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Plant Geography of Chile

An Essay on Postmodern Biogeography

Den Naturwissenschaftlichen Fakultten der Friedrich-Alexander-Universitt Erlangen-Nrnberg zur Erlangung des Doktorgrades

vorgelegt von Andrs Moreira-Muoz aus Los Angeles, Chile

Gedruckt mit Untersttzung des Deutschen Akademischen Austauschdienstes (DAAD)

Als Dissertation genehmigt von den Naturwissenschaftlichen Fakultten der Universitt Erlangen-Nrnberg

Tag der mndlichen Prfung: Vorsitzender der Promotioskommision: Erstberichterstatter: Zweitberichterstatter:

Donnerstag 4. Juni 007 Prof. Dr. Eberhard Bnsch Prof. Dr. Michael Richter Prof. Dr. Tod Stuessy

A Paola, Silene, Coyn y Sayn

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 ................................................ 7 2.2 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 ........................................................ 0 2.5 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......................................................................................... 77 3.5 South-temperate floristic element ................................................................................. 80 3.6 Endemic floristic element ............................................................................................. 8 3.7 Cosmopolitan floristic element ..................................................................................... 8 4 Biogeographic analysis ........................................................................................................ 89 4.. To be or not to be disjunct ............................................................................................ 90 4.2 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 Earths surface .......................................... 7 . Oceanic transgressions over the continental surface ..................................................  . The Andean uplift........................................................................................................ 8 .4 The last 0 000 years: surviving the Ice Age .............................................................. 9 5.5 Alternative palaeogeographies: against scientific consensus ...................................... 


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 ................................................................................ 7 7. 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 .................................................. 47 Appendix C: Matrix for PAE: Distribution of Endemic Genera .......................................  Appendix D: Native Genera in the Chilean Pacific Islands .............................................  Agradecimientos / Danksagung / Acknowledgements .........................................................  Lebenslauf ............................................................................................................................. 7

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 groen Herausforderung stellen, sich der Moderne und der Postmoderne anzupassen. Die vorliegende Arbeit analysiert die wichtigsten phytogeographischen Gegebenheiten in Chile unter besonderer Bercksichtigung der geographischen Verbreitung von Gattungen und ihrer globalen Beziehungen. Die Auswirkungen dieser Verhltnisse auf die Entwicklung der chilenischen Flora werden dabei besonders unter verschiedenen palogeographischen Szenarien diskutiert. Auerdem wird die Situation im Kontext der modernen und postmodernen Wissenschaftstheorie betrachtet. Das erlaubt eine Kontext-orientierte Synthese der chilenischen Pflanzengeographie und stellt den Ausgangspunkt fr weitere Forschungen in dieser bergangsdisziplin zwischen Geographie und Biologie. Schlagwrter: 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.

 Introduction

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

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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 biologytrained 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 Concepcin, in a plan that comprises six volumes, from which the second has been almost completed (Marticorena & Rodrguez 99-00). It seems to be also the time for a renewal of Reiches 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 98, 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 Humboldts Essai sur la gographie 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 (80) called the subject just Gographie botanique and his son Alphonse de Candolle (8) added the adjective [Gographie botanique] raisonn. Later Schrter wrote about genetische Pflanzengeographie oder Epiontologie (Schrter 9), while Meusel preferred Vergleichende Arealkunde or Geobotanik (Meusel 94), and Walter added the adjective floristisch-historische [Geobotanik] or reduced it to Arealkunde (Walter 94, Walter & Straka 970). Schmithsen 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 (Schmithsen 98). 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. Cains 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).



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 () = the residence, the distribution district.
Unter Chorologie verstehen wir die gesammte Wissenschaft von der rumlichen Verbreitung der Organismen, von ihrer geographischen und topographischen Ausdehnung ber die Erdoberflche. [] Im weitesten Sinne gehrt mithin die gesammte Geographie und Topographie der Thiere und Pflanzen hierher, sowie die Statistik der Organismen, welche diese Verbreitungs-Verhltnisse mathematisch darstellt (Haeckel 8, pp. 887).

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, 99). Suessenguth (98) 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 98). 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 (97), 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 0s 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

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

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 48 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 Walters Arealkunde (Walter 94, . 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 . Donats distribution maps in Hannig & Winkler (9-940)

arealkundliche school at Halle/Saale brought Arealkunde/Chorologie to its highest point, with the work Vergleichende Chorologie der Zentraleuropischen Flora, published in three volumes (Meusel et al. 9, 978; Meusel & Jger 99). From the 0s to 70s we can mention also the large mapping work of NW Europe from Swedish E. Hultn, and his Circumpolar plants (Hultn 90, 94, 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 90, Milbradt 97, Wel & Lindacher 994). The earlier maps were an auxiliary tool for solving more general problems about the relationships



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 & Schnfelder 988), of Bavaria (Schnfelder & 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 Zrich-Montpellier school (Braun-Blanquet) and phytosociology (Txen). 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 Bergmanns rule, Allens rule, and perhaps the most famous Rapoports 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 Rapoports rule is still
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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 & Lsch 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 Ebachs 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 Croizats panbiogeography (Croizat 98). For the description and cartography of distributions, the most suited concept is still the old-fashion term Chorography.

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1.3 Naming the plant world


The 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 Stuessys 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 00 BC - 40 Issues Pioneer Greek Theophrastus wrote several manuscripts dealing with plants, translated as Enquiry into Plants (Hort 9, quoted by Stuessy 00). Age of the Herbalists, practical knowledge against diseases Classification per se, searching for revealing Gods 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. The search for natural systems (e.g. Jussies Genera Plantarum, de Candolles Prodromus). Phylogenetic systematics, mainly enforced by Darwins Origin of Species (89). 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). Phenetic studies avoiding phylogenetic speculation and returning to a more natural approach, sometimes using numerical methods Resurgence of phylogenetics mainly based on cladistics and molecular information.

