic

otia

naA

ster

ids

Sol

anal

esS

olan

acea

e67

Am

, US

A to

Chi

le, A

ust,

New

Cal

edon

ia, N

amib

ia, J

F1

10-1

8,47

-43,

62-3

1,04

25,1

5

Nie

rem

berg

iaA

ster

ids

Sol

anal

esS

olan

acea

e20

Mex

to A

rg, B

ras

W-N

T3

1-3

4,83

-51,

00-4

2,92

16,1

7

Nitr

ophi

laE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

8U

S, M

ex, C

hile

, Arg

AN

T-P

41

-22,

50-2

4,50

-23,

502,

00

Noc

caea

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e20

N te

mp,

few

S A

mW

-AN

T4

1-3

3,50

-54,

83-4

4,17

21,3

3

Nol

ana

Ast

erid

sS

olan

ales

Sol

anac

eae

77P

eru

to C

hile

, Gal

apag

osA

ND

336

-18,

25-4

0,17

-29,

2121

,92

Not

anth

era

Eud

icot

sS

anta

lale

sLo

rant

hace

ae1

Chi

le te

mpe

rate

, JF

61

-32,

57-4

3,00

-37,

7810

,43

Not

hofa

gus

Ros

ids

Faga

les

Not

hofa

gace

ae37

New

Gui

nea,

Aus

, Tas

m, N

Z, N

ew C

aled

, tem

p S

Am

SP

T2

10-3

2,95

-55,

97-4

4,46

23,0

2

Not

hola

ena

Filic

opsi

daP

terid

ales

Pte

ridac

eae

40w

arm

to tr

op A

m, S

W U

S, M

ex, M

adag

, JF

12

-17,

58-2

7,28

-22,

439,

70

Not

hosc

ordu

mM

onoc

ots

Asp

arag

ales

Alli

acea

e26

Am

W-N

T3

6-2

9,90

-40,

25-3

5,08

10,3

5

Not

icas

trum

Ast

erid

sA

ster

ales

Ast

erac

eae

20P

eru,

Bol

, Chi

le, A

rg, E

cuad

or, B

razi

lS

-AM

Z3

5-3

1,92

-38,

73-3

5,33

6,82

Not

otric

heR

osid

sM

alva

les

Mal

vace

ae10

0E

cuad

or, P

eru,

Bol

, Chi

le, A

rgA

ND

324

-17,

72-3

4,67

-26,

1916

,95

Och

agav

iaM

onoc

ots

Poa

les

Bro

mel

iace

ae4

Chi

le, J

F 6

3-3

1,67

-38,

25-3

4,96

6,58

Oen

othe

raR

osid

sM

yrta

les

Ona

grac

eae

121

tem

p &

sub

trop

N C

& S

Am

W-N

T3

14-1

8,00

-52,

00-3

5,00

34,0

0

Ols

yniu

mM

onoc

ots

Asp

arag

ales

Irida

ceae

20A

ndea

n an

d te

mp

S A

m a

nd N

W N

Am

W-N

T3

16-2

3,50

-55,

00-3

9,25

31,5

0

Om

brop

hytu

mE

udic

ots

San

tala

les

Bal

anop

hora

ceae

4P

eru,

W B

rasi

l, N

Arg

, Chi

leA

ND

31

-18,

20-2

2,12

-20,

163,

92

Onu

risR

osid

sB

rass

ical

esB

rass

icac

eae

5C

hile

, Pat

agon

ia5

5-3

7,58

-54,

67-4

6,13

17,0

8

Oph

iogl

ossu

mFi

licop

sida

Oph

iogl

ossa

les

Oph

iglo

ssac

eae

26S

ubco

smop

, Mad

ag, J

F7

3-3

3,10

-54,

83-4

3,97

21,7

3

Oph

ryos

poru

sA

ster

ids

Ast

eral

esA

ster

acea

e37

trop

& s

ubtro

p S

Am

, Bra

zil

S-A

MZ

35

-17,

58-3

6,67

-27,

1319

,08

Ore

obol

usM

onoc

ots

Poa

les

Cyp

erac

eae

15M

ales

ia, A

us, T

asm

, NZ,

Haw

aii,

JF, t

rop

& te

mp

Am

, C

hile

, Tfu

ego,

Fal

klan

dS

PT

21

-36,

50-5

6,00

-46,

2519

,50

Ore

ocer

eus

Eud

icot

sC

aryo

phyl

lale

sC

acta

ceae

7A

ndes

Per

u, B

ol, C

hile

, Arg

AN

D3

3-1

8,17

-22,

50-2

0,33

4,33

Ore

omyr

rhis

Ast

erid

sA

pial

esA

piac

eae

23E

Asi

a, B

orne

o, N

Gui

nea,

SE

Aus

, NZ,

Mex

, C A

m,

And

es, T

fueg

o, F

alkl

and

SP

T2

1-5

2,00

-54,

00-5

3,00

2,00

Ore

opol

usA

ster

ids

Gen

tiana

les

Rub

iace

ae1

Chi

le, A

rg, P

atag

onia

51

-33,

00-5

4,50

-43,

7521

,50

Orit

esE

udic

ots

Pro

teal

esP

rote

acea

e9

6 te

mp

E A

us, 3

And

es S

Am

AU

S2

1-3

5,83

-38,

25-3

7,04

2,42

Oro

banc

heA

ster

ids

Lam

iale

sO

roba

ncha

ceae

130

tem

p, s

ubtro

p, N

Hem

is7

2-1

8,00

-33,

50-2

5,75

15,5

0

Ort

achn

eM

onoc

ots

Poa

les

Poa

ceae

2A

rg, C

hile

52

-39,

48-5

6,00

-47,

7416

,52

Osm

orhi

zaA

ster

ids

Api

ales

Api

acea

e10

N A

m, E

Asi

a, te

mp

S A

mW

-AN

T4

3-3

0,83

-54,

93-4

2,88

24,1

0

Oth

olob

ium

Ros

ids

Faba

les

Faba

ceae

36E

& S

Afr,

tem

p S

Am

12

-29,

83-4

0,00

-34,

9210

,17

Our

isia

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

25Ta

sm, N

Z, A

ndes

S A

mA

US

213

-18,

23-5

6,00

-37,

1237

,77

Ovi

dia

Ros

ids

Mal

vale

sTh

ymel

aeac

eae

2te

mp

S A

m5

2-3

7,80

-45,

93-4

1,87

8,13

Oxa

lisR

osid

sO

xalid

ales

Oxa

lidac

eae

600

Cos

mop

711

7-1

8,00

-56,

00-3

7,00

38,0

0

Oxy

chlo

ë M

onoc

ots

Poa

les

Junc

acea

e6

And

es S

Am

AN

D3

1-1

8,00

-34,

25-2

6,13

16,2

5

Oxy

phyl

lum

Ast

erid

sA

ster

ales

Ast

erac

eae

1N

Chi

le6

1-2

4,83

-26,

15-2

5,49

1,32

Oxy

thec

aE

udic

ots

Car

yoph

ylla

les

Pol

ygon

acea

e7

W N

Am

, Chi

le, A

rgA

NT-

P4

1-2

4,00

-31,

50-2

7,75

7,50

��8

Ozi

roe

Mon

ocot

sA

spar

agal

esH

yaci

ntha

ceae

5C

hile

, Arg

, Per

ú, B

ol, P

arag

uay

AN

D3

4-1

8,40

-36,

00-2

7,20

17,6

0

Pach

ylae

naA

ster

ids

Ast

eral

esA

ster

acea

e2

And

es N

Chi

le A

rgA

ND

32

-28,

00-3

5,80

-31,

907,

80

Pala

uaR

osid

sM

alva

les

Mal

vace

ae15

Coa

stal

Per

u an

d C

hile

AN

D3

4-2

0,38

-25,

20-2

2,79

4,82

Pani

cum

Mon

ocot

sP

oale

sP

oace

ae60

0te

mp,

sub

trop,

trop

regs

72

-33,

50-3

7,67

-35,

584,

17

Para

stre

phia

Ast

erid

sA

ster

ales

Ast

erac

eae

5A

ndes

Bol

, Chi

le, P

erú,

Arg

AN

D3

5-1

7,58

-25,

00-2

1,29

7,42

Parie

taria

Ros

ids

Ros

ales

Urti

cace

ae15

subc

osm

op, J

F, ID

72

-20,

80-3

5,62

-28,

2114

,82

Paro

nych

iaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

100

Nor

th te

mpe

rate

, aus

tral

W-A

NT

45

-18,

18-3

7,80

-27,

9919

,62

Pasi

thea

Mon

ocot

sA

spar

agal

esH

emer

ocal

lidac

eae

1P

eru,

Chi

leA

ND

31

-22,

00-4

0,00

-31,

0018

,00

Pasp

alum

Mon

ocot

sP

oale

sP

oace

ae33

0te

mp

& tr

op re

gs, I

P7

5-2

0,50

-42,

50-3

1,50

22,0

0

Passiflora

Ros

ids

Mal

pigh

iale

sP

assi

flora

ceae

500

trop

& s

ubtro

p A

m, 1

Mad

ag, 2

0 tro

p E

Asi

a, M

ales

ia,

3 A

us, N

Z1

1-3

1,97

-32,

50-3

2,23

0,53

Pato

sia

Mon

ocot

sP

oale

sJu

ncac

eae

1A

ndes

Chi

le A

rg B

olA

ND

31

-24,

00-3

8,00

-31,

0014

,00

Pect

ocar

yaA

ster

ids

Unp

lace

dB

orag

inac

eae

13W

N A

m, S

Am

AN

T-P

44

-23,

58-3

5,97

-29,

7812

,38

Pella

eaFi

licop

sida

Pte

ridal

esP

terid

acea

e35

trop

& w

arm

-tem

p, M

adag

12

-18,

27-4

0,32

-29,

2922

,05

Pelle

tiera

Ast

erid

sE

rical

esM

yrsi

nace

ae2

1 su

btro

p S

Am

, 1 M

acar

ones

ia1

1-3

2,50

-40,

00-3

6,25

7,50

Penn

iset

umM

onoc

ots

Poa

les

Poa

ceae

115

trop

& te

mp

regs

W-A

NT

41

-20,

00-3

3,53

-26,

7713

,53

Pepe

rom

iaM

agno

liids

Pip

eral

esP

iper

acea

e10

00tro

p &

war

m e

sp A

m, J

F, H

awai

i1

4-2

4,83

-42,

50-3

3,67

17,6

7

Pere

zia

Ast

erid

sA

ster

ales

Ast

erac

eae

32A

ndes

from

Col

ombi

a to

Chi

le, S

E B

razi

lA

ND

323

-17,

58-5

6,00

-36,

7938

,42

Perit

yle

Ast

erid

sA

ster

ales

Ast

erac

eae

63S

W N

Am

, 1 C

hile

Per

uA

NT-

P4

1-2

0,80

-29,

90-2

5,35

9,10

Pers

eaM

agno

liids

Laur

ales

Laur

acea

e12

0tro

ps &

sub

trops

Am

& A

sia,

Mac

aron

esia

W-A

NT

42

-33,

08-4

3,18

-38,

1310

,10

Peum

usM

agno

liids

Laur

ales

Mon

imia

ceae

1C

hile

61

-30,

67-4

0,50

-35,

589,

83

Phac

elia

Ast

erid

sU

npla

ced

Bor

agin

acea

e20

0W

N A

m, S

Am

AN

T-P

46

-18,

00-5

4,83

-36,

4236

,83

Phal

aris

Mon

ocot

sP

oale

sP

oace

ae20

tem

pera

te E

uras

ia, A

mer

ica

W-A

NT

42

-30,

83-3

9,88

-35,

369,

05

Phile

sia

Mon

ocot

sLi

liale

sP

hile

siac

eae

1C

hile

, Arg

51

-39,

83-5

5,50

-47,

6715

,67

Phili

bert

iaA

ster

ids

Gen

tiana

les

Apo

cyna

ceae

40A

ndes

Per

ú, C

hile

, Bol

ivia

, Arg

entin

aA

ND

31

-18,

33-2

2,27

-20,

303,

93

Phili

ppie

llaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

1P

atag

onia

Chi

le-A

rg.

51

-48,

00-5

0,00

-49,

002,

00

Phle

umM

onoc

ots

Poa

les

Poa

ceae

15te

mpe

rate

Eur

asia

, Am

eric

aW

-AN

T4

1-2

9,50

-55,

17-4

2,33

25,6

7

Phra

gmite

sM

onoc

ots

Poa

les

Poa

ceae

4C

osm

op7

1-1

9,67

-40,

00-2

9,83

20,3

3

Phro

dus

Ast

erid

sS

olan

ales

Sol

anac

eae

1C

hile

(Ata

cam

a &

Coq

uim

bo)

61

-26,

43-3

1,33

-28,

884,

90

Phyc

ella

Mon

ocot

sA

spar

agal

esA

mar

yllid

acea

e3

Chi

le, A

rgA

ND

33

-29,

67-3

6,62

-33,

146,

95

Phyl

lach

neA

ster

ids

Ast

eral

esS

tylid

iace

ae4

Tasm

, NZ,

tem

p S

Am

AU

S2

1-4

5,00

-56,

00-5

0,50

11,0

0

Phyl

losc

irpus

Mon

ocot

sP

oale

sC

yper

acea

e2

Arg

, Chi

le5

2-2

2,00

-51,

00-3

6,50

29,0

0

Phyt

olac

caE

udic

ots

Car

yoph

ylla

les

Phy

tola

ccac

eae

25tro

p &

war

m1

1-3

7,13

-41,

88-3

9,51

4,75

Picr

osia

Ast

erid

sA

ster

ales

Ast

erac

eae

2su

btro

p S

Am

, Chi

le, P

eru,

Arg

, Bra

zil

S-A

MZ

31

-34,

50-3

6,72

-35,

612,

22

Pile

aR

osid

sR

osal

esU

rtica

ceae

250

trop

& w

arm

exc

l Aus

trala

sia

12

-37,

62-4

5,42

-41,

527,

80

Pilg

erod

endr

onP

inop

sida

Pin

ales

Cup

ress

acea

e1

Chi

le A

rg5

1-3

9,90

-54,

25-4

7,08

14,3

5

Pilo

styl

esR

osid

sM

alpi

ghia

les

Raf

flesi

acea

e25

Am

US

to M

agal

lane

s, A

sia,

2 W

Aus

11

-18,

18-3

6,90

-27,

5418

,72

��9

Pilu

laria

Filic

opsi

daM

arsi

leal

esM

arsi

leac

eae

6E

uro,

Aus

, NZ,

Eth

iopi

a, W

S A

m7

1-3

2,15

-40,

28-3

6,22

8,13

Ping

uicu

laA

ster

ids

Lam

iale

sLe

ntib

ular

iace

ae85

Am

, Med

it, c

ircum

bore

alW

-AN

T4

2-3

5,50

-56,

00-4

5,75

20,5

0

Pint

oaR

osid

sZy

goph

ylla

les

Zygo

phyl

lace

ae1

Chi

le6

1-2

7,33

-28,

88-2

8,11

1,55

Pipt

ocha

etiu

mM

onoc

ots

Poa

les

Poa

ceae

30A

m U

S to

Arg

, JF

W-N

T3

7-2

9,50

-40,

00-3

4,75

10,5

0

Pita

via

Ros

ids

Sap

inda

les

Rut

acea

e1

Chi

le6

1-3

5,33

-37,

53-3

6,43

2,20

Pitr

aea

Ast

erid

sLa

mia

les

Verb

enac

eae

1S

Am

, Chi

le, P

eru,

Arg

AN

D3

1-1

8,47

-33,

83-2

6,15

15,3

7

Pity

rogr

amm

aFi

licop

sida

Pte

ridal

esP

terid

acea

e16

Am

trop

des

de S

US

, Afr,

Mad

ag1

1-1

8,50

-18,

67-1

8,58

0,17

Plac

eaM

onoc

ots

Asp

arag

ales

Am

aryl

lidac

eae

5C

hile

65

-31,

00-3

4,55

-32,

783,

55

Plag

iobo

thry

sA

ster

ids

Unp

lace

dB

orag

inac

eae

70W

N A

m, S

outh

Am

, E A

sia,

Aus

C-P

AC

419

-29,

83-5

4,00

-41,

9224

,17

Plan

tago

Ast

erid

sLa

mia

les

Pla

ntag

inac

eae

270

Cos

mop

, JF,

ID7

18-1

8,17

-56,

00-3

7,08

37,8

3

Plaz

iaA

ster

ids

Ast

eral

esA

ster

acea

e3

Per

u, B

ol, A

rg, C

hile

AN

D3

2-1

8,18

-21,

95-2

0,07

3,77

Plec

triti

sA

ster

ids

Dip

saca

les

Vale

riana

ceae

54

W N

Am

, 1 C

hile

AN

T-P

41

-32,

50-3

7,33

-34,

924,

83

Pleo

carp

hus

Ast

erid

sA

ster

ales

Ast

erac

eae

1N

Chi

le6

1-2

9,15

-33,

50-3

1,33

4,35

Pleo

pelti

sFi

licop

sida

Pol

ypod

iale

sP

olyp

odia

ceae

10tro

p A

m, 1

Afr-

Mad

ag, I

ndia

, Sri

Lank

a1

1-2

4,00

-42,

40-3

3,20

18,4

0

Pleu

roph

ora

Ros

ids

Myr

tale

sLy

thra

ceae

9S

Am

, Bra

zil

S-A

MZ

33

-25,

00-3

4,42

-29,

719,

42

Pleu

roso

rus

Filic

opsi

daA

sple

nial

esA

sple

niac

eae

4E

spañ

a, M

arru

ecos

, Aus

, Tas

m, N

Z, C

hile

, Arg

71

-32,

25-4

5,58

-38,

9213

,33

Pluc

hea

Ast

erid

sA

ster

ales

Ast

erac

eae

80A

sia,

N &

S A

m, A

fr, A

usW

-AN

T4

1-1

8,42

-19,

12-1

8,77

0,70

Plum

bago

Eud

icot

sC

aryo

phyl

lale

sP

lum

bagi

nace

ae20

trop

& w

arm

-tem

p re

gs7

1-2

4,75

-31,

93-2

8,34

7,18

Poa

Mon

ocot

sP

oale

sP

oace

ae35

0te

mp

& c

old

& m

onta

ne tr

ops

W-A

NT

461

-18,

00-5

6,00

-37,

0038

,00

Poda

gros

tisM

onoc

ots

Poa

les

Poa

ceae

4A

rg, C

hile

, Can

adá,

US

AA

NT-

P4

1-3

5,25

-40,

33-3

7,79

5,08

Poda

nthu

sA

ster

ids

Ast

eral

esA

ster

acea

e2

Chi

le6

2-3

0,28

-38,

25-3

4,27

7,97

Podo

carp

usP

inop

sida

Pin

ales

Pod

ocar

pace

ae10

0A

fr, M

adag

, Asi

a, A

us, T

asm

, NZ,

Mex

, C &

S A

m, W

I1

2-3

5,83

-50,

38-4

3,11

14,5

5

Podo

phor

usM

onoc

ots

Poa

les

Poa

ceae

1JF

61

0,00

Pole

mon

ium

Ast

erid

sE

rical

esP

olem

onia

ceae

40te

mp

Eur

asia

, N A

m, C

hile

W-A

NT

41

-30,

83-5

4,00

-42,

4223

,17

Poly

achy

rus

Ast

erid

sA

ster

ales

Ast

erac

eae

7P

eru,

Chi

leA

ND

37

-17,

58-3

4,73

-26,

1617

,15

Poly

carp

onE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

16C

osm

op7

1-3

0,00

-30,

00-3

0,00

0,00

Poly

gala

Ros

ids

Faba

les

Pol

ygal

acea

e50

0su

bcos

mop

78

-29,

00-5

4,00

-41,

5025

,00

Poly

gonu

mE

udic

ots

Car

yoph

ylla

les

Pol

ygon

acea

e90

Cos

mop

71

-30,

00-4

6,75

-38,

3816

,75

Poly

lepi

sR

osid

sR

osal

esR

osac

eae

15A

ndes

Ven

ezue

la to

Arg

, Chi

leA

ND

32

-17,

72-2

2,00

-19,

864,

28

Poly

podi

umFi

licop

sida

Pol

ypod

iale

sP

olyp

odia

ceae

150

Cos

mop

, JF

73

-23,

52-5

2,88

-38,

2029

,37

Poly

pogo

nM

onoc

ots

Poa

les

Poa

ceae

13C

osm

op7

4-1

8,00

-46,

67-3

2,33

28,6

7

Poly

stic

hum

Filic

opsi

daD

ryop

terid

ales

Dry

opte

ridac

eae

160

Cos

mop

, Mad

ag, I

P7

5-3

1,30

-55,

00-4

3,15

23,7

0

Porli

eria

Ros

ids

Zygo

phyl

lale

sZy

goph

ylla

ceae

5S

US

A, M

exic

o, S

Am

AN

T-P

41

-30,

00-3

4,28

-32,

144,

28

Port

ulac

aE

udic

ots

Car

yoph

ylla

les

Por

tula

cace

ae40

trop,

sub

trop

& te

mp

regi

ons

72

-18,

33-2

8,80

-23,

5710

,47

Pota

mog

eton

Mon

ocot

sA

lism

atal

esP

otam

oget

onac

eae

95C

osm

op7

6-2

1,50

-55,

00-3

8,25

33,5

0

Pout

eria

Ast

erid

sE

rical

esS

apot

acea

e20

0tro

p. A

m, A

fr, A

sia,

Aus

, NZ,

New

Cal

ed1

1-3

2,10

-33,

03-3

2,57

0,93

�40

Pozo

aA

ster

ids

Api

ales

Api

acea

e2

And

es C

hile

Arg

52

-31,

50-3

8,58

-35,

047,

08

Prat

iaA

ster

ids

Ast

eral

esC

ampa

nula

ceae

15S

E A

sia

tropi

cal,

Aus

, NZ,

Am

Aus

tral

SP

T2

1-1

8,83

-56,

00-3

7,42

37,1

7

Prim

ula

Ast

erid

sE

rical

esP

rimul

acea

e40

0N

Hem

isp,

trop

Asi

a, te

mp

S A

mW

-AN

T4

1-3

8,50

-55,

83-4

7,17

17,3

3

Pros

opis

Ros

ids

Faba

les

Faba

ceae

44tro

p &

sub

trop

Am

, Afr,

W A

sia

16

-19,

00-3

4,33

-26,

6715

,33

Prou

stia

Ast

erid

sA

ster

ales

Ast

erac

eae

3B

ol, C

hile

, Arg

, Per

uA

ND

33

-18,

00-4

0,33

-29,

1722

,33

Prum

nopi

tys

Pin

opsi

daP

inal

esP

odoc

arpa

ceae

8C

osta

Ric

a to

Chi

le, N

ew C

aled

, NZ,

E A

usA

US

21

-35,

83-3

9,50

-37,

673,

67

Pseu

dopa

nax

Ast

erid

sA

pial

esA

ralia

ceae

10Ta

sm, N

Z, C

hile

AU

S2

2-3

5,33

-52,

33-4

3,83

17,0

0

Psilo

carp

hus

Ast

erid

sA

ster

ales

Ast

erac

eae

8W

US

, tem

p S

Am

AN

T-P

42

-30,

00-3

8,00

-34,

008,

00

Psilo

tum

Psi

loto

psid

aP

silo

tale

sP

silo

tace

ae2

pant

rop,

Haw

aii,

Aus

tralia

, NZ,

Mad

ag, I

P1

10,

00

Pter

isFi

licop

sida

Pte

ridal

esP

terid

acea

e28

0C

osm

op, M

adag

, JF

72

-32,

55-4

9,70

-41,

1317

,15

Pter

ocac

tus

Eud

icot

sC

aryo

phyl

lale

sC

acta

ceae

9P

atag

onia

Arg

, Chi

le5

1-4

6,50

-46,

50-4

6,50

0,00

Pter

omon

nina

Ros

ids

Faba

les

Pol

ygal

acea

e13

0A

m S

W U

S to

Chi

leW

-NT

34

-29,

00-3

7,83

-33,

428,

83

Pucc

inel

liaM

onoc

ots

Poa

les

Poa

ceae

80N

tem

p: A

sia,

N A

m, S

tem

p: S

Afr,

Aus

, S A

m7

8-2

0,35

-56,

00-3

8,18

35,6

5

Puya

Mon

ocot

sP

oale

sB

rom

elia

ceae

190

C &

S A

m, B

razi

lW

-NT

38

-22,

50-3

7,80

-30,

1515

,30

Pycn

ophy

llum

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae17

And

es P

eru,

Chi

le, A

rg, B

olA

ND

35

-18,

17-2

4,73

-21,

456,

57

Pycr

eus

Mon

ocot

sP

oale

sC

yper

acea

e70

pant

rops

11

0,00

Qui

llaja

Ros

ids

Faba

les

Qui

llaja

ceae

3te

mp

S A

m, A

rg, C

hile

, Per

u, B

razi

lS

-AM

Z3

1-3

0,00

-38,

23-3

4,12

8,23

Qui

ncha

mal

ium

Eud

icot

sS

anta

lale

sS

anta

lace

ae25

And

es B

ol, C

hile

, Per

ú, A

rgA

ND

316

-18,

18-5

1,00

-34,

5932

,82

Ran

uncu

lus

Eud

icot

sR

anun

cula

les

Ran

uncu

lace

ae50

0C

osm

op, R

aca

ulis

dis

j NZ,

Fal

klan

d, R

bon

arie

nsis

di

sj N

Am

S A

m7

17-1

7,78

-54,

92-3

6,35

37,1

3

Rei

chee

llaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

1C

hile

61

-18,

00-2

2,58

-20,

294,

58

Rel

chel

aM

onoc

ots

Poa

les

Poa

ceae

1C

hile

Arg

51

-35,

85-4

2,08

-38,

976,

23

Ret

anill

aR

osid

sR

osal

esR

ham

nace

ae3

Chi

le, A

rg5

2-3

1,62

-37,

80-3

4,71

6,18

Rey

esia

Ast

erid

sS

olan

ales

Sol

anac

eae

4C

hile

Arg

AN

D3

4-1

9,10

-30,

20-2

4,65

11,1

0

Rha

mnu

sR

osid

sR

osal

esR

ham

nace

ae16

0N

Hem

isp,

S A

fr, S

Am

W-A

NT

41

-35,

83-4

1,00

-38,

425,

17

Rha

phith

amnu

sA

ster

ids

Lam

iale

sVe

rben

acea

e2

Chi

le, A

rg, J

F5

1-3

0,67

-45,

42-3

8,04

14,7

5

Rho

doph

iala

Mon

ocot

sA

spar

agal

esA

mar

yllid

acea

e30

Chi

le, A

rg, B

ol, B

ras

S-A

MZ

39

-25,

00-4

0,00

-32,

5015

,00

Rho

mbo

lytr

umM

onoc

ots

Poa

les

Poa

ceae

2S

Bra

sil,

Uru

g, C

hile

S-A

MZ

31

-33,

00-3

5,62

-34,

312,

62

Rhy

ncho

spor

aM

onoc

ots

Poa

les

Cyp

erac

eae

250

cosm

p7

1-3

7,83

-39,

23-3

8,53

1,40

Rib

esE

udic

ots

Sax

ifrag

ales

Gro

ssul

aria

ceae

150

N H

emis

, N A

fr, S

Am

Can

ada

to T

fueg

oW

-AN

T4

13-2

9,83

-56,

00-4

2,92

26,1

7

Rob

inso

nia

Ast

erid

sA

ster

ales

Ast

erac

eae

7JF

66

0,00

Ror

ippa

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e75

cosm

op7

6-3

3,60

-43,

83-3

8,72

10,2

3

Ros

tkov

iaM

onoc

ots

Poa

les

Junc

acea

e2

NZ,

S S

Am

, Tfu

ego,

Fal

klan

dA

US

21

-46,

43-5

6,00

-51,

229,

57

Ros

trar

iaM

onoc

ots

Poa

les

Poa

ceae

2A

rg, C

arib

e, C

hil,

Mex

ico,

Per

u, U

SA

,Uru

gW

-NT

31

-25,

00-2

6,38

-25,

691,

38

Rub

usR

osid

sR

osal

esR

osac

eae

500

cosm

op, J

F7

2-3

6,90

-55,

67-4

6,28

18,7

7

Rum

exE

udic

ots

Car

yoph

ylla

les

Pol

ygon

acea

e20

0co

smop

710

-30,

00-5

5,00

-42,

5025

,00

Rum

ohra

Filic

opsi

daD

ryop

terid

ales

Dry

opte

ridac

eae

7ci

rcum

aust

ral,

J Fe

rnan

dez,

Mad

agC

-AU

S2

1-3

0,67

-55,

08-4

2,88

24,4

2

�4�

Rup

pia

Mon

ocot

sA

lism

atal

esR

uppi

acea

e10

cosm

op7

2-2

1,00

-55,

00-3

8,00

34,0

0

Ryt

idos

perm

aM

onoc

ots

Poa

les

Poa

ceae

28A

ustra

lia, N

Z, A

rg, C

hile

, IP

AU

S2

6-3

0,67

-51,

00-4

0,83

20,3

3

Sagi

naE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

25te

mp

N H

emis

, E A

fr, A

sia,

N G

uine

a, A

us, N

Z, A

ndes

71

-31,

50-4

2,50

-37,

0011

,00

Sagi

ttaria

Mon

ocot

sA

lism

atal

esA

lism

atac

eae

25C

osm

op7

1-3

6,50

-40,

00-3

8,25

3,50

Salix

Ros

ids

Mal

pigh

iale

sS

alic

acea

e45

0A

rctic

tem

p, fe

w in

trop

s, te

mp

S A

m, 1

S A

frW

-AN

T4

1-1

8,58

-38,

78-2

8,68

20,2

0

Salp

ichr

oaA

ster

ids

Sol

anal

esS

olan

acea

e15

S A

m: E

cuad

, Per

u, B

ol, B

ra, P

arag

, Uru

, Chi

le, A

rgA

ND

32

-18,

25-1

9,87

-19,

061,

62

Salp

iglo

ssis

Ast

erid

sS

olan

ales

Sol

anac

eae

2C

hile

, Arg

52

-27,

00-4

0,00

-33,

5013

,00

Salv

iaA

ster

ids

Lam

iale

sLa

mia

ceae

900

cosm

op7

4-2

2,00

-25,

33-2

3,67

3,33

Sam

olus

Ast

erid

sE

rical

esS

amol

acea

e12

Cos

mop

, IP

73

-33,

42-5

5,00

-44,

2121

,58

Sanc

tam

bros

iaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

1D

esve

ntur

adas

Isla

nds

(Chi

le)