 

c. 40-0 0 - 7

4 

770-880 880-90

 7

90-980 980 onwards

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 9

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 nave 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 nave, 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 [99, p. 4] (Cassirer 9, as quoted by Griffiths 1974, p. 88). Poppers example is reproduced here as figure 1.3.

Figure 1.3 Similarities between simple geometrical figures. 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)

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 Darwins Origin of Species, and the consequent changes in biological theory. Woodger (1952) explained this change in biological theory: In the Linnean system of classification

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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 90s into the 90s), 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 Hennigs Grundzge einer Theorie der phylogenetischen Systematik into English (Hennig 90, 9).

Figure .4. David Maddisons graphical representation of the debate between cladistics and phenetics during the 80s. [http://david.bembidion.org/]

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 Hennigs work, but they were not impressed by the various rules Hennig set out for inferring phylogeny, nor by Hennigs 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).



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).

Box . Basics of cladistics

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

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.

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

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/glossphylo.html Understanding evolution: http://evolution.berkeley.edu/evolibrary/article/phylogenetics_0

The methods proposed by Hennig (90, 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.


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;


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 didnt 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 wasnt 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 APGs 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 Stdte gleicher Gre durch gerade Linien, einmal, um festzustellen, ob im Eisenbahn- oder Straennetz gewisse Regeln erkennbar seien, ob es regelhafte Verkehrsnetze gbe, zum anderen, um die Abstnde zwischen gleich groen Stdten zu messen. Dabei fllten sich meine Karten mit Dreiecken, oft gleichseitigen Dreiecken die Abstnde gleich groer Stdte untereinander waren also annhrend gleich --, die sich zu Sechsecken zusammenschlossen. Ich stellte weiter fest, da in Sddeutschland die kleinen Landstdte sehr oft sehr genau einen Abstand von  km voneinander haben. Mein Ziel war abgesteckt: Gesetze zu finden, nach denen Anzahl, Gre und Verteilung der Stdte bestimmt sind... (Christaller 98, 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
4

(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


C A B C D A B A C

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 vicariance. When formation of barriers or splitting up of areas triggered speciation (i.e. vicariance), all species 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 cladograms 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-



1.4.1 Vicariance biogeography v/s dispersalism Gareth Nelson and Norman Platnick developed this approach based on the work of Len 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 Ratzels 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 Hennigs 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 Hennigs 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 Platnicks 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 (18711930), K.P. Schmidt (89097), G.G. Simpson (90984), P.J. Darlington, Jr (90498) and G.S. Myers (9098), (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 Simpsons 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 98). 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 80s and 90s 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


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 Christallers 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-). . . . 4. . . 7. 8. 9. 0. . . Location of greatest differentiation of a type Location of dominance or greatest abundance of individuals Location of synthetic or closely related forms Location of maximum size of individuals Location of greatest productiveness Continuity and convergence of lines of dispersal Location of least dependence upon a restricted habitat Continuity and directness of individual variations radiating from the centre along highways of dispersal Direction indicated by geographical affinities 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 biogeography


Willi Hennig, the great theorist behind the current cladistics paradigm, explained the criterion of veracity of his phylogenetic approach, with the reconstruction of a fragmented map:
Suppose a geographer has obtained fragments of a topographic map of an unknown land. He will make every effort to reconstruct the map from the fragments. How can he succeed if the original map is unknown to him? He was not present when the map was torn up. The geographer must try to assign each fragment to its original place in the 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.

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a
Figure . Hennigs fragmented map as test of the criterion of veracity (Hennig 9)

In section 1.3 cladistics was briefly introduced, therefore Hennigs 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 pretectonic 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 & Moores 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 populationsThe dismissal of Earth process from historical and ecological biogeography may have a sociological, political and perhaps an emotional raison detre. 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)

8

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 hegemnico de la ciencia para denunciar sus excesos y exponer sus lmites (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).

9

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 Len Croizats analytical and synthetical biogeography: Panbiogeography (Croizat 90). In fact, Croizats 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). Croizats 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 Ferr 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 (Ferr 997). The reaction of Cotterill (999) for such a nave 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 disciplines scientific integrity (Cotterill 999). Ferrs 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
0

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 & Cotterills (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 valuefree 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 concerns0. 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 ancestordescendant 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 (00a) 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 (97) 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 anyones 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. (Fernndez-Armesto 997, Truth, p 8).

Following Fernndez-Armestos 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 divisin fitogeogrfica deber, entre tanto, considerarse como provisional o aproximada. Podrn expresarse opiniones, pero discutir divisiones, lmites o nomenclatura, o tratar de imponer ideas en forma dogmtica, ser 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 

2 Chile, a Remote Corner on Earth


Durante la Conquista, cuando los espaoles recorran Amrica buscando esos metales [plata y oro] y llevndose todo lo que encontraban al paso, Chile se consideraba el culo del mundo, porque comparado con las riquezas del resto del continente tena 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 world


The discovery of the Chilean plant world can be tracked back to the circum-navigation of Hernando de Magallanes and Sebastin 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 Muoz-Schick & Moreira-Muoz 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 Magallanes and his crew ( october 0) set the start point for the European exploration of South America. Magallanes gave the name Cabo de Las Vrgenes 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 canelo, Drimys winteri. The sailors early noted the properties of canelos bark and herbs like Cardamine and Apium against scurvy. The first notes from sailors refer of course the medicinal, nutritional and wooden properties of the new plants found.

Hiplito Ruiz and Jos Pavn 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 Pavn. 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.