61

0,00

Sani

cula

Ast

erid

sA

pial

esA

piac

eae

40su

bcos

mop

, exc

Aus

& N

Z7

2-3

0,67

-44,

50-3

7,58

13,8

3

Sant

alum

Eud

icot

sS

anta

lale

sS

anta

lace

ae25

Indo

mal

asia

to A

us, H

awai

i, JF

(ex)

SP

T2

10,

00

Sarc

ocor

nia

Eud

icot

sC

aryo

phyl

lale

sA

mar

anth

acea

e16

Eur

asia

, Afr,

Aus

, Tas

m, N

Z, N

& S

Am

, JF

73

-20,

63-5

4,00

-37,

3233

,37

Sarm

ient

aA

ster

ids

Lam

iale

sG

esne

riace

ae1

Chi

le6

1-3

0,60

-43,

83-3

7,22

13,2

3

Satu

reja

Ast

erid

sLa

mia

les

Lam

iace

ae38

tem

p &

sub

trop,

Med

it7

4-1

8,50

-54,

00-3

6,25

35,5

0

Saxe

-got

haea

Pin

opsi

daP

inal

esP

odoc

arpa

ceae

1C

hile

, Arg

51

-36,

00-4

5,75

-40,

889,

75

Saxi

frag

aE

udic

ots

Sax

ifrag

ales

Sax

ifrag

acea

e35

0N

tem

p, E

uras

ia, W

N A

m, A

ndes

S A

mW

-AN

T4

1-3

6,00

-55,

00-4

5,50

19,0

0

Saxi

frag

ella

Eud

icot

sS

axifr

agal

esS

axifr

agac

eae

1A

ntar

ctic

S A

m5

1-5

0,72

-56,

00-5

3,36

5,28

Saxi

frag

odes

Eud

icot

sS

axifr

agal

esS

axifr

agac

eae

1Tf

uego

51

-51,

00-5

4,83

-52,

923,

83

Schi

nus

Ros

ids

Sap

inda

les

Ana

card

iace

ae27

Mex

ico

to A

rgW

-NT

39

-18,

00-4

6,55

-32,

2828

,55

Schi

zaea

Filic

opsi

daS

chiz

aeal

esS

chiz

aece

ae30

trop

& a

ustra

l, N

Am

, Mad

ag1

1-3

9,88

-49,

15-4

4,52

9,27

Schi

zant

hus

Ast

erid

sS

olan

ales

Sol

anac

eae

12C

hile

, Arg

AN

D3

12-2

1,03

-41,

00-3

1,02

19,9

7

Schi

zeile

ma

Ast

erid

sA

pial

esA

piac

eae

1311

NZ,

1 A

us, 1

S S

Am

AU

S2

1-3

8,58

-54,

48-4

6,53

15,9

0

Schi

zope

talo

nR

osid

sB

rass

ical

esB

rass

icac

eae

10C

N C

hile

, Arg

AN

D3

10-2

6,43

-35,

33-3

0,88

8,90

Schk

uhria

Ast

erid

sA

ster

ales

Ast

erac

eae

6tro

p &

sub

trop

Am

W-N

T3

2-1

7,58

-31,

83-2

4,71

14,2

5

Scho

enus

Mon

ocot

sP

oale

sC

yper

acea

e10

0su

bcos

mop

74

-35,

83-5

6,00

-45,

9220

,17

Scirp

usM

onoc

ots

Poa

les

Cyp

erac

eae

20N

orth

tem

pera

te, a

ustra

l, JF

, IP

720

-17,

67-5

6,00

-36,

8338

,33

Scut

ella

riaA

ster

ids

Lam

iale

sLa

mia

ceae

360

cosm

op7

3-3

5,33

-53,

00-4

4,17

17,6

7

Scyp

hant

hus

Ast

erid

sC

orna

les

Loas

acea

e2

Chi

le6

2-3

1,00

-38,

23-3

4,62

7,23

Selk

irkia

Ast

erid

sU

npla

ced

Bor

agin

acea

e1

JF6

10,

00

Selli

era

Ast

erid

sA

ster

ales

Goo

deni

acea

e2

Aus

, NZ,

Chi

leA

US

21

-28,

45-4

2,72

-35,

5814

,27

Sene

cio

Ast

erid

sA

ster

ales

Ast

erac

eae

1250

Cos

mop

721

8-1

7,58

-56,

00-3

6,79

38,4

2

Senn

aR

osid

sFa

bale

sFa

bace

ae30

0pa

ntro

ps1

9-1

8,00

-40,

00-2

9,00

22,0

0

Serp

yllo

psis

Filic

opsi

daH

ymen

ophy

lla-

les

Hym

enop

hylla

ceae

1S

S A

m, J

F, F

alkl

and

51

-39,

63-5

6,00

-47,

8216

,37

Sicy

osR

osid

sC

ucur

bita

les

Cuc

urbi

tace

ae50

Aus

, Tas

m, N

Z, H

awai

i, N

& S

Am

, ID

C-P

AC

41

-20,

17-3

3,40

-26,

7813

,23

Sige

sbec

kia

Ast

erid

sA

ster

ales

Ast

erac

eae

3su

btro

& tr

op re

gs1

1-3

2,83

-42,

72-3

7,78

9,88

�4�

Sile

neE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

600

tem

p N

hem

is, M

edit,

S A

fr7

8-1

8,00

-55,

00-3

6,50

37,0

0

Sisy

mbr

ium

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e90

tem

p E

uras

ia, M

edit,

S A

fr, A

ndes

, N A

mW

-AN

T4

23-1

8,20

-55,

00-3

6,60

36,8

0

Sisy

rinch

ium

Mon

ocot

sA

spar

agal

esIri

dace

ae60

N, C

& S

Am

W-N

T3

11-2

4,50

-55,

25-3

9,88

30,7

5

Skyt

anth

usA

ster

ids

Gen

tiana

les

Apo

cyna

ceae

3B

rasi

l, C

hile

S-A

MZ

31

-25,

20-2

8,93

-27,

073,

73

Sola

num

Ast

erid

sS

olan

ales

Sol

anac

eae

1700

subc

osm

op, J

F, ID

, IP

751

-18,

20-4

6,50

-32,

3528

,30

Sola

riaM

onoc

ots

Asp

arag

ales

Alli

acea

e2

Chi

le, A

rgA

ND

31

-33,

00-3

6,83

-34,

923,

83

Sole

nom

elus

Mon

ocot

sA

spar

agal

esIri

dace

ae2

Chi

le, A

rg5

2-3

1,00

-50,

00-4

0,50

19,0

0

Solid

ago

Ast

erid

sA

ster

ales

Ast

erac

eae

150

Tem

pera

te A

mer

ica,

Mac

aron

esia

, Eur

asia

W-A

NT

42

-30,

95-4

4,00

-37,

4813

,05

Soliv

aA

ster

ids

Ast

eral

esA

ster

acea

e8

S A

m, N

Am

, Aus

C-P

AC

44

-29,

95-4

1,83

-35,

8911

,88

Soph

ora

Ros

ids

Faba

les

Faba

ceae

47tro

p &

mos

t N te

mp,

JF,

IP7

2-3

2,00

-45,

42-3

8,71

13,4

2

Spar

tina

Mon

ocot

sP

oale

sP

oace

ae20

Am

, W E

uro,

NW

Afr

W-A

NT

41

-36,

67-3

7,50

-37,

080,

83

Spee

aM

onoc

ots

Asp

arag

ales

Alli

acea

e1

Chi

le6

2-3

2,92

-34,

27-3

3,59

1,35

Sper

gula

riaE

udic

ots

Car

yoph

ylla

les

Car

yoph

ylla

ceae

30co

smop

, JF,

ID7

13-1

8,00

-40,

00-2

9,00

22,0

0

Spha

eral

cea

Ros

ids

Mal

vale

sM

alva

ceae

60W

N A

m, M

ex, t

empe

rate

S A

m

AN

T-P

42

-29,

45-3

7,78

-33,

628,

33

Spila

nthe

sA

ster

ids

Ast

eral

esA

ster

acea

e6

trops

11

-18,

47-1

8,47

-18,

470,

00

Spor

obol

usM

onoc

ots

Poa

les

Poa

ceae

160

Am

, Asi

a, A

fr, 1

Eur

o7

1-2

3,62

-23,

62-2

3,62

0,00

Stac

hys

Ast

erid

sLa

mia

les

Lam

iace

ae30

0tro

p, s

ubtro

p, te

mp

regi

ons

exc.

Aus

trala

sia

711

-22,

00-4

4,00

-33,

0022

,00

Stan

gea

Ast

erid

sD

ipsa

cale

sVa

leria

nace

ae5

Arg

,Chi

l,Per

úA

ND

31

-17,

67-1

8,08

-17,

880,

42

Stel

laria

Eud

icot

sC

aryo

phyl

lale

sC

aryo

phyl

lace

ae15

0co

smop

75

-18,

00-5

5,83

-36,

9237

,83

Stem

odia

Ast

erid

sLa

mia

les

Gra

tiola

ceae

45Tr

ops

and

Sub

trops

, Mad

agW

-AN

T4

1-2

9,00

-38,

00-3

3,50

9,00

Sten

andr

ium

Ast

erid

sLa

mia

les

Aca

ntha

ceae

40S

US

to C

Chi

leW

-NT

31

-30,

00-3

7,80

-33,

907,

80

Sten

omes

son

Mon

ocot

sA

spar

agal

esA

mar

yllid

acea

e40

And

esA

ND

31

-18,

00-1

8,17

-18,

080,

17

Stev

iaA

ster

ids

Ast

eral

esA

ster

acea

e23

0S

US

to C

hile

W-N

T3

2-1

7,58

-25,

43-2

1,51

7,85

Stip

aM

onoc

ots

Poa

les

Poa

ceae

300

tem

p N

Hem

is, m

onta

ne tr

ops,

IP7

10,

00

Stuc

keni

aM

onoc

ots

Alis

mat

ales

Pot

amog

eton

acea

e6

Sub

cosm

op7

3-1

8,00

-53,

00-3

5,50

35,0

0

Suae

daE

udic

ots

Car

yoph

ylla

les

Am

aran

thac

eae

100

cosm

op, I

D7

4-2

0,25

-55,

00-3

7,63

34,7

5

Tage

tes

Ast

erid

sA

ster

ales

Ast

erac

eae

50N

& S

Am

W-N

T3

3-1

7,58

-36,

60-2

7,09

19,0

2

Tape

inia

Mon

ocot

sA

spar

agal

esIri

dace

ae1

Chi

le, A

rg5

1-4

0,50

-56,

00-4

8,25

15,5

0

Tara

Ros

ids

Faba

les

Faba

ceae

3M

exic

o, C

Am

to S

Am

W-N

T3

1-1

8,47

-18,

47-1

8,47

0,00

Tara

saR

osid

sM

alva

les

Mal

vace

ae30

2 sp

Mex

ico,

dis

junc

t to

Per

u, C

hile

, Arg

AN

T-P

410

-18,

17-3

8,78

-28,

4820

,62

Tara

xacu

mA

ster

ids

Ast

eral

esA

ster

acea

e60

tem

p N

Hem

is &

tem

p S

Am

, NZ,

JF

W-A

NT

42

-33,

45-5

6,00

-44,

7322

,55

Teco

ma

Ast

erid

sLa

mia

les

Big

noni

acea

e12

Am

trop

ical

Ariz

ona

to C

hile

, 1 S

Afr,

Aus

?1

1-1

8,47

-18,

50-1

8,48

0,03

Teco

phila

eaM

onoc

ots

Asp

arag

ales

Teco

phila

eace

ae2

Chi

le6

2-3

0,00

-33,

00-3

1,50

3,00

Tepu

alia

Ros

ids

Myr

tale

sM

yrta

ceae

1C

hile

, Arg

51

-35,

40-5

3,00

-44,

2017

,60

Tess

aria

Ast

erid

sA

ster

ales

Ast

erac

eae

1S

Am

, Bra

zil

S-A

MZ

31

-18,

42-3

6,50

-27,

4618

,08

Tetil

laR

osid

sG

eran

iale

sFr

anco

acea

e1

Chi

le6

1-3

1,50

-35,

42-3

3,46

3,92

�4�

Tetr

acho

ndra

Ast

erid

sLa

mia

les

Tetra

chon

drac

eae

21

NZ,

1 P

atag

onia

, Tfu

ego

AU

S2

1-4

9,00

-54,

83-5

1,92

5,83

Tetr

aglo

chin

Ros

ids

Ros

ales

Ros

acea

e8

Arg

, Chi

l, P

erú

AN

D3

3-2

8,72

-46,

17-3

7,44

17,4

5

Tetr

agon

iaE

udic

ots

Car

yoph

ylla

les

Aiz

oace

ae70

N &

S A

fr, S

E A

sia,

Aus

, Tas

m, N

Z, P

olin

esia

, Per

u,

Chi

leW

-AN

T4

10-2

1,03

-31,

92-2

6,48

10,8

8

Tetr

onci

umM

onoc

ots

Alis

mat

ales

Junc

agin

acea

e1

tem

p S

Am

, Mag

alla

nes

51

-43,

00-5

6,00

-49,

5013

,00

Teuc

rium

Ast

erid

sLa

mia

les

Lam

iace

ae25

0C

osm

op7

2-2

5,00

-40,

00-3

2,50

15,0

0

Tham

nose

risA

ster

ids

Ast

eral

esA

ster

acea

e1

Des

vent

urad

as Is

land

s (C

hile

)6

10,

00

Thel

ypte

risFi

licop

sida

Ble

chna

les

Thel

ypte

ridac

eae

800

trops

& s

ubtro

ps, M

adag

11

-18,

27-3

7,33

-27,

8019

,07

Thyr

sopt

eris

Filic

opsi

daD

enns

taed

tiale

sD

icks

onia

ceae

1JF

61

0,00

Tigr

idia

Mon

ocot

sA

spar

agal

esIri

dace

ae35

Mex

, Gua

tem

ala

disj

unct

to P

eru,

Chi

le, B

ol, B

razi

lA

NT-

P4

1-2

2,00

-25,

40-2

3,70

3,40

Tilla

ndsi

aM

onoc

ots

Poa

les

Bro

mel

iace

ae54

0tro

p &

sub

trop

Am

, WI

W-N

T3

4-1

8,00

-38,

00-2

8,00

20,0

0

Tiqu

ilia

Ast

erid

sU

npla

ced

Bor

agin

acea

e27

arid

N &

S A

mA

NT-

P4

5-1

8,18

-29,

00-2

3,59

10,8

2

Trau

bia

Mon

ocot

sA

spar

agal

esA

mar

yllid

acea

e1

Chi

le6

1-3

1,50

-34,

00-3

2,75

2,50

Trev

oa

Ros

ids

Ros

ales

Rha

mna

ceae

1C

hile

61

-30,

42-3

5,83

-33,

135,

42

Trib

eles

Eud

icot

sS

axifr

agal

esS

axifr

agac

eae

1C

hile

& T

Fueg

o5

1-4

0,50

-55,

50-4

8,00

15,0

0

Tric

hocl

ine

Ast

erid

sA

ster

ales

Ast

erac

eae

22S

Am

, 1 W

Aus

AU

S2

5-1

8,43

-37,

82-2

8,13

19,3

8

Tric

hom

anes

Filic

opsi

daH

ymen

ophy

llale

sH

ymen

ophy

llace

ae80

patro

p, M

adag

, JF

11

-39,

87-4

3,10

-41,

483,

23

Tric

hone

ura

Mon

ocot

sP

oale

sP

oace

ae7

Ara

bia,

trop

Afr,

S U

S, P

eru

11

-18,

47-1

8,47

-18,

470,

00

Tric

hope

talu

mM

onoc

ots

Asp

arag

ales

Laxm

anni

acea

e2

Chi

le, A

rg5

1-2

5,43

-38,

23-3

1,83

12,8

0

Trifo

lium

Ros

ids

Faba

les

Faba

ceae

250

tem

p &

sub

trop

exc

Aus

W-A

NT

48

-25,

00-4

0,00

-32,

5015

,00

Trig

loch

inM

onoc

ots

Alis

mat

ales

Junc

agin

acea

e12

N te

mpe

rate

, circ

umbo

real

, aus

tral

W-A

NT

43

-20,

33-5

5,00

-37,

6734

,67

Trio

dani

sA

ster

ids

Ast

eral

esC

ampa

nula

ceae

8N

Am

, Mex

, Gua

tem

, Med

itW

-AN

T4

1-2

5,00

-40,

00-3

2,50

15,0

0

Trip

ogon

Mon

ocot

sP

oale

sP

oace

ae30

trops

& s

ubtro

ps1

1-1

8,38

-23,

58-2

0,98

5,20

Trip

tilio

nA

ster

ids

Ast

eral

esA

ster

acea

e12

C C

hile

, 1 P

atag

onia

512

-23,

00-4

2,50

-32,

7519

,50

Tris

etum

Mon

ocot

sP

oale

sP

oace

ae70

cold

& te

mp,

mon

tane

trop

ics,

JF

714

-25,

00-5

6,00

-40,

5031

,00

Tris

tagm

aM

onoc

ots

Asp

arag

ales

Alli

acea

e6

Chi

le, A

rg, U

ru5

12-2

9,00

-54,

83-4

1,92

25,8

3

Tris

terix

Eud

icot

sS

anta

lale

sLo

rant

hace

ae11

And

es C

olom

bia

to C

hile

AN

D3

3-2

8,50

-42,

50-3

5,50

14,0

0

Triu

mfe

ttaR

osid

sM

alva

les

Mal

vace

ae70

Pan

trop

11

0,00

Trix

isA

ster

ids

Ast

eral

esA

ster

acea

e50

SW

N A

m to

Chi

leW

-NT

31

-18,

55-2

0,93

-19,

742,

38

Trop

aeol

umR

osid

sB

rass

ical

esTr

opae

olac

eae

50M

exic

o to

Chi

leW

-NT

318

-22,

08-4

4,00

-33,

0421

,92

Trop

idoc

arpu

mR

osid

sB

rass

ical

esB

rass

icac

eae

4C

alifo

rnia

, Chi

leA

NT-

P4

1-3

0,67

-33,

38-3

2,03

2,72

Tuni

llaE

udic

ots

Car

yoph

ylla

les

Cac

tace

ae9

And

es P

eru,

Bol

, Chi

le, A

rgA

ND

32

-18,

00-2

2,00

-20,

004,

00

Twee

dia

Ast

erid

sG

entia

nale

sA

pocy

nace

ae6

Bol

, Uru

, Chi

le, A

rgS

-AM

Z3

2-2

6,75

-37,

67-3

2,21

10,9

2

Typh

aM

onoc

ots

Poa

les

Typh

acea

e13

Cos

mop

72

-19,

17-3

9,00

-29,

0819

,83

Ugn

iR

osid

sM

yrta

les

Myr

tace

ae10

Am

from

Mex

to C

hile

, JF

W-N

T3

2-3

4,17

-45,

38-3

9,78

11,2

2

Unc

inia

Mon

ocot

sP

oale

sC

yper

acea

e60

Mal

esia

, Pac

ific,

Aus

, SE

Asi

a, C

& S

Am

, JF

SP

T2

10-3

0,67

-56,

00-4

3,33

25,3

3

Urb

ania

Ast

erid

sLa

mia

les

Verb

enac

eae

1C

hile

, Arg

AN

D3

1-1

8,00

-24,

73-2

1,37

6,73

�44

Urm

enet

eaA

ster

ids

Ast

eral

esA

ster

acea

e1

N C

hile

, NW

Arg

AN

D3

1-2

1,50

-23,

18-2

2,34

1,68

Urt

ica

Ros

ids

Ros

ales

Urti

cace

ae75

subc

osm

op, t

empe

rate

710

-17,

72-5

5,50

-36,

6137

,78

Utr

icul

aria

Ast

erid

sLa

mia

les

Lent

ibul

aria

ceae

200

cosm

op7

1-2

8,72

-43,

00-3

5,86

14,2

8

Vald

ivia

Ast

erid

sU

npla

ced

Esc

allo

niac

eae

1C

hile

61

-39,

88-4

0,25

-40,

070,

37

Vale

riana

Ast

erid

sD

ipsa

cale

sVa

leria

nace

ae23

0N

tem

p E

uras

ia, S

Afr,

And

esW

-AN

T4

43-1

8,00

-55,

00-3

6,50

37,0

0

Vasc

once

llea

Ros

ids

Bra

ssic

ales

Car

icac

eae

20M

exic

o to

N A

rg &

S B

rasi

lW

-NT

31

-29,

67-3

3,00

-31,

333,

33

Verb

ena

Ast

erid

sLa

mia

les

Verb

enac

eae

225

tem

p &

trop

Am

, few

OW

W-A

NT

433

-18,

00-4

0,00

-29,

0022

,00

Verb

esin

aA

ster

ids

Ast

eral

esA

ster

acea

e30

0tro

p &

sub

trop

Am

, WI

W-N

T3

2-2

2,00

-29,

90-2

5,95

7,90

Vest

iaA

ster

ids

Sol

anal

esS

olan

acea

e1

Chi

le6

1-3

3,03

-40,

00-3

6,52

6,97

Vici

aR

osid

sFa

bale

sFa

bace

ae14

0te

mp

N H

emis

, Med

it, fe

w tr

op A

fr, S

Am

, Haw

aii

W-A

NT

428

-25,

00-5

5,33

-40,

1730

,33

Vigu

iera

Ast

erid

sA

ster

ales

Ast

erac

eae

180

trop

& s

ubtro

p A

m, W

IW

-NT

36

-18,

18-3

7,80

-27,

9919

,62

Villa

nova

Ast

erid

sA

ster

ales

Ast

erac

eae

10M

ex to

Chi

leW

-NT

32

-18,

20-2

5,00

-21,

606,

80

Viol

aR

osid

sM

alpi

ghia

les

Viol

acea

e43

0co

smop

770

-18,

00-5

6,00

-37,

0038

,00

Vitta

riaFi

licop

sida

Pte

ridal

esVi

ttaria

ceae

70pa

ntro

p1

10,

00

Vivi

ania

Ros

ids

Ger

ania

les

Vivi

ania

ceae

6S

Bra

sil,

Chi

le, A

rgS

-AM

Z3

3-2

8,72

-36,

83-3

2,78

8,12

Vulp

iaM

onoc

ots

Poa

les

Poa

ceae

25N

& S

Am

, Eur

o, N

Afr,

M A

sia

72

-23,

33-3

7,33

-30,

3314

,00

Wah

lenb

ergi

aA

ster

ids

Ast

eral

esC

ampa

nula

ceae

150

Sou

ther

n H

emis

pher

e, E

urop

e, S

E A

sia

W-A

NT

41

-30,

00-4

0,00

-35,

0010

,00

Web

erba

uera

Ros

ids

Bra

ssic

ales

Bra

ssic

acea

e18

And

es P

eru,

Chi

le, A

rg, B

olA

ND

37

-30,

67-3

6,90

-33,

786,

23

Wed

elia

Ast

erid

sA

ster

ales

Ast

erac

eae

100

pant

rops

, W A

ustra

lia1

1-2

3,58

-27,

37-2

5,48

3,78

Wei

nman

nia

Ros

ids

Oxa

lidal

esC

unon

iace

ae16

0M

adag

, Aus

trala

sia,

Mex

, C &

S A

mC

-AU

S2

1-3

5,33

-49,

42-4

2,38

14,0

8

Wen

dtia

Ros

ids

Ger

ania

les

Ledo

carp

acea

e3

Chi

le, A

rg5

1-3

0,50

-41,

73-3

6,12

11,2

3

Wer

derm

anni

aR

osid

sB

rass

ical

esB

rass

icac

eae

3N

Chi

le, A

rgA

ND

32

-21,

27-2

6,33

-23,

805,

07

Wer

neria

Ast

erid

sA

ster

ales

Ast

erac

eae

40A

ndes

Ecu

ador

to C

hile

AN

D3

9-1

7,58

-33,

82-2

5,70

16,2

3

Wolffia

Mon

ocot

sA

lism

atal

esLe

mna

ceae

11co

smop

, war

m te

mpe

rate

and

trop

ical

regi

ons

71

-36,

75-3

6,75

-36,

750,

00

Wolffiella

Mon

ocot

sA

lism

atal

esLe

mna

ceae

10tro

p &

war

m A

m, 1

S A

fr1

1-3

3,33

-36,

75-3

5,04

3,42

Woo

dsia

Filic

opsi

daD

ryop

terid

ales

Dry

opte

ridac

eae

25te

mp

& c

ool t

emp

Eur

asia

, Afr,

Am

, exc

Aus

W-A

NT

41

-18,

22-1

8,35

-18,

280,

13

Xeno

phyl

lum

Ast

erid

sA

ster

ales

Ast

erac

eae

21A

ndes

Col

ombi

a to

N A

rg, C

hile

AN

D3

5-1

8,00

-23,

58-2

0,79

5,58

Xero

drab

aR

osid

sB

rass

ical

esB

rass

icac

eae

6S

Arg

, Chi

le5

2-5

1,00

-51,

00-5

1,00

0,00

xMar

gyra

caen

aR

osid

sR

osal

esR

osac

eae

1JF

61

0,00

Yunq

uea

Ast

erid

sA

ster

ales

Ast

erac

eae

1JF

61

0,00

Zam

eios

cirp

usM

onoc

ots

Poa

les

Cyp

erac

eae

3A

rg, C

hile

, Bol

AN

D3

3-1

7,58

-33,

67-2

5,63

16,0

8

Zann

iche

llia

Mon

ocot

sA

lism

atal

esP

otam

oget

onac

eae

5co

smop

71

-22,

00-5

4,00

-38,

0032

,00

Zant

hoxy

lum

Ros

ids

Sap

inda

les

Rut

acea

e25

0A

m, A

fr, A

sia,

JF

12

0,00

Zeph

yra

Mon

ocot

sA

spar

agal

esTe

coph

ilaea

ceae

1C

hile

61

-20,

82-2

9,00

-24,

918,

18

Zoel

lner

alliu

mM

onoc

ots

Asp

arag

ales

Alli

acea

e2

Chi

le, A

rgA

ND

32

-30,

00-3

3,83

-31,

923,

83

�4�

ELEMENTS N° of genera

1. Pantropical 88

2. Australasiatic 59

3. Neotropical 216

4. Antitropical 152

5. South-temperate 81

6. Endemic 83

7. Cosmopolitan 134

TOTAL 813

Appendix A (continuation): Legend and summary for tracks and elements

TRACKS (follow subsections in the text) N° of genera3.2.1 Austral-antarctic track AUS 313.2.2 South Pacific tropical track SPT 253.2.3 Circum-austral track C-AUS 3