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

The Jesuit Juan Ignacio Molina (1737-1829) is considered the first Chilean naturalist. After the expulsion of Chile of the Jesuit Company (78), Molina published in Italy his Compendio della storia geografica, naturale, e civile del Regno del Chile (Molina 77). This and coming works (Molina 78, 80) were for a long time the main source of knowledge on natural sciences of Chile. The illustration represents araucaria, the Chilean palm (Jubaea) and culn (Otholobium glandulosum). In 87 the German naturalist Eduard Poeppig 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. Endlicher), 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 nrdlichen Halbkugel schieden, einer der feierlichten und bedeutsamsten eines langen Reiselebens Fast will es dem Reisenden dnken als trete er mir dem Eintritte in eine neue Welt auch in ein neues Leben. (after Morawetz & Rsser 998). The illustration corresponds to Puya alpestris.

The English naturalist Joseph Dalton Hooker (87-9) accompanied the expedition to the South Pole of John Clark Ross (1839 - 1843). He was the first who collected intensively in the southern lands (Kerguelen, Tasmania, New Zealand, Falkland, Tierra del Fuego) and who noticed the close floristic relationships between these territories. His Botany of the Antarctic Voyage was published in several volumes between 844-80.

8

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


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 848 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, bounded by the Andes and the Pacific, was seen as in a map. The pleasure from the scenery, in itself beautiful, was heightened by the many reflections which arose from the mere view of the grand range, with its lesser parallel ones, and of the broad valley of Quillota directly intersecting the 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 89). After its Independence from the Spanish dominion in 80, the young Republic of Chile had to better know its natural resources. The French professor of natural sciences, Claude Gay (800-87), already living in Chile, was contracted by the government to do the first intensive scientific exploration of the territory. He intensively traveled between Copiap and Chilo (80 - 84) (Muoz Pizarro 944). Afterwards he published his masterpiece Historia Fsica y Poltica de Chile, which consists of 8 volumes (8 of botany) and  illustration atlases (one cultural and one physical) (Gay 84-84). This was a work without precedents in America to that date. The mission 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 77 new species are described. Illustration of Desfontainia spinosa (source: www.memoriachilena.cl)

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 got soon the collaboration of his son Federico Philippi (88-90), exploring vast regions in Chile and the Norte Grande, territories just annexed to Chile after the Pacific war. There were R.A. Philippi a great amount of collaborators who provided material to the Philippi. With the impulse and the botanical explorations of father and son, the amount of 77 species described to that date rises to more that 7497 species. Of the 70 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.

F. Philippi

9

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


Karl Reiche (80-99), 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 (Grndzuge der Pflanzenverbreitung in Chile, 907, translated to Spanish by Gualterio Looser as Geografa Botnica de Chile).

Federico Johow (89-9) arrived in Chile 889 to teach natural sciences at the Instituto Pedaggico, Instituto Nacional and Universidad de Chile. He is the author of the milestone Estudios sobre la flora de las Islas de Juan Fernndez, 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.

Francisco Fuentes (879-94) 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 Reiches Flora.

Ivan Murray Johnston (898-90), US American prominent botanist, studied the coastal desert flora and vegetation. His papers The coastal flora of the departments of Chaaral and Taltal, and The flora of the Nitrate Coast (99 a,b), are considered classical studies of the northern Chilean coast, where vegetation is so highly related to sporadic precipitacion due to El Nio events (Dillon et al. 00) and the fogs water content (MuozSchick et al. 00).

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 Fernndez 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.

40

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


Carl Skottsberg (880-9), the Swedish botanical eminence, president of the 7th International Botanical Congress in 90, is one of the most impressive botanists that explored the Chilean territory. He came first with the Swedish expedition to the South Pole between 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 Vegetationsverhltnisse lngs der Cordillera de los Andes sdlich von 41 Grad... (9) and his Natural History of Juan Fernandez and Easter Island (90-9) are till today the most complete works that treat these territories.

Carlos Muoz Pizarro (9-97), was a student of Francisco Fuentes. After a Guggenheim scholarship 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, Muoz Pizarro explored, together with his wife Ruth Schick, the principal European herbaria, carrying out the photographic registry of the type collections of Chilean plants. The acquired knowledge allowed him to publish several important books: Sinopsis de la Flora Chilena (1959, 2d ed. 1966), the principal synthetical work of the Chilean flora still in use; Flores Silvestres de Chile (9); Chile: Plantas en extincin (97).

Mutisia decurrens, from Sinopsis de la Flora Chilena (9)

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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 Villagrn (00) suggests to recognize this millenary task as a ciencia indgena. 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).
ch e Ly oph co y ph tes Eu yte ph s y Sp llop hy er m te M ato s on ph ilo y ph tes yt es P = te fe rid rn s F oph er yt ns es a .. .fe nd. .. rn al E lie up s or an gi at e fe rn s
x x x x x x x x x x x x x Monophyletic x x x x x x x x x x x x x x x x x x x x x x x x x x x x Paraphyletic

Monilophytes = ferns

Leptosporangiate ferns Horsetails Marattioid ferns Ophioglossoid ferns Wisk ferns

Euphyllophytes

Spermatophytes

Tracheophytes

Gnetophytes Conifers Ginkgo Cycads

Lycophytes

Angiosperms Quillworts Spikemosses Clubmosses

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

2.2.1 Floristic composition: major families and genera Reiches (907) statistics were composed of 4 families and 7 genera for the Chilean Flora (table .), as he critically revised previous Philippis 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 Reiches revision. Muoz Pizarro (9) in his Sinopsis de la Flora Chilena gave a number of 8 families and 90 native genera, and Marticorenas (990) checklist considered 7 families an 87 genera (table .).

Tr a
x x x x x x x x x x x x x

4

Amborellaceae Nymphaeaceae

angiosperms

Austrobaileyales
Chloranthaceae

Canellales 1/1 Piperales 3/3 magnoliids Laurales 4/6 Magnoliales Acorales Alismatales 7/15 Asparagales 10/38 Dioscoreales 1/2 Liliales 4/7 Pandanales Arecales 1/2 Poales 7/102 commelinids Commelinales Zingiberales
Ceratophyllales 1/1 Ranunculales 4/10 Proteales 1/4

monocots

eudicots

Gunnerales Caryophyllales Santalales Saxifragales

1/1 12/69 3/9 4/10 + Vitales 1/1

Berberidopsidales 2/2

Crossosomatales Geraniales 4/9 Myrtales 3/17 Celastrales 1/1 Malpighiales 10/20 Oxalidales 3/6 Fabales 2/25 Rosales 3/21 Cucurbitales 2/2 Fagales 2/2 Brassicales 4/28 Malvales 2/13 Sapindales 3/9 Cornales 2/8 Ericales 6/17 Garryales Gentianales 3/16 Lamiales 15/58 Solanales 2/29 Aquifoliales Apiales 3/ 26 Asterales 5/137 Dipsacales 1/3

core eudicots

eurosids I
+ Zygophyllales 2/7

rosids

eurosids II

asterids

+ unplaced Boraginaceae /13 + unplaced Icacinaceae /1

euasterids I

+ unplaced Desfontainiaceae /1 + unplaced Escalloniaceae /2

euasterids II

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

Table . Historical statistics for the Chilean native vascular plants Orders 9 Families 4 8 7 Genera 8 7 90* 87

F. Philippi 88 Reiche 907 Muoz Pizarro 9 Marticorena 990 * 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
4

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) ferns and fern allies gymnosperms monocots dicots TOTAL Orders 8    9 Families 4 4 0  79 Genera 0 9  9 8 Native species   880 , 4,

Table 2.3 Genera not found in continental Chile (* = family also absent from the continent)
Genus (family) Dicksonia (Dicksoniaceae)* Thyrsopteris (Dicksoniaceae)* Arthropteris (Oleandraceae)* Juania (Arecaceae) Machaerina (Cyperaceae) Megalachne (Poaceae) Podophorus (Poaceae) 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)

Juan Fernndez Islands

ferns and fern allies gymnosperms monocots

dicots

44

Desventuradas Islands

ferns and fern allies gymnosperms monocots dicots

Lycapsus (Asteraceae) Thamnoseris (Asteraceae) Nesocaryum (Boraginaceae) Sanctambrosia (Caryophyllaceae) Davallia (Davalliaceae)* Microlepia (Dennstaedtiaceae) Diplazium (Dryopteridaceae) Dryopteris (Dryopteridaceae) Microsorum (Polypodiaceae) Psilotum (Psilotaceae)* Vittaria (Vittariaceae)* Doodia (Blechnaceae) Kyllinga (Cyperaceae) Pycreus (Cyperaceae) Axonopus (Poaceae) Stipa (Poaceae) Triumfetta (Malvaceae) Ipomoea (Convolvulaceae)

Isla de Pascua

ferns and fern allies

gymnosperms monocots

dicots

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
Asteraceae Poaceae Apiaceae Brassicaceae Fabaceae Solanaceae Caryophyllaceae Cactaceae Cyperaceae Boraginaceae Rosaceae Verbenaceae Malvaceae Alliaceae Myrtaceae Amaranthaceae Iridaceae Campanulaceae Lamiaceae Plantaginaceae

FAM

N genera N Chilean species


 7 4    8       9 9 9 9 8 8 8 8 9 9  7 4  9 4 07 4 8 7 9  47 8 7 9 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 Per 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 Adesmia Oxalis Viola Haplopappus Poa Carex Solanum Calceolaria Berberis Astragalus Baccharis Valeriana Leucheria Dioscorea Cryptantha Chaetanthera Nolana Loasa Hypochaeris, Alstroemeria, Verbena 8  7 70 4    0 47 4 4 4 4 4 9 7   

2.2.2 Endemic families The Chilean flora comprises three monotypic or dytipic endemic families, Gomortegaceae, Francoaceae, and Lactoridaceae, this latter endemic to Juan Fernndez (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 Gomortegaceae Francoaceae Lactoridaceae Genera Gomortega Francoa Tetilla Lactoris N species     (Juan Fernndez)

Table 2.7 Subendemic families of the Chilean flora (+adjacent areas in Argentina/Peru)
Family Aextoxicaceae Misodendraceae Philesiaceae Malesherbiaceae Genera Aextoxicon Misodendrum Lapageria Philesia Malesherbia N species (Chile)  () 8 (8)  ()  () 4 (8)

4

Just as a brief comparison, Argentina harbours one endemic family (Halophytaceae) and 4 endemic genera (Zuloaga et al. 1999). The flora of Per, composed of more than 17.000 species, and around .400 native genera4, 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 (Jrgensen & Len-Yez 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 G.R. Treviranus 80 August Pyramus de Candolle 80 Schouw 8 Alphonse de Candolle 8 Grisebach 87 Engler 879-88 Drude 884 Drude 890 Diels 908 Good 947 (974) Mattick 94 8 Flor  4 4 4    N regions 0 4 4    7 7 4 Antarktische Flor le Chili and les terres Magellaniques South of 42 = Antarktisches Florenreich; 42 - 23 = Reich der Holzartigen Synanthereen (Compositae); North of 23 = Reich der Cactus und Piper Region , Le Chili, and region , la Patagonie, la terre de Feu et les iles Malouines (Falkland). Tropische Andean Flora = North to 23; Chilenisches bergangsgebiet = 23-34, Antarktisches Waldgebiet = 34 Sdamerikanisches Florenreich = to 41, Altoceanisches Florenreich = south from 41 South of 41 = Antarktisches Florenreich; North of 41 = Andines Florenreich Andines Florenreich and Antarktisches Florenreich Antarktisches Florenreich and Neotropisches Florenreich Antarctic kingdom, Patagonian region, South of 4; Neotropical kingdom, Andean region, North of 4. South = Antarktisches Florenreich; North = Neotropisches Florenreich