3.3.1 Wide Neotropical track W-NT 643.3.2 Andean track AND 1133.3.3 South Amazonian track S-AMZ 39

3.4.1 Wide antitropical track W-ANT 843.4.2 Antitropical disjunct Pacific track ANT-P 563.4.3 Circum-Pacific track C-PAC 9

�4�

�47

Appendix B: Genera shared by several Chilean regions

�48

GEN Summe ANT BIO COQ JF MAG GEN Summe ANT BIO COQ JF MAG

Aa 1 1 Laretia 1 1

Abrotanella 2 1 1 Larrea 2 1 1

Acacia 2 1 1 Lastarriaea 1 1

Acaena 5 1 1 1 1 1 Lasthenia 3 1 1 1

Acantholippia 2 1 1 Lathyrus 4 1 1 1 1

Acrisione 2 1 1 Laurelia 1 1

Adenocaulon 1 1 Laureliopsis 1 1

Adenopeltis 2 1 1 Lebetanthus 1 1

Adesmia 4 1 1 1 1 Legrandia 1 1

Adiantum 5 1 1 1 1 1 Lemna 3 1 1 1

Aextoxicon 2 1 1 Lenzia 2 1 1

Agalinis 1 1 Lepidium 4 1 1 1 1

Agallis 1 1 Lepidoceras 1 1

Ageratina 3 1 1 1 Lepidophyllum 1 1

Agoseris 3 1 1 1 Lepidothamnus 1 1

Agrostis 4 1 1 1 1 Leptinella 1 1

Allionia 1 1 Leptocarpha 1 1

Alona 2 1 1 Leptophyllochloa 2 1 1

Alonsoa 2 1 1 Leptostigma 1 1

Alopecurus 3 1 1 1 Lepuropetalon 1 1

Aloysia 1 1 Leucheria 4 1 1 1 1

Alstroemeria 4 1 1 1 1 Leucocoryne 3 1 1 1

Alternanthera 1 1 Leunisia 1 1

Amaranthus 1 1 Libertia 3 1 1 1

Amblyopappus 2 1 1 Ligaria 2 1 1

Ambrosia 3 1 1 1 Lilaeopsis 4 1 1 1 1

Amomyrtus 1 1 Limonium 1 1

Amphibromus 1 1 Limosella 4 1 1 1 1

Amsimkia 1 1 Linanthus 1 1

Amsinckia 3 1 1 1 Lindernia 1 1

Anagallis 4 1 1 1 1 Linum 3 1 1 1

Anarthrophyllum 2 1 1 Lippia 2 1 1

Anatherostipa 1 1 Lithrea 2 1 1

Androsace 1 1 Littorella 1 1

Anemone 3 1 1 1 Llagunoa 2 1 1

Anisomeria 2 1 1 Loasa 4 1 1 1 1

Antennaria 3 1 1 1 Lobelia 3 1 1 1

Anthochloa 1 1 locenes 1 1

Anthoxanthum 2 1 1 Lomatia 3 1 1 1

Antidaphne 1 1 Lophosoria 2 1 1

Aphanes 1 1 Lotus 3 1 1 1

Apium 5 1 1 1 1 1 Ludwigia 2 1 1

Apodasmia 2 1 1 Luma 2 1 1

Arachnitis 2 1 1 Lupinus 3 1 1 1

Araeoandra 1 1 Luzula 4 1 1 1 1

Araucaria 1 1 Luzuriaga 2 1 1

Arenaria 4 1 1 1 1 Lycium 2 1 1

Argemone 2 1 1 Lycopersicon 1 1

Argylia 3 1 1 1 Lycopodium 3 1 1 1

Aristeguietia 2 1 1 Lysimachia 1 1

Aristida 3 1 1 1 Machaerina 1 1

Aristolochia 1 1 Macrachaenium 1 1

Aristotelia 2 1 1 Madia 3 1 1 1

�49

Arjona 2 1 1 Maihuenia 1 1

Armeria 3 1 1 1 Maihueniopsis 2 1 1

Artemisia 2 1 1 Malacothamnus 1 1

Arthropteris 1 1 Malacothrix 2 1 1

Asplenium 5 1 1 1 1 1 Malesherbia 2 1 1

Astelia 1 1 Malvella 2 1 1

Aster 3 1 1 1 Mancoa 1 1

Asteranthera 2 1 1 Margyricarpus 4 1 1 1 1

Asteriscium 3 1 1 1 Marsilea 1 1

Astragalus 4 1 1 1 1 Marsippospermum 2 1 1

Atriplex 4 1 1 1 1 Mastigostyla 1 1

Austrocedrus 1 1 Mathewsia 2 1 1

Azara 3 1 1 1 Maytenus 3 1 1 1

Azolla 4 1 1 1 1 Megalachne 1 1

Azorella 4 1 1 1 1 Megalastrum 3 1 1 1

Baccharis 4 1 1 1 1 Melica 2 1 1

Bacopa 2 1 1 Melosperma 2 1 1

Bahia 3 1 1 1 Menodora 1 1

Bakerolimon 2 1 1 Menonvillea 4 1 1 1 1

Balbisia 2 1 1 Mentzelia 2 1 1

Balsamocarpon 1 1 Microphyes 3 1 1 1

Barneoudia 1 1 Micropsis 2 1 1

Bartsia 1 1 Microseris 3 1 1 1

Beilschmiedia 1 1 Microsteris 3 1 1 1

Belloa 2 1 1 Miersia 1 1

Benthamiella 1 1 Mikania 1 1

Berberidopsis 1 1 Mimulus 5 1 1 1 1 1

Berberis 5 1 1 1 1 1 Minuartia 1 1

Bidens 3 1 1 1 Miqueliopuntia 1 1

Bipinnula 3 1 1 1 Mirabilis 2 1 1

Blechnum 4 1 1 1 1 Misodendrum 2 1 1

Blennosperma 1 1 Mitraria 3 1 1 1

Blepharocalyx 1 1 Montia 3 1 1 1

Boehmeria 1 1 Montiopsis 3 1 1 1

Boerhavia 2 1 1 Monttea 2 1 1

Boisduvalia 2 1 1 Moscharia 2 1 1

Bolax 1 1 Moschopsis 1 1

Bomarea 1 1 Moschopsìs 1 1

Boopis 2 1 1 Muehlenbeckia 2 1 1

Boquila 1 1 Muhlenbergia 2 1 1

Botrychium 1 1 Mulinum 4 1 1 1 1

Bouteloua 1 1 Munroa 1 1

Bowlesia 4 1 1 1 1 Mutisia 3 1 1 1

Brachyclados 1 1 Myoschilos 3 1 1 1

Brachystele 2 1 1 Myosotis 1 1

Bridgesia 1 1 Myosurus 3 1 1 1

Bromidium 2 1 1 Myrceugenia 3 1 1 1

Bromus 5 1 1 1 1 1 Myrcianthes 1 1

Buddleja 2 1 1 Myriophyllum 4 1 1 1 1

Bulbostylis 1 1 Myrteola 3 1 1 1

Bulnesia 1 1 Nama 1 1

Caesalpinia 2 1 1 Nanodea 1 1

��0

Caiophora 3 1 1 1 Nardophyllum 3 1 1 1

Calamagrostis 4 1 1 1 1 Nassauvia 3 1 1 1

Calandrinia 4 1 1 1 1 Nassella 5 1 1 1 1 1

Calceolaria 4 1 1 1 1 Nastanthus 3 1 1 1

Caldcluvia 1 1 Nasturtium 1 1

Calliandra 1 1 Navarretia 2 1 1

Callitriche 4 1 1 1 1 Nertera 4 1 1 1 1

Calopappus 1 1 Nicandra 1 1

Calotheca 1 1 Nicotiana 4 1 1 1 1

Caltha 3 1 1 1 Nierembergia 1 1

Calycera 3 1 1 1 Nitrophila 2 1 1

Calydorea 2 1 1 Nolana 3 1 1 1

Calystegia 2 1 1 Notanthera 2 1 1

Camissonia 2 1 1 Nothofagus 2 1 1

Campsidium 2 1 1 Notholaena 2 1 1

Cardamine 4 1 1 1 1 Nothoscordum 2 1 1

Cardionema 3 1 1 1 Noticastrum 1 1

Carex 5 1 1 1 1 1 Nototriche 2 1 1

Carica 1 1 Ochagavia 3 1 1 1

Carpha 1 1 Oenothera 4 1 1 1 1

Carpobrotus 3 1 1 1 Olsynium 4 1 1 1 1

Castilleja 2 1 1 Ombrophytum 1 1

Catabrosa 3 1 1 1 Onuris 1 1

Centaurea 3 1 1 1 Ophioglossum 4 1 1 1 1

Centaurium 3 1 1 1 Ophryosporus 2 1 1

Centaurodendron 1 1 Oreobolus 3 1 1 1

Centella 2 1 1 Oreocereus 1 1

Centipeda 2 1 1 Oreomyrrhis 1 1

Cerastium 3 1 1 1 Oreopolus 2 1 1

Ceratophyllum 1 1 Orites 1 1

Cestrum 3 1 1 1 Ornithopus 1 1

Chaetanthera 3 1 1 1 Orobanche 1 1

Chaetotropis 2 1 1 Ortachne 1 1

Chaptalia 1 1 Osmorhiza 3 1 1 1

Chascolytrum 2 1 1 Otholobium 2 1 1

Cheilanthes 3 1 1 1 Ourisia 3 1 1 1

Chenopodium 5 1 1 1 1 1 Ovidia 1 1

Chersodoma 1 1 Oxalis 4 1 1 1 1

Chevreulia 1 1 Oxychloe 2 1 1

Chiliophyllum 1 1 Oxyphyllum 1 1

Chiliotrichum 2 1 1 Oxytheca 2 1 1

Chiropetalum 3 1 1 1 Oziroe 3 1 1 1

Chloraea 3 1 1 1 Pachylaena 1 1

Chorizanthe 2 1 1 Palaua 1 1

Chrysosplenium 1 1 Panicum 1 1

Chuquiraga 2 1 1 Parastrephia 1 1

Chusquea 3 1 1 1 Parietaria 3 1 1 1

Cicendia 2 1 1 Paronychia 3 1 1 1

Ciclospermum 1 1 Pasithea 3 1 1 1

Cissarobryon 1 1 Paspalum 2 1 1

Cissus 2 1 1 Passiflora 1 1

Cistanthe 3 1 1 1 Patosia 3 1 1 1

���

Citronella 2 1 1 Pectocarya 3 1 1 1

Clarkia 2 1 1 Pellaea 3 1 1 1

Cleome 1 1 Pelletiera 1 1

Cliococca 1 1 Pennisetum 2 1 1

Codonorchis 2 1 1 Peperomia 3 1 1 1

Colletia 3 1 1 1 Perezia 4 1 1 1 1

Colliguaja 3 1 1 1 Perityle 2 1 1

Collomia 2 1 1 Persea 1 1

Colobanthus 4 1 1 1 1 Peumus 2 1 1

Combera 1 1 Phacelia 4 1 1 1 1

Conanthera 3 1 1 1 Phalaris 2 1 1

Convolvulus 3 1 1 1 Philesia 1 1

Conyza 4 1 1 1 1 Philibertia 1 1

Copiapoa 2 1 1 Philippiella 1 1

Coprosma 1 1 Phleum 2 1 1

Cordia 1 1 Phragmites 3 1 1 1

Coriaria 1 1 Phrodus 1 1

Coronopus 2 1 1 Phycella 2 1 1

Corrigiola 2 1 1 Phyllachne 1 1

Cortaderia 4 1 1 1 1 Phylloscirpus 1 1

Corynabutilon 2 1 1 Phytolacca 1 1

Cotula 1 1 Picrosia 1 1

Crassula 4 1 1 1 1 Pilea 1 1

Cressa 2 1 1 Pilgerodendron 1 1

Crinodendron 1 1 Pilostyles 3 1 1 1

Cristaria 2 1 1 Pilularia 2 1 1