47

Takhtajan 9 Takhtajan 978 Cox 00 Morrone 00

   

7  

Antarctic kingdom, Patagonian region, South of 40; Neotropical kingdom, Andean region, North of 40. Holantarctic kingdom, Chile-Patagonian region, South of  to Antarctic peninsula and Malvinas Islands (Falkland Is.) Southamerican kingdom Austral kingdom, Andean region

2.3.1. The global classification Gottfried R. Treviranus (77-87), 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 (740-80), J. Banks (74-80), and G. Forster (74-794). 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 franaise (Ebach & Goujet 00). A-P. de Candolle, in his Gographie 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 Candolles 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 40S 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; . of the genera had to be endemic or have their maximal distribution there; . A plant realm had to have some endemic families. De Candolles 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

48

floristic classification: Je tiens donc les divisions du globe par rgions, proposes jusqu present, pour des systmes 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 compltement diffrent de celui auquel mon pre pensait, car les documents etaient devenus plus nombreus, et mes ides staient singulirement loignees de celles qui rgnaient dans la science depuis vingt ans 7(A. de Candolle Mmoires, 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 (87-9). While sailing on board James Cooks Endeavour, he notably improved the floristic knowledge of the southern hemisphere. Hookers publications compiled as The Botany of the Antarctic Voyage (Hooker 844-80), were almost as epoch-making as Darwins 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 (84-879) to publish his Vegetation der Erde, which related explicitly the plant world with the regional climates (Grisebach 87). In Grisebachs view, Chile had to be classified into 3 regions =Gebiete: 1. a Chilean floristic core, Chilenisches bergangsgebiet (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 (844-90), 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 Sdamerikanische Florenreich (recognizing the Nordchilenische Provinz as the Chilenische bergangsgebiet from Grisebach) and in the Altoceanisches Florenreich, South of S. This Altoceanisches Florenreich grouped southern Chile with New Zealands South Island, the subAntarctic 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. ).

49

0

Figure .4 Vegetation of the world, after Grisebach (87, d edition 884)



Figure 2.5 Botanical regions and realms, floristic and physiognomic (Engler 1882)

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



Figure . Floristic realms from Drude (884)

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 4S corresponds to the Andines Florenreich and southern Chile to the Antarktisches Florenreich (figure 2.6). Drudes 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 Kppen (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 Drudes classification into six floristic realms, following the early proposal from Engler (88) but obviating the oceanic realm (Ozeanisches Florenreich), and dividing Englers 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) 

- and considered Malesia and New Zealand as part of the Palotropis. 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 (94) 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 & Lsch 004).

Figure .8 Floristic regions from Good (947)

English botanist Ronald Goods (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. Diels 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.
4

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 Takhtajans Holantarctic kingdom listing the  endemic families and 4 endemic genera proposed by Takhtajan (978). Takhtajans 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 Coxs 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 Englers early intention (879, 88). In fact the result is remarkably similar to Englers, but grouping the paleotropis and neotropis in one tropical realm. In Morrones 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.





Figure 2.10 Botanical realms from Takhtajan (1978). I = Holarctis, II = Paleotropis, III = Neotropis, IV = Capensis, V = Australis, VI = Holantarctis.

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 Reiches 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 Reiches 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 4S. In spite of appearing so simple, it is notably the first intend of a floristic cartography for Chile. 50 years later, Schmithsen drew a schematic representation of the different floristic elements composing the woody flora between 30 and 42S. (Schmithsen 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-Martnez & 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)

7

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

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

8

1. General floristic area of neotropical origin 2. Forest flora of south-hemispheric-subtropical origin . Laurophyllous neotropical and suth-hemispheric origin 4. Neotropical sclerophyllous element . Temperate rainforest of neotropical and south-hemispheric origin 6. Neotropical endemic flora from La Serena 7. Decoduous forest of subantarctic origin 8. Evergreen forest of subantarctic origin

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

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

9

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 (Kppen 90), followed by bioclimatic proposals for the country (di Castri 98, di Castri & Hajek 1976, Quintanilla 1974, Amigo & Ramrez 1998), the vegetation cartography has developed in hand of Schmithsen (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 90a, 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 Skottsbergs three volumes on the Natural History of Juan Fernndez and Easter Island (Skottsberg 90-9), 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 . Schmithsens vegetation map (9): 1= north-Andean vegetation; 2= desert; 3= semi-desertic scrub; 4= fog-forest; 5= xeric scrub; 6= sclerophyllous matorral; 7= temperate forest; 8a= Valdivian rainforest; 8b= northpatagonic rainforest; 8c=subantarctic rainforest; 9= tundra; 10=subantarctic deciduous-forest, 11=east-patagonic steppe; 12=south-Andean vegetation. 0

Figure .7 Phytogeographic map from Skottsberg (90a)



Figure .8 Vegetationsbild from Skottsberg (90 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 Muoz-Schick & Moreira-Muoz 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, Henrquez et al. 1995, Marticorena et al. 1998a, 2001) and local floras and checklists (e.g. Muoz-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, Villagrn 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, Ozire, 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 ().



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Myricaceae Balanophoraceae Cleomaceae Krameriaceae Nyctaginaceae Cactaceae Malesherbiaceae Cucurbitaceae Papaveraceae Malpighiaceae Hyacinthaceae Rafflesiaceae Thelypteridaceae Hydrocharitaceae Zygophyllaceae Anacardiaceae Verbenaceae Tecophilaeaceae Typhaceae Aizoaceae Acanthaceae Lythraceae Valerianaceae Amaryllidaceae Ledocarpaceae Pteridaceae Zosteraceae Loasaceae Azollaceae Aristolochiaceae Solanaceae Amaranthaceae Hemerocallidaceae Portulacaceae Alstroemeriaceae Fabaceae Caricaceae Malvaceae Equisetaceae Apocynaceae Bignoniaceae Caryophyllaceae Sapindaceae Laxmanniaceae Asteraceae Dioscoreaceae Plumbaginaceae Orobanchaceae Passifloraceae Oleaceae Sapotaceae Convolvulaceae Bromeliaceae Vivianiaceae Arecaceae Poaceae Salicaceae Lamiaceae Polygonaceae Piperaceae Brassicaceae Phrymaceae Euphorbiaceae Marsileaceae Scrophulariaceae Linaceae Boraginaceae Quillajaceae Polemoniaceae Alliaceae Molluginaceae Icacinaceae Geraniaceae Gentianaceae Campanulaceae Gratiolaceae Rosaceae Ceratophyllaceae Urticaceae Ephedraceae Goodeniaceae Monimiaceae Francoaceae Polypodiaceae Onagraceae Dryopteridaceae Iridaceae Lauraceae Apiaceae Agavaceae Loranthaceae Juncaceae Phytolaccaceae Rutaceae Vitaceae Gomortegaceae Frankeniaceae Violaceae Crassulaceae Oxalidaceae Griseliniaceae Aextoxicaceae Elatinaceae Calceolariaceae Potamogetonaceae Haloragaceae Rhamnaceae Berberidopsidaceae Polygalaceae Elaeocarpaceae Alismataceae Orchidaceae Lardizabalaceae Calyceraceae Escalloniaceae Aspleniaceae Ruppiaceae Rubiaceae Myrsinaceae Linderniaceae Hypericaceae Araucariaceae Myrtaceae Cyperaceae Coriariaceae Gesneriaceae Tropaeolaceae Lemnaceae Hydrangeaceae Plantaginaceae Atherospermataceae Berberidaceae Dennstaedtiaceae Proteaceae Lentibulariaceae Lophosoriaceae Ranunculaceae Santalaceae Corsiaceae Juncaginaceae Cupressaceae Philesiaceae Celastraceae Lomariopsidaceae Cunoniaceae Podocarpaceae Grossulariaceae Gunneraceae Winteraceae Blechnaceae Araliaceae Restionaceae Hymenophyllaceae Desfontainiaceae Samolaceae Nothofagaceae Misodendraceae Schizaeceae Luzuriagaceae Isoetaceae Ericaceae Thymelaeaceae Gleicheniaceae Grammitidaceae Droseraceae Centrolepidaceae Asteliaceae Saxifragaceae Ophiglossaceae Stylidiaceae Primulaceae Lycopodiaceae Tetrachondraceae

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 7S (Mediterranean-type zone) (figure 2.21); and 37 to 56S (temperate zone) (figure 2.22).

Figure .9 Latitudinal ranges for Chilean families 4

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Stangea Mniodes Dissanthelium Cremolobus Stenomesson Dielsiochloa Coreopsis Alchemilla Achyrocline Woodsia Chloris Spilanthes Tara Trichoneura Tecoma Neowerdermannia Dunalia Pityrogramma Pluchea Browningia Morella Haplorhus Lophopappus Salpichroa Haageocereus Distichia Trixis Eremodraba Polylepis Corryocactus Anthochloa Tunilla Plazia Neuontobothrys Ombrophytum Gomphrena Reicheella Diplostephium Heterosperma Philibertia Oreocereus Metharme Luciliocline Mastigostyla Eudema Chersodoma Helogyne Xenophyllum Lampaya Tripogon Aa Munroa Cuatrecasasiella Cotula Bouteloua Exodeconus Leptochloa Parastrephia Urbania Enneapogon Pycnophyllum Mancoa Stevia Villanova Dalea Allionia Urmenetea Notholaena Nasa Anatherostipa Palaua Cleome Aphyllocladus Nitrophila Nama Alternanthera Portulaca Lycopersicon Tiquilia Sporobolus Salvia Drymaria Tigridia Domeykoa Werdermannia Ipomopsis Balbisia Acantholippia Huidobria Microsteris Eremocharis Dinemandra Fuertesimalva Reyesia Maihueniopsis Schkuhria Erechtites Zephyra Encelia Geoffroea Dicliptera Caesalpinia Croton Bacopa Fagonia Perityle Eulychnia Grabowskia Wedelia Oxyphyllum Cumulopuntia Aloysia Krameria Gypothamnium Deuterocohnia Zameioscirpus Rostraria Werneria Hoffmannseggia Orobanche Cynodon Verbesina Doniophyton Oxychlo Pitraea Polyachyrus Nototriche Malacothrix Amblyopappus Tetragonia Copiapoa Lenzia Prosopis Lycium Chuquiraga Bakerolimon Echinopsis Malesherbia Pennisetum Sicyos Cristaria Argemone

Figure .0 Latitudinal ranges for Chilean genera with average distribution north of 7S 