Croton 1 1 Pinguicola 1 1

Cruckshanksia 2 1 1 Pinguicula 1 1

Cryptantha 3 1 1 1 Piptochaetium 3 1 1 1

Cryptocarya 2 1 1 Pitavia 1 1

Cryptogramma 2 1 1 Pitraea 2 1 1

Cuatrecasasiella 1 1 Placea 1 1

Cuminia 1 1 Plagiobothrys 3 1 1 1

Cumulopuntia 2 1 1 Plantago 5 1 1 1 1 1

Cuscuta 3 1 1 1 Plazia 1 1

Cyclospermum 2 1 1 Plectritis 1 1

Cynodon 2 1 1 Pleocarphus 1 1

Cynoglossum 1 1 Pleopeltis 3 1 1 1

Cyperus 4 1 1 1 1 Pleurophora 2 1 1

Cyphocarpus 1 1 Pleurosorus 2 1 1

Cystopteris 4 1 1 1 1 Pluchea 1 1

Dalea 1 1 Plumbago 2 1 1

Danthonia 3 1 1 1 Poa 4 1 1 1 1

Dasyphyllum 1 1 Podagrostis 1 1

Daucus 3 1 1 1 Podanthus 2 1 1

Dendroseris 1 1 Podocarpus 2 1 1

Dennstaedtia 1 1 Podophorus 1 1

Deschampsia 3 1 1 1 Polemonium 2 1 1

Descurainia 3 1 1 1 Polyachyrus 2 1 1

Desfontainia 2 1 1 Polycarpon 1 1

Desmaria 1 1 Polygala 3 1 1 1

Deuterocohnia 1 1 Polygonum 2 1 1

Dichondra 4 1 1 1 1 Polylepis 1 1

Dicksonia 1 1 Polypodium 5 1 1 1 1 1

���

Dicllptera 1 1 Polypogon 4 1 1 1 1

Dielsiochloa 1 1 Polystichum 4 1 1 1 1

Dinemagonum 1 1 Porlieria 1 1

Dinemandra 1 1 Portulaca 1 1

Dioscorea 3 1 1 1 Potamogeton 4 1 1 1 1

Diostea 2 1 1 Pouteria 1 1

Diplachne 1 1 Pozoa 2 1 1

Diplolepis 3 1 1 1 Pratia 2 1 1

Diplostephium 1 1 Primula 1 1

Diposis 1 1 Prosopis 2 1 1

Discaria 3 1 1 1 Proustia 3 1 1 1

Distichia 1 1 Prumnopitys 1 1

Distichlis 3 1 1 1 Pseudopanax 2 1 1

Dodonaea 1 1 Psilocarphus 2 1 1

Domeykoa 1 1 Pteris 3 1 1 1

Donatia 1 1 Pteromonnina 2 1 1

Doniophyton 2 1 1 Puccinellia 3 1 1 1

Downingia 2 1 1 Puya 3 1 1 1

Draba 3 1 1 1 Pycnophyllum 1 1

Drapetes 1 1 Quillaja 2 1 1

Drimys 4 1 1 1 1 Quinchamalium 3 1 1 1

Drosera 1 1 Raimundochloa 2 1 1

Drymaria 1 1 Ranunculus 5 1 1 1 1 1

Dysopsis 4 1 1 1 1 Reicheella 1 1

Eccremocarpus 1 1 Relchela 1 1

Echinopsis 2 1 1 Retanilla 2 1 1

Elaphoglossum 1 1 Reyesia 2 1 1

Elatine 1 1 Rhamnus 1 1

Eleocharis 5 1 1 1 1 1 Rhaphithamnus 3 1 1 1

Elodea 1 1 Rhodophiala 3 1 1 1

Elymus 4 1 1 1 1 Ribes 3 1 1 1

Elytropus 1 1 Robinsonia 1 1

Embothrium 2 1 1 Rorippa 1 1

Empetrum 3 1 1 1 Rostkovia 1 1

Encelia 2 1 1 Rubus 3 1 1 1

Enneapogon 1 1 Rumex 3 1 1 1

Ephedra 4 1 1 1 1 Rumohra 4 1 1 1 1

Epilobium 4 1 1 1 1 Ruppia 3 1 1 1

Epipetrum 3 1 1 1 Rytidosperma 3 1 1 1

Equisetum 2 1 1 Sagittaria 1 1

Eragrostis 3 1 1 1 Salix 3 1 1 1

Ercilla 1 1 Salpiglossis 2 1 1

Erechtites 1 1 Salvia 1 1

Eremocharis 1 1 Samolus 2 1 1

Eriachaenium 1 1 Sanicula 2 1 1

Erigeron 4 1 1 1 1 Santalum 1 1

Eriosyce 3 1 1 1 Sarcocornia 5 1 1 1 1 1

Errazurizia 2 1 1 Sarmienta 2 1 1

Eryngium 4 1 1 1 1 Satureja 4 1 1 1 1

Escallonia 4 1 1 1 1 Saxegothaea 1 1

Eucryphia 1 1 Saxifraga 2 1 1

Eudema 2 1 1 Saxifragella 1 1

Eulychnia 2 1 1 Saxifragodes 1 1

Euphorbia 4 1 1 1 1 Schinus 2 1 1

���

Euphrasia 4 1 1 1 1 Schizaea 1 1

Evolvulus 1 1 Schizanthus 3 1 1 1

Exodeconus 1 1 Schizeilema 1 1

Fabiana 3 1 1 1 Schizopetalon 1 1

Facelis 3 1 1 1 Schkuhria 2 1 1

Fagonia 2 1 1 Schlnus 1 1

Fascicularia 1 1 Schoenus 2 1 1

Festuca 4 1 1 1 1 Scirpus 5 1 1 1 1 1

Flaveria 3 1 1 1 Scutellaria 2 1 1

Flourensia 2 1 1 Scyphanthus 2 1 1

Fragaria 1 1 Selkirkia 1 1

Francoa 1 1 Selliera 2 1 1

Frankenia 4 1 1 1 1 Senecio 4 1 1 1 1

Fuchsia 3 1 1 1 Senna 3 1 1 1

Fuertesimalva 2 1 1 Serpyllopsis 2 1 1

Gaimardia 1 1 Sicyos 2 1 1

Galinsoga 3 1 1 1 Sigesbeckia 1 1

Galium 5 1 1 1 1 1 Silene 4 1 1 1 1

Gamocarpha 2 1 1 Sisymbrium 4 1 1 1 1

Gamochaeta 5 1 1 1 1 1 Sisyrinchium 4 1 1 1 1

Gamochaetopsis 1 1 Skytanthus 2 1 1

Gaultheria 5 1 1 1 1 1 Solanum 4 1 1 1 1

Gavilea 4 1 1 1 1 Solaria 1 1

Gayophytum 2 1 1 Solenomelus 3 1 1 1

Gentiana 4 1 1 1 1 Solidago 2 1 1

Gentianella 4 1 1 1 1 Soliva 2 1 1

Geoffroea 2 1 1 Sophora 3 1 1 1

Geranium 4 1 1 1 1 Spartina 1 1

Gethyum 1 1 Spergularia 4 1 1 1 1

Geum 2 1 1 Sphacele 2 1 1

Gevuina 1 1 Sphaeralcea 2 1 1

Gilia 3 1 1 1 Spirodela 1 1

Gilliesia 2 1 1 Stachys 3 1 1 1

Glandularia 3 1 1 1 Stellaria 4 1 1 1 1

Gleichenia 3 1 1 1 Stemodia 2 1 1

Glinus 2 1 1 Stenandrium 2 1 1

Glyceria 1 1 Stevia 1 1

Glycyrrhiza 1 1 Stipa 4 1 1 1 1

Gnaphalium 4 1 1 1 1 Suaeda 3 1 1 1

Gochnatia 2 1 1 Tagetes 3 1 1 1

Gomortega 1 1 Tapeinia 1 1

Gomphrena 1 1 Tarasa 2 1 1

Grabowskia 1 1 Taraxacum 2 1 1

Grammitis 3 1 1 1 Tecophilaea 1 1

Grammosperma 1 1 Tepualia 2 1 1

Gratiola 1 1 Tessaria 2 1 1

Greigia 2 1 1 Tetilla 1 1

Griselinia 4 1 1 1 1 Tetrachondra 1 1

Guindilia 1 1 Tetraglochin 2 1 1

Gunnera 4 1 1 1 1 Tetragonia 2 1 1

Gutierrezia 3 1 1 1 Tetroncium 1 1

Guynesomia 1 1 Teucrium 3 1 1 1

Gymnachne 1 1 Thelypteris 3 1 1 1

Gymnophyton 2 1 1 Thlaspi 1 1

��4

Gypothamnium 1 1 Thyrsopteris 1 1

Habenaria 2 1 1 Tigridia 1 1

Haloragis 1 1 Tillandsia 3 1 1 1

Hamadryas 1 1 Tiquilia 1 1

Haplopappus 3 1 1 1 Traubia 1 1

Hebe 1 1 Trevoa 2 1 1

Hedyotis 2 1 1 Tribeles 1 1

Helenium 3 1 1 1 Trichocline 3 1 1 1

Helictotrichon 1 1 Trichomanes 1 1

Heliotropium 4 1 1 1 1 Trichopetalum 2 1 1

Helogyne 1 1 Trifolium 3 1 1 1

Herbertia 1 1 Triglochin 4 1 1 1 1

Herreria 1 1 Triodanis 1 1

Heterosperma 1 1 Triptilion 3 1 1 1

Heterozostera 1 1 Trisetum 5 1 1 1 1 1

Hieracium 2 1 1 Tristagma 3 1 1 1

Hierochloe 2 1 1 Tristerix 2 1 1

Hippuris 2 1 1 Tropaeolum 3 1 1 1

Histiopteris 2 1 1 Tunilla 1 1

Hoffmannseggia 2 1 1 Tweedia 3 1 1 1

Homalocarpus 2 1 1 Typha 3 1 1 1

Hordeum 4 1 1 1 1 Ugni 2 1 1

Hornungia 1 1 Uncinia 4 1 1 1 1

Huanaca 2 1 1 Urbania 1 1

Huidobria 2 1 1 Urmenetea 1 1

Huperzia 1 1 Urtica 5 1 1 1 1 1

Hybanthus 1 1 Utricularia 2 1 1

Hydrangea 1 1 Valeriana 4 1 1 1 1

Hydrocotyle 3 1 1 1 Verbena 3 1 1 1

Hymenoglossum 3 1 1 1 Verbesina 2 1 1

Hymenophyllum 4 1 1 1 1 Vestia 1 1

Hypericum 2 1 1 Vicia 4 1 1 1 1

Hypochaeris 4 1 1 1 1 Viguiera 3 1 1 1

Hypolepis 4 1 1 1 1 Villanova 1 1

Hypsela 2 1 1 Viola 4 1 1 1 1

Imperata 2 1 1 Viviania 2 1 1

Ipomopsis 1 1 Vulpia 3 1 1 1

Isoetes 3 1 1 1 Wahlenbergia 3 1 1 1

Jaborosa 4 1 1 1 1 Weberbauera 3 1 1 1

Jarava 3 1 1 1 Weinmannia 2 1 1

Jovellana 1 1 Wendtia 2 1 1

Juania 1 1 Werdermannia 1 1

Jubaea 1 1 Werneria 2 1 1

Juncus 5 1 1 1 1 1 Wolffia 1 1

Junellia 3 1 1 1 Wolffiella 1 1

Kageneckia 2 1 1 Xenophyllum 1 1

Koeleria 1 1 Xerodraba 1 1

Krameria 2 1 1 xMargyracaena 1 1

Kurzamra 1 1 Yunquea 1 1

Lactoris 1 1 Zannichellia 3 1 1 1

Lagenophora 3 1 1 1 Zanthoxylum 1 1

Lampaya 1 1 Zephyra 2 1 1

Lapageria 2 1 1 Zoellnerallium 1 1

Lardizabala 2 1 1

���

Appendix C: Matrix for PAE: Distribution of Endemic Ge-nera

���

N° GENERO A an B co B an C co C an D co D an E co E an F co F an G co G in G an H co

1 Acrisione 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

2 Adenopeltis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

3 Alona 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1

4 Anisomeria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

5 Araeoandra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

6 Avellanita 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

7 Bakerolimon 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1

8 Balsamocarpon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

9 Bridgesia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

10 Calopappus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

11 Cissarobryon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

12 Conanthera 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1

13 Copiapoa 0 0 0 0 0 0 0 1 0 1 0 1 0 0 1

14 Cyphocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

15 Desmaria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

16 Dinemagonum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

17 Dinemandra 0 0 0 0 0 0 0 1 0 1 1 1 1 0 1

18 Epipetrum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

19 Ercilla 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

20 Fascicularia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

21 Francoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

22 Gethyum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

23 Gomortega 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

24 Guynesomia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

25 Gymnachne 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

26 Gypothamnium 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

27 Hollermayera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

28 Homalocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

29 Huidobria 0 0 0 1 1 0 1 1 0 1 0 1 0 1 1

30 Hymenoglossum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

31 Ivania 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

32 Jubaea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

33 Lapageria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

34 Latua 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

35 Legrandia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

36 Leontochir 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

37 Leptocarpha 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

38 Leucocoryne 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0

39 Leunisia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

40 Marticorenia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

41 Metharme 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0

42 Microphyes 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

43 Miersia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

44 Miqueliopuntia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

45 Moscharia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

46 Neuontobothrys 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

47 Notanthera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

48 Ochagavia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

49 Oxyphyllum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

50 Peumus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

51 Phrodus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

52 Pintoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

53 Pitavia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

54 Placea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

55 Pleocarphus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

56 Podanthus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

57 Reicheella 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0

58 Sarmienta 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

59 Scyphanthus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

60 Speea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

61 Tecophilaea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

62 Tetilla 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

63 Traubia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

64 Trevoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

65 Valdivia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

66 Vestia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

67 Zephyra 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0

TOT 0 0 0 1 1 4 2 4 1 6 2 5 1 1 9

��7

N° GENERO H an I co I an J co J an K co K an L co L an M co M an N co N an

1 Acrisione 0 0 0 0 0 0 0 0 0 0 0 0 0

2 Adenopeltis 0 0 0 0 0 0 0 0 0 0 0 1 0

3 Alona 0 1 0 1 0 1 0 1 0 1 0 1 0

4 Anisomeria 0 1 0 0 0 0 0 1 0 1 0 1 0

5 Araeoandra 0 0 0 0 0 0 0 0 0 1 0 1 0

6 Avellanita 0 0 0 0 0 0 0 0 0 0 0 0 0

7 Bakerolimon 0 1 0 1 0 1 0 0 0 1 0 0 0

8 Balsamocarpon 0 0 0 0 0 1 0 1 1 1 0 0 0

9 Bridgesia 0 0 0 0 0 0 0 1 0 1 0 1 1

10 Calopappus 0 0 0 0 0 0 0 0 0 0 0 0 0

11 Cissarobryon 0 0 0 0 0 0 0 0 0 0 0 0 0

12 Conanthera 0 1 0 0 0 1 0 1 0 1 0 1 0

13 Copiapoa 0 1 0 1 0 1 0 1 0 1 0 1 0

14 Cyphocarpus 0 0 0 0 0 1 0 1 0 1 0 1 1

15 Desmaria 0 0 0 0 0 0 0 0 0 0 0 0 0

16 Dinemagonum 0 0 0 0 0 1 0 1 0 1 1 1 1

17 Dinemandra 0 1 0 1 0 1 0 0 0 0 0 0 0

18 Epipetrum 0 1 0 0 0 0 0 0 0 0 0 1 0

19 Ercilla 0 0 0 0 0 0 0 0 0 0 0 0 0

20 Fascicularia 0 0 0 0 0 0 0 0 0 0 0 0 0

21 Francoa 0 0 0 0 0 0 0 0 0 0 0 0 0

22 Gethyum 0 0 0 0 0 0 0 0 0 1 0 1 0

23 Gomortega 0 0 0 0 0 0 0 0 0 0 0 0 0

24 Guynesomia 0 0 0 0 0 0 0 0 0 0 1 0 1

25 Gymnachne 0 0 0 0 0 0 0 0 0 0 0 0 0

26 Gypothamnium 0 1 0 1 0 0 0 0 0 0 0 0 0

27 Hollermayera 0 0 0 0 0 0 0 0 0 0 0 0 0

28 Homalocarpus 0 1 0 0 0 1 0 1 1 1 0 1 1

29 Huidobria 0 1 0 1 1 1 0 0 1 1 0 0 0

30 Hymenoglossum 0 0 0 0 0 0 0 0 0 0 0 0 0

31 Ivania 0 0 0 0 0 1 0 0 0 0 0 0 0

32 Jubaea 0 0 0 0 0 0 0 0 0 0 0 0 0

33 Lapageria 0 0 0 0 0 0 0 0 0 0 0 1 0

34 Latua 0 0 0 0 0 0 0 0 0 0 0 0 0

35 Legrandia 0 0 0 0 0 0 0 0 0 0 0 0 0

36 Leontochir 0 0 0 0 0 1 0 1 0 0 0 0 0

37 Leptocarpha 0 0 0 0 0 0 0 0 0 0 0 0 0

38 Leucocoryne 0 1 0 1 0 1 0 1 0 1 1 1 0

39 Leunisia 0 0 0 0 0 0 0 0 0 0 0 1 0

40 Marticorenia 0 0 0 0 0 0 0 0 0 0 0 0 0

41 Metharme 0 0 0 0 0 0 0 0 0 0 0 0 0

42 Microphyes 0 1 0 1 0 1 0 1 0 1 1 1 1

43 Miersia 0 0 0 0 0 0 0 0 0 0 0 0 0

44 Miqueliopuntia 0 0 0 0 0 1 0 1 0 1 0 0 0

45 Moscharia 0 0 0 0 0 0 0 0 0 1 0 1 0

46 Neuontobothrys 0 0 0 0 0 0 0 0 0 0 0 0 0

47 Notanthera 0 0 0 0 0 0 0 0 0 0 0 0 0

48 Ochagavia 0 0 0 0 0 0 0 0 0 0 0 0 0

49 Oxyphyllum 0 1 0 1 0 0 0 0 0 0 0 0 0

50 Peumus 0 0 0 0 0 0 0 0 0 0 0 1 0

51 Phrodus 0 0 0 0 1 1 0 1 1 1 1 1 1

52 Pintoa 0 0 0 0 0 1 0 0 1 0 0 0 0

53 Pitavia 0 0 0 0 0 0 0 0 0 0 0 0 0

54 Placea 0 0 0 0 0 0 0 0 0 0 0 0 0

55 Pleocarphus 0 0 0 0 0 0 0 0 0 1 0 1 0

56 Podanthus 0 0 0 0 0 0 0 0 0 0 0 1 0

57 Reicheella 0 0 0 0 0 0 0 0 0 0 0 0 0

58 Sarmienta 0 0 0 0 0 0 0 0 0 0 0 1 0

59 Scyphanthus 0 0 0 0 0 0 0 0 0 0 0 0 0

60 Speea 0 0 0 0 0 0 0 0 0 0 0 0 0

61 Tecophilaea 0 0 0 0 0 0 0 0 0 0 0 1 0

62 Tetilla 0 0 0 0 0 0 0 0 0 0 0 0 0

63 Traubia 0 0 0 0 0 0 0 0 0 0 0 0 0

64 Trevoa 0 0 0 0 0 0 0 0 0 0 0 1 0

65 Valdivia 0 0 0 0 0 0 0 0 0 0 0 0 0

66 Vestia 0 0 0 0 0 0 0 0 0 0 0 0 0

67 Zephyra 0 1 0 0 0 1 0 1 0 1 0 0 0

TOT 0 14 0 9 2 18 0 15 5 20 5 24 7

��8

N° GENERO O co O an P co P an Q co Q an R co R an S co S an T co T an U co U an V co

1 Acrisione 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1

2 Adenopeltis 1 0 1 0 1 0 1 0 1 0 1 0 0 0 0

3 Alona 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0

4 Anisomeria 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0

5 Araeoandra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

6 Avellanita 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0

7 Bakerolimon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

8 Balsamocarpon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

9 Bridgesia 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0

10 Calopappus 0 0 0 1 0 1 0 1 0 1 0 0 0 0 0

11 Cissarobryon 0 0 0 0 1 1 0 1 0 1 0 1 1 1 0

12 Conanthera 1 0 1 1 1 1 1 1 1 1 1 1 1 0 0

13 Copiapoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

14 Cyphocarpus 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0

15 Desmaria 0 0 0 0 0 0 0 0 0 1 0 1 1 1 1

16 Dinemagonum 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0

17 Dinemandra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

18 Epipetrum 1 0 1 0 1 0 1 1 1 0 0 0 0 1 0

19 Ercilla 0 0 0 0 1 1 1 1 1 1 1 0 1 0 1

20 Fascicularia 0 0 0 0 1 0 0 0 1 0 1 0 1 0 1

21 Francoa 0 0 1 0 1 0 1 0 1 1 1 1 1 1 1

22 Gethyum 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0

23 Gomortega 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0

24 Guynesomia 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

25 Gymnachne 0 0 0 0 1 0 1 0 1 0 1 0 1 0 1

26 Gypothamnium 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

27 Hollermayera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

28 Homalocarpus 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0

29 Huidobria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

30 Hymenoglossum 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1

31 Ivania 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

32 Jubaea 1 0 1 0 1 0 1 0 1 0 0 0 0 0 0

33 Lapageria 0 0 0 0 1 0 0 0 1 0 1 1 1 0 1

34 Latua 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

35 Legrandia 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0

36 Leontochir 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

37 Leptocarpha 0 0 0 0 0 0 0 0 1 0 1 1 1 1 1

38 Leucocoryne 1 1 1 1 1 1 1 0 1 1 1 0 1 0 0

39 Leunisia 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

40 Marticorenia 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0

41 Metharme 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

42 Microphyes 1 1 1 0 1 1 0 0 0 1 1 0 1 1 0

43 Miersia 1 0 1 0 1 0 1 0 1 0 0 0 0 0 0

44 Miqueliopuntia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

45 Moscharia 1 0 1 0 1 1 1 0 1 0 1 0 0 0 0

46 Neuontobothrys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

47 Notanthera 0 0 1 0 1 0 1 1 1 1 1 1 1 1 1

48 Ochagavia 1 0 1 0 1 0 1 1 1 1 1 1 1 1 0

49 Oxyphyllum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

50 Peumus 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1

51 Phrodus 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0

52 Pintoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

53 Pitavia 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0

54 Placea 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0

55 Pleocarphus 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0

56 Podanthus 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1

57 Reicheella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

58 Sarmienta 0 0 0 0 0 0 0 0 1 0 1 0 1 0 1

59 Scyphanthus 1 1 1 1 0 1 0 1 1 1 1 1 0 1 0

60 Speea 0 0 1 0 1 1 0 1 0 0 0 0 0 0 0

61 Tecophilaea 1 0 1 0 1 1 0 0 0 0 0 0 0 0 0

62 Tetilla 1 0 1 0 1 1 1 1 1 0 0 0 0 0 0

63 Traubia 1 0 1 0 1 0 1 0 0 0 0 0 0 0 0

64 Trevoa 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0

65 Valdivia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

66 Vestia 0 0 1 0 1 0 0 1 1 0 1 1 1 1 1

67 Zephyra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

TOT 26 12 28 12 31 20 20 18 26 17 22 14 21 13 14

��9

N° GENERO V an W co W an X co X an Y co Y an Z co Z an AA co AA an AB co AB an

1 Acrisione 0 1 0 0 0 0 0 0 0 0 0 0 0

2 Adenopeltis 0 0 0 0 0 0 0 0 0 0 0 0 0

3 Alona 0 0 0 0 0 0 0 0 0 0 0 0 0

4 Anisomeria 0 0 0 0 0 0 0 0 0 0 0 0 0

5 Araeoandra 0 0 0 0 0 0 0 0 0 0 0 0 0

6 Avellanita 0 0 0 0 0 0 0 0 0 0 0 0 0

7 Bakerolimon 0 0 0 0 0 0 0 0 0 0 0 0 0

8 Balsamocarpon 0 0 0 0 0 0 0 0 0 0 0 0 0

9 Bridgesia 0 0 0 0 0 0 0 0 0 0 0 0 0

10 Calopappus 0 0 0 0 0 0 0 0 0 0 0 0 0

11 Cissarobryon 0 0 0 0 0 0 0 0 0 0 0 0 0

12 Conanthera 0 0 0 0 0 0 0 0 0 0 0 0 0

13 Copiapoa 0 0 0 0 0 0 0 0 0 0 0 0 0

14 Cyphocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0

15 Desmaria 1 1 0 1 0 0 0 0 0 0 0 0 0

16 Dinemagonum 0 0 0 0 0 0 0 0 0 0 0 0 0

17 Dinemandra 0 0 0 0 0 0 0 0 0 0 0 0 0

18 Epipetrum 0 0 0 0 0 0 0 0 0 0 0 0 0

19 Ercilla 1 1 0 1 1 1 1 1 1 0 0 0 1

20 Fascicularia 0 1 0 1 1 0 1 1 0 0 0 0 0

21 Francoa 1 1 1 1 1 1 1 1 0 1 0 0 0

22 Gethyum 0 0 0 0 0 0 0 0 0 0 0 0 0

23 Gomortega 0 0 0 0 0 0 0 0 0 0 0 0 0

24 Guynesomia 0 0 0 0 0 0 0 0 0 0 0 0 0

25 Gymnachne 0 1 0 0 0 0 0 0 0 0 0 0 0

26 Gypothamnium 0 0 0 0 0 0 0 0 0 0 0 0 0

27 Hollermayera 1 0 0 0 1 0 0 0 0 0 0 0 0

28 Homalocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0

29 Huidobria 0 0 0 0 0 0 0 0 0 0 0 0 0

30 Hymenoglossum 0 1 1 1 1 1 1 1 1 1 1 1 0

31 Ivania 0 0 0 0 0 0 0 0 0 0 0 0 0

32 Jubaea 0 0 0 0 0 0 0 0 0 0 0 0 0

33 Lapageria 1 1 0 1 1 0 0 0 0 0 0 0 0

34 Latua 0 0 0 1 0 1 0 1 0 0 0 0 0

35 Legrandia 0 0 0 0 0 0 0 0 0 0 0 0 0

36 Leontochir 0 0 0 0 0 0 0 0 0 0 0 0 0

37 Leptocarpha 0 1 1 1 1 0 0 0 0 0 0 0 0

38 Leucocoryne 0 1 0 0 0 0 0 0 0 0 0 0 0

39 Leunisia 0 0 0 0 0 0 0 0 0 0 0 0 0

40 Marticorenia 0 0 0 0 0 0 0 0 0 0 0 0 0

41 Metharme 0 0 0 0 0 0 0 0 0 0 0 0 0

42 Microphyes 0 0 0 0 0 0 0 0 0 0 0 0 0

43 Miersia 0 0 1 0 0 0 0 0 0 0 0 0 0

44 Miqueliopuntia 0 0 0 0 0 0 0 0 0 0 0 0 0

45 Moscharia 0 0 0 0 0 0 0 0 0 0 0 0 0

46 Neuontobothrys 0 0 0 0 0 0 0 0 0 0 0 0 0

47 Notanthera 1 1 0 0 1 0 0 1 0 0 0 0 0

48 Ochagavia 1 0 0 0 0 0 0 0 0 0 0 0 0

49 Oxyphyllum 0 0 0 0 0 0 0 0 0 0 0 0 0

50 Peumus 0 1 0 1 1 0 0 0 0 0 0 0 0

51 Phrodus 0 0 0 0 0 0 0 0 0 0 0 0 0

52 Pintoa 0 0 0 0 0 0 0 0 0 0 0 0 0

53 Pitavia 0 0 0 0 0 0 0 0 0 0 0 0 0

54 Placea 0 0 0 0 0 0 0 0 0 0 0 0 0

55 Pleocarphus 0 0 0 0 0 0 0 0 0 0 0 0 0

56 Podanthus 0 0 0 0 0 0 0 0 0 0 0 0 0

57 Reicheella 0 0 0 0 0 0 0 0 0 0 0 0 0

58 Sarmienta 0 1 0 1 1 1 0 1 0 1 0 0 0

59 Scyphanthus 1 0 0 0 0 0 0 0 0 0 0 0 0

60 Speea 0 0 0 0 0 0 0 0 0 0 0 0 0

61 Tecophilaea 0 0 0 0 0 0 0 0 0 0 0 0 0

62 Tetilla 0 0 0 0 0 0 0 0 0 0 0 0 0

63 Traubia 0 0 0 0 0 0 0 0 0 0 0 0 0

64 Trevoa 0 0 0 0 0 0 0 0 0 0 0 0 0

65 Valdivia 0 1 0 1 0 0 0 0 0 0 0 0 0

66 Vestia 1 1 0 0 0 0 0 1 0 0 0 0 0

67 Zephyra 0 0 0 0 0 0 0 0 0 0 0 0 0

TOT 9 15 4 11 10 5 4 8 2 3 1 1 1

��0

N° GENERO AC co AC an AD co AD in AD an AE co AE an AF co AF an AG co AG an AH co AH an AI co AI an