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Lippia Skytanthus Tagetes Ophryosporus Mirabilis Oziroe Alona Bartsia Bidens Ivania Tessaria Pilostyles Eriosyce Chorizanthe Argylia Maireana Mentzelia Errazurizia Oxytheca Flaveria Thelypteris Elodea Cressa Cardionema Cruckshanksia Leontochir Caiophora Paronychia Viguiera Tillandsia Eragrostis Pintoa Trichocline Cerastium Parietaria Cistanthe Monttea Aristida Plumbago Heliotropium Tarasa Bomarea Balsamocarpon Grindelia Salix Miqueliopuntia Boerhavia Phrodus Chaetanthera Bahia Bulnesia Lupinus Senna Spergularia Verbena Gymnophyton Typha Proustia Nolana Pellaea Malvella Homalocarpus Agalinis Dinemagonum Cestrum Kurzamra Glandularia Pleurophora Myrcianthes Calliandra Cyphocarpus Pectocarya Cuscuta Limonium Cordia Phragmites Centaurea Polycarpon Montiopsis Conanthera Galinsoga Puya Ambrosia Menonvillea Heterozostera Chiropetalum Muhlenbergia Linum Vulpia Azolla Anisomeria Ciclospermum Leucocoryne Aristolochia Araeoandra Guynesomia Bridgesia Ageratina Schizopetalon Brachyclados Patosia Calycera Pasithea Schizanthus Nicotiana Dodonaea Microphyes Helenium Pleocarphus Vasconcellea Epipetrum Equisetum Marsilea Microseris Tecophilaea Cicendia Cyperus Paspalum Asteriscium Flourensia Cryptantha Carpobrotus Leunisia Trichopetalum Gethyum Pachylaena Lastarriaea Zoellnerallium Llagunoa Fabiana Mathewsia Tropidocarpum Barneoudia Porlieria Tweedia Passiflora Lemna Cheilanthes Schinus Polypogon Menodora Solanum Lotus Distichlis Melica Convolvulus Rhodophiala Teucrium Trifolium Triodanis Pouteria Acacia Muehlenbeckia Facelis Andeimalva Traubia Gayophytum Bipinnula Dioscorea Triptilion Placea Larrea Viviania Laretia Eryngium Dichondra Aristeguietia Stachys Jubaea Tropaeolum Erodium Alonsoa Moscharia Trevoa Phycella Marticorenia Miersia Pleopeltis Diposis Mulinum Centaurium Mutisia Adiantum Haplopappus Pteromonnina Tetilla Guindilia Corrigiola Stemodia Salpiglossis Speea Sphaeralcea

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Adenopeltis Peperomia Melosperma Ligaria Mimulus Weberbauera Calydorea Nassella Stenandrium Micropsis Avellanita Psilocarphus Calopappus Quillaja Lithrea Dennstaedtia Gochnatia Podanthus Rhombolytrum Glinus Linanthus Margyricarpus Cryptogramma Gilia Kageneckia Lobelia Ludwigia Quinchamaliu Diplolepis Beilschmiedia Scyphanthus Buddleja Retanilla Citronella Downingia Piptochaetium Chaptalia Minuartia Lepechinia Otholobium Solaria Plectritis Ochagavia Wahlenbergia Oenothera Clarkia Blennosperma Wolffiella Pozoa Hydrocotyle Nothoscordum Bowlesia Colliguaja Ceratophyllum Junellia Camissonia Aphanes Cryptocarya Gilliesia Noticastrum Phalaris Artemisia Castilleja Mecardonia Cissarobryon Imperata Nastanthus Stuckenia Tristerix Loasa Astragalus Euphorbia Ephedra Panicum Peumus Selliera Jarava Picrosia Atriplex Descurainia Acrisione Escallonia Lapageria Hypochaeris Glycyrrhiza Lucilia Bromidium Daucus Utricularia Soliva Centipeda Lilaeopsis Legrandia Chenopodium Hypsela Jaborosa Gnaphalium Calotheca Wendtia Herreria Lardizabala Mikania Pilularia Pelletiera Satureja Calandrinia Lepuropetalon Conyza Leucheria Eleocharis Moschopsis Ranunculus Gamochaeta Phacelia Pitavia Austrocedrus Calystegia Gymnachne Cynoglossum Herbertia Cissus Phylloscirpus Arenaria Cystopteris Draba Gentiana Lepidium Limosella Silene Valeriana Vestia Adesmia Maihuenia Calceolaria Evolvulus Geranium Hordeum Eccremocarpu Bromus Sisymbrium Corynabutilon Urtica Gomortega Calamagrostis Alisma Gutierrezia Azorella Epilobium Wolffia Hybanthus Amaranthus Baccharis Colobanthus Juncus Perezia Senecio Frankenia Festuca Scirpus Helictotrichon Navarretia Blepharocalyx Stellaria Asplenium Catabrosa



Figure . Latitudinal ranges for Chilean genera with average distribution in Central Chile between 7S and 7S

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Dichanthelium Crassula Galium Oxalis Poa Viola Sagina Orites Griselinia Plantago Spartina Ourisia Brachystele Alstroemeria Aextoxicon Sarmienta Elatine Cortaderia Myriophyllum Danthonia Sarcocornia Apium Berberidopsis Aristotelia Pratia Diostea Tetraglochin Glyceria Luma Solidago Gamochaetopsis Elymus Crinodendron Sanicula Cliococca Amsinckia Suaeda Lilaea Triglochin Prumnopitys Dasyphyllum Sigesbeckia Notanthera Podagrostis Belloa Francoa Jovellana Desmaria Laurelia Chascolytrum Azara Nardophyllum Zannichellia Ruppia Chevreulia Rhaphithamnus Persea Puccinellia Polypodium Sagittaria Potamogeton Leptocarpha Colletia Polygonum Rhamnus Carex Gentianella Bulbostylis Rhynchospora Myrceugenia Lindernia Gratiola Hedyotis Hypericum Elytropus Chusquea Megalastrum Sophora Rorippa Araucaria Fascicularia Pleurosorus Relchela Leptophyllochloa Acaena Lysimachia Ercilla Centella Amphibromus Greigia Olsynium Boquila Coriaria Phytolacca Leptostigma Blumenbachia Lepidoceras Ugni Hollermayera Sisyrinchium Hypolepis Legenere Anagallis Antidaphne Valdivia Hydrangea Lomatia Austrocactus Vicia Habenaria Mitraria Gevuina Berberis Combera Solenomelus Callitriche Trisetum Libertia Fragaria Fonkia Eucryphia Lophosoria Rytidosperma Saxe-gothaea Amphiscirpus Amomyrtus Lasthenia Pteris Lathyrus Asteranthera Fitzroya Gamocarpha Trichomanes Arachnitis Polygala Pilea Anarthrophyllum Latua