1 Acrisione 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

2 Adenopeltis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

3 Alona 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

4 Anisomeria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

5 Araeoandra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

6 Avellanita 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

7 Bakerolimon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

8 Balsamocarpon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

9 Bridgesia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

10 Calopappus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

11 Cissarobryon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

12 Conanthera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

13 Copiapoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

14 Cyphocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

15 Desmaria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

16 Dinemagonum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

17 Dinemandra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

18 Epipetrum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

19 Ercilla 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

20 Fascicularia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

21 Francoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

22 Gethyum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

23 Gomortega 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

24 Guynesomia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

25 Gymnachne 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

26 Gypothamnium 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

27 Hollermayera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

28 Homalocarpus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

29 Huidobria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

30 Hymenoglossum 1 0 1 1 1 0 0 0 0 1 0 0 0 0 0

31 Ivania 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

32 Jubaea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

33 Lapageria 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

34 Latua 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

35 Legrandia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

36 Leontochir 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

37 Leptocarpha 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

38 Leucocoryne 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

39 Leunisia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

40 Marticorenia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

41 Metharme 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

42 Microphyes 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

43 Miersia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

44 Miqueliopuntia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

45 Moscharia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

46 Neuontobothrys 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

47 Notanthera 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

48 Ochagavia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

49 Oxyphyllum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

50 Peumus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

51 Phrodus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

52 Pintoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

53 Pitavia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

54 Placea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

55 Pleocarphus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

56 Podanthus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

57 Reicheella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

58 Sarmienta 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

59 Scyphanthus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

60 Speea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

61 Tecophilaea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

62 Tetilla 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

63 Traubia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

64 Trevoa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

65 Valdivia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

66 Vestia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

67 Zephyra 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

TOT 1 1 1 1 1 0 0 0 0 1 0 0 0 0 0

���

N° GENERO AJ co AJ in AJ an AK co AK in AK an AL co AL an AM co AM an SUM

1 Acrisione 0 0 0 0 0 0 0 0 0 0 14

2 Adenopeltis 0 0 0 0 0 0 0 0 0 0 7

3 Alona 0 0 0 0 0 0 0 0 0 0 10

4 Anisomeria 0 0 0 0 0 0 0 0 0 0 15

5 Araeoandra 0 0 0 0 0 0 0 0 0 0 2

6 Avellanita 0 0 0 0 0 0 0 0 0 0 2

7 Bakerolimon 0 0 0 0 0 0 0 0 0 0 6

8 Balsamocarpon 0 0 0 0 0 0 0 0 0 0 4

9 Bridgesia 0 0 0 0 0 0 0 0 0 0 9

10 Calopappus 0 0 0 0 0 0 0 0 0 0 4

11 Cissarobryon 0 0 0 0 0 0 0 0 0 0 7

12 Conanthera 0 0 0 0 0 0 0 0 0 0 19

13 Copiapoa 0 0 0 0 0 0 0 0 0 0 10

14 Cyphocarpus 0 0 0 0 0 0 0 0 0 0 7

15 Desmaria 0 0 0 0 0 0 0 0 0 0 8

16 Dinemagonum 0 0 0 0 0 0 0 0 0 0 7

17 Dinemandra 0 0 0 0 0 0 0 0 0 0 9

18 Epipetrum 0 0 0 0 0 0 0 0 0 0 9

19 Ercilla 0 0 0 0 0 0 0 0 0 0 19

20 Fascicularia 0 0 0 0 0 0 0 0 0 0 10

21 Francoa 0 0 0 0 0 0 0 0 0 0 19

22 Gethyum 0 0 0 0 0 0 0 0 0 0 5

23 Gomortega 0 0 0 0 0 0 0 0 0 0 3

24 Guynesomia 0 0 0 0 0 0 0 0 0 0 3

25 Gymnachne 0 0 0 0 0 0 0 0 0 0 7

26 Gypothamnium 0 0 0 0 0 0 0 0 0 0 2

27 Hollermayera 0 0 0 0 0 0 0 0 0 0 2

28 Homalocarpus 0 0 0 0 0 0 0 0 0 0 13

29 Huidobria 0 0 0 0 0 0 0 0 0 0 14

30 Hymenoglossum 0 0 0 0 0 0 0 0 0 0 18

31 Ivania 0 0 0 0 0 0 0 0 0 0 1

32 Jubaea 0 0 0 0 0 0 0 0 0 0 5

33 Lapageria 0 0 0 0 0 0 0 0 0 0 11

34 Latua 0 0 0 0 0 0 0 0 0 0 3

35 Legrandia 0 0 0 0 0 0 0 0 0 0 2

36 Leontochir 0 0 0 0 0 0 0 0 0 0 2

37 Leptocarpha 0 0 0 0 0 0 0 0 0 0 10

38 Leucocoryne 0 0 0 0 0 0 0 0 0 0 22

39 Leunisia 0 0 0 0 0 0 0 0 0 0 3

40 Marticorenia 0 0 0 0 0 0 0 0 0 0 2

41 Metharme 0 0 0 0 0 0 0 0 0 0 2

42 Microphyes 0 0 0 0 0 0 0 0 0 0 18

43 Miersia 0 0 0 0 0 0 0 0 0 0 6

44 Miqueliopuntia 0 0 0 0 0 0 0 0 0 0 3

45 Moscharia 0 0 0 0 0 0 0 0 0 0 9

46 Neuontobothrys 0 0 0 0 0 0 0 0 0 0 1

47 Notanthera 0 0 0 0 0 0 0 0 0 0 15

48 Ochagavia 0 0 0 0 0 0 0 0 0 0 12

49 Oxyphyllum 0 0 0 0 0 0 0 0 0 0 3

50 Peumus 0 0 0 0 0 0 0 0 0 0 17

51 Phrodus 0 0 0 0 0 0 0 0 0 0 10

52 Pintoa 0 0 0 0 0 0 0 0 0 0 2

53 Pitavia 0 0 0 0 0 0 0 0 0 0 3

54 Placea 0 0 0 0 0 0 0 0 0 0 7

55 Pleocarphus 0 0 0 0 0 0 0 0 0 0 5

56 Podanthus 0 0 0 0 0 0 0 0 0 0 15

57 Reicheella 0 0 0 0 0 0 0 0 0 0 2

58 Sarmienta 0 0 0 0 0 0 0 0 0 0 11

59 Scyphanthus 0 0 0 0 0 0 0 0 0 0 12

60 Speea 0 0 0 0 0 0 0 0 0 0 4

61 Tecophilaea 0 0 0 0 0 0 0 0 0 0 5

62 Tetilla 0 0 0 0 0 0 0 0 0 0 7

63 Traubia 0 0 0 0 0 0 0 0 0 0 4

64 Trevoa 0 0 0 0 0 0 0 0 0 0 12

65 Valdivia 0 0 0 0 0 0 0 0 0 0 2

66 Vestia 0 0 0 0 0 0 0 0 0 0 12

67 Zephyra 0 0 0 0 0 0 0 0 0 0 6

TOT 0 0 0 0 0 0 0 0 0 0

���

���

Appendix D: Native Genera in the Chilean Pacific Islands

��4

Islas Desventuradas Isla de Pascua Juan Fernández Juan Fernández (cont.)

Atriplex Agrostis Abrotanella LeptophyllochloaChenopodium Apium Acaena LibertiaCristaria Asplenium Adiantum LobeliaEragrostis Axonopus Agrostis LophosoriaFrankenia Davallia Apium LuzulaFuertesimalva Diplazium Arthropteris LycopodiumLepidium Doodia Asplenium MachaerinaLycapsus Dryopteris Azara MargyricarpusMaireana Elaphoglossum Berberis MegalachneNesocaryum Ipomoea Blechnum MegalastrumParietaria Juncus Boehmeria MimulusPlantago Kyllinga Bromus MyrceugeniaSanctambrosia Lycium Calystegia MyrteolaSicyos Microlepia Cardamine NassellaSolanum Microsorum Carex NerteraSpergularia Ophioglossum Centaurodendron NicotianaSuaeda Paspalum Centella NotantheraTetragonia Polystichum Chenopodium NotholaenaThamnoseris Psilotum Chusquea Ochagavia

Pycreus Colletia OphioglossumRytidosperma Coprosma OreobolusSamolus Cuminia ParietariaScirpus Cyperus PeperomiaSolanum Cystopteris PiptochaetiumSophora Danthonia PlantagoStipa Dendroseris PleopeltisThelypteris Dichondra PodophorusTriumfetta Dicksonia PolypodiumVittaria Drimys Polypogon

Dysopsis PolystichumElaphoglossum PterisEleocharis RanunculusEmpetrum RhaphithamnusErigeron RobinsoniaEryngium RubusEscallonia RumohraEuphrasia SantalumGalium SarcocorniaGamochaeta ScirpusGaultheria SelkirkiaGavilea SerpyllopsisGleichenia SolanumGrammitis SophoraGreigia SpergulariaGunnera TaraxacumHaloragis ThyrsopterisHedyotis TrichomanesHistiopteris TrisetumHymenoglossum UgniHymenophyllum UnciniaHypolepis UrticaJuania WahlenbergiaJuncus xMargyracaenaLactoris YunqueaLagenophora Zanthoxylum

���

Agradecimientos / Danksagung / Aknowledgements

Quiero, en primer lugar, dar las gracias a Paola Soto, mi mujer; sin ella ni este trabajo ni nada hubiese sido posible. A ella se lo dedico. Y a mi pequeña flor, mi pequeño roble, y mi pequeño amor.

Agradezco también a Mélica Muñoz, no sólo por ser mi madre que ya es mucho, sino por compartir siempre sus inagotables conocimientos botánicos y por haber sido desde que tengo memoria el pilar fundamental de mi formación naturalista. A través de ella no puedo dejar de mencionar a mi abuelo Carlos Muñoz y su esposa Ruth, quienes aún nos iluminan en los momentos de desconcierto. A través de ellos he tenido la suerte de conocer a Calvin y Linda Heusser, quienes me iniciaron en los intrincados caminos de la ciencia.

Vaya mi gratitud a mi padre Sergio, no sólo por ser mi padre sino por haberme impulsado siempre a viajar en busca de nuevos horizontes. Mis hermanos han colaborado cada uno a su manera: con Tiarella compartimos el nervio de una tesis, Iván ha enviando imprescindible música y videos; Simón ha descuidando su propia tesis para conseguir literatura ‚indispensable‘.

Federico Luebert ha revisado la tesis en forma exhaustiva y crítica, y ha sido junto con Patricio Pliscoff siempre partícipe de una discusión enriquecedora. Espero contar con su amistad por muchos años (viva Berlín!) También Sergio Elórtegui ha sido un gran compañero en la distancia, mágico creador de ilustraciones e ilusiones.

Agradezco también la amistad y la compañía musical de Carlos Ledermann, y el duende de los Mártires del Compás; sin él hubiese perdío el rumbo hace tiempo.

Meinem Doktorvater, Herrn Prof. Dr. Michael Richter, danke ich für die engagierte Betreuung der Arbeit am Anfang und dafür, mir am Ende die notwendigen Freiräume gewährt zu haben. Frau Prof. Dr. Perdita Pohle und Herrn Prof. Dr. Achim Bräuning danke ich herzlich für die Bereitschaft, bei meinem Rigorosum den Prüfungsvorsitz übernommen bzw. mich geprüft zu haben.

Dank dafür gebührt natürlich auch Herrn Prof. Dr. Werner Nezadal, meinem Prüfer im Nebenfach; ihm verdanke ich auch, viele Pflanzen am Cabo de Gata (Spanien) kennengelernt zu haben. Walter Welß hieß mich in der Mitte seiner umfangreichen Bibliothek im Botanischen Garten willkommen; langsam sind viele Bücher auch zu mir gewandert... Er ist in Teilen verantwortlich für die Ergebnisse meiner Arbeit (zumindest die guten). Herrn Prof. Dr. Tod Stuessy (Wien) danke ich für die Übernahme des Zweitgutachtens.

Lieben Dank an Sabine Donner für die stetige Hilfe in allen bürokratischen Fragen und danke allen Mitarbeitern des Instituts für Geographie für die freundliche Atmosphäre und die Möglichkeit zum fachlichen Austausch!

���

Besonders danke ich dabei meinen Mit-Doktoranden (viele sind mittlerweile schon promoviert): Luisa Vogt, Hendrik Wagenseil, Natalie Schulz, Cristina Dall‘ Ozzo, Markus Pingold, Michaela Ise, Paul Emck, Daniel Lingenhöhl, Henning Schröder, Tobias Bolch, Frieda Grüninger und Tom Fickert. Klaus Geiselhart verdient einen speziellen Dank für seine grundlegende Erfindung: den Geofanten.

Auch Alex Brenning muss ich hier erwähnen, obwohl ich ihn nicht leiden kann (…nur ein Scherz!). Und von einer erlebnisreichen Exkursion nach Tunesien bleibt mir die gute Freundschaft mit den Studierenden Viktor Kollmannsberger, Eva Neubauer, Matthias Patrzek und vor allem Ales Macik erhalten. Es war ein Glück Euch kennenzulernen!

Die sehr hilfreiche Englisch-Korrektur der Dissertation übernahm Iris Burchardt. Nochmals vielen Dank! Marian Jüngling gehört zwar nicht zur Geographie, doch war seine sprachliche Hilfe entscheidend für den guten Verlauf des Rigorosums.

Speziellen Dank an Thomas Sokoliuk für seine langjährige Unterstützung und seine unerschöpfliche Feststimmung (danke für den Fisch, Thomas!) Bei Rolf Kastner möchte ich mich unter anderem für die Überlassung seines Computers bedanken und bei Stefan Adler für die große Hilfe beim Einleben in Deutschland im ersten Jahr. Irma Richter, Anette Welß, Arietta Eberstadt sowie Coni Augustin sorgten auch für das gute Eingewöhnen und stets für das seelische und leibliche Wohl.

Bei Frau Dr. Brigitte Perlick, Akademisches Auslandsamt der Universität, möchte ich mich für ihre unendliche Energie bedanken und bei Frau Merker vom Promotionsbüro für den besten Rat in einer schwierigen Situation.

Schließlich danke ich dem DAAD für die gewährte finanzielle Unterstützung. Besonders der zuständigen Referentin Maria Hartmann gebührt für die schnelle Lösung unlösbarer Probleme mein herzlicher Dank. Der Zantner-Busch Stiftung gilt auch mein Dank für die finanzielle Unterstützung des Besuchs verschiedener Kongresse und Seminare.

Malte Ebach, Michael Heads, and Dennis McCarthy contributed with the review and edition of several papers. I hope our friendship grows and never suffers vicariance.

S. Liede, S. Renner, P. Endress, V. Funck, M. Gandolfo, M. Griffith, and J.J. Morrone collaborated providing important papers. Of course I’m alone responsible for the interpretation of these papers.

Am Schluss möchte ich den Erlangern danken, die Stadt bot uns fast vier Jahre lang ein Zuhause. Und einen Ratschlag möchte ich zu guter Letzt geben: Nehmt alles nicht so ernst! Das Leben ist zu kurz dafür! Ich wünsche mir, Deutschland hätte die WM 2006 doch gewonnen… Die gute Stimmung hätte vielleicht ein paar Jahre angedauert.

��7

LEBENSLAUF

Persönliche DatenName: Andrés Moreira-MuñozEltern: Sergio Moreira / Mélica Muñoz-SchickGeburtsdatum: ��. Feb. �97� in Los Angeles, ChileFamilienstand: verheiratet mit Paola Soto aus Viña del Mar, ChileKinder: Sayén (4), Silene (�), Coyán (�/4)

Schule�97� - �988 Deutsche Schule, Santiago de ChileAbschluß: Prueba de Aptitud Académica (entspricht Abitur)

Studium�989 - �994 Studium der Geographie an der Universidad Católica de Chile

Abschluß: Diplom-Geograph. Thema der Diplomarbeit: „Naturschutzgebiete an der Küste Zentral Chiles“

�994 - �99� Studium der Angewandte Geographie an der Universidad Católica de Chile

Abschluß: Geógrafo Profesional. Thema der Abschlußarbeit:: „Umwelt-Erziehung im Natur-Reservat Yerba Loca, Andes Zentral Chile“

Promotion�00� - �007 Institut für Geographie, Friedrich Alexander Universität Erlangen - Nürnberg Thema: Plant Geography of Chile

Andrés Moreira-Muñoz

Erlangen, �4. Juni �007