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Myosurus Fuchsia Maytenus Elaphoglossum Discaria Ovidia Plagiobothrys Tristagma Anemone Antennaria Montia Phleum Caldcluvia Weinmannia Deschampsia Polemonium Euphrasia Madia Rumex Erigeron Laureliopsis Alopecurus Arjona Dysopsis Nassauvia Rumohra Osmorhiza Gunnera Ribes Nierembergia Chloraea Caltha Collomia Cardamine Luzula Gavilea Podocarpus Polystichum Drimys Koeleria Myoschilos Blechnum Aster Armeria Gaultheria Hymenophyllum Nertera Uncinia Hymenoglossum Agoseris Agrostis Oreopolus Pseudopanax Apodasmia Ophioglossum Campsidium Desfontainia Scutellaria Noccaea Tepualia Samolus Empetrum Nothofagus Geum Misodendrum Schizaea Taraxacum Huanaca Hieracium Lagenophora Boopis Luzuriaga Embothrium Saxifraga Codonorchis Isoetes Pinguicula Adenocaulon Schoenus Hippuris Chiliotrichum Myrteola Onuris Anthoxanthum Oreobolus Rubus Lycopodium Marsippospermum Pterocactus Magallana Schizeilema Gleichenia Macrachaenium Grammitis Donatia Drosera Pilgerodendron Primula Littorella Lepidothamnus Philesia Ortachne Serpyllopsis Gaimardia Leptinella Tribeles Histiopteris Astelia Nanodea Carpha Tapeinia Philippiella Abrotanella Tetroncium Lebetanthus Benthamiella Phyllachne Grammosperma Xerodraba Drapetes Eriachaenium Rostkovia Hebe Myosotis Tetrachondra Lecanophora Bolax Lepidophyllum Hamadryas Saxifragodes Chiliophyllum Oreomyrrhis Botrychium Androsace Saxifragella Chrysosplenium Huperzia Landoltia

7

Figure . Latitudinal ranges for Chilean genera with average distribution in the temperate zone south of 7S

8

 Geographic Relationships of the Chilean Flora

9

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

70

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 90, 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 Mxico (Qian 1999), the East Asian flora (Qian et al. 2003), the flora of the Bolivian Andean valleys (Lpez 00), and the Ecuadorian superpramo (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, Villagrn & 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)    4   7 Tropical-American Andes Chile/Argentina California/Mexican Antarctic Boreal (Europa, S Chile) Pantropic Introduced flora Villagrn & Hinojosa 997 Neotropical (amplio / disyunto) Endmico bosques subantrticos / Endmico de Chile Australasiano (clido/antrtico) Amplio Pantropical -

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, 98)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 . Pantropical . Australasiatic . Neotropical 4. Antitropical . South-temperate . Endemic 7. Cosmopolitan Total N Genera 88 9   8 8 4 8 %  7 7 9 0 0  00 %

Table . Generalized tracks for the vascular plant Chilean genera (update 07 February 007)
Tracks (follow subsections in the text) .. Austral-antarctic track 3.2.2 South Pacific tropical track .. Circum-austral track .. Wide Neotropical track .. Andean track .. South Amazonian track .4. Wide antitropical track 3.4.2 Antitropical disjunct Pacific track 3.4.3 Circum-Pacific track N of genera    4  9 84  9

3.1 Pantropical floristic element


Genera 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,

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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 90

Acacia (Fabaceae)

Bacopa (Gratiolaceae), sensu Dawson 98

Cotula (Asteraceae), Meusel & Jger 99, Bremer 994

Lobelia (Campanulaceae), sensu Meusel & Jger 99

Nicotiana (Solanaceae), sensu van Steenis & van Balgooy 9

Figure 3.1 Examples of the pantropical floristic element

3.2 Australasiatic floristic element


This 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.
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Azorella distribution, sensu Baumann 988

Berberidopsis distribution, sensu Ronse De Craene 004

Coprosma distribution, sensu van Steenis & van Balgooy 9

Hebe distribution, sensu van Steenis & van Balgooy9

Jovellana distribution, sensu Dawson 98, Heads 994

Lagenophora distribution, sensu Cabrera 9

Leptinella distribution, sensu Bremer 994

Oreomyrrhis distribution, sensu Mathias & Constance 9

Ourisia distribution, sensu Meudt & Simpson 00

Rytidosperma distribution, sensu Linder & Barker 000

Figure 3.2 Examples of the australasiatic floristic element 74

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 Fernndez 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 Fernndez Islands, also absent from the continent9. 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 element


Strictly 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

Alstroemeria

Azara

Calceolaria

Cremolobus

Eremodraba

Escallonia

?
Eudema Malesherbia Mathewsia

Mecardonia

Nototriche

Vasconcellea

Figures 3.3 Examples of the neotropical floristic element (distribution from monographs and checklists) 7

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 Per, 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/Per (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 Brazil It 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 element


This 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. Wel (pers. comm.) correctly noted, the most appropriate term would be antitropic, since amphitropical means both tropics. In Coxs (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).
77

Hydrangea distribution, sensu Mabberley 997

Antennaria distribution, sensu Meusel & Jger 99

Artemisia distribution, sensu Meusel & Jger 99

Euphrasia distribution, sensu Van Steenis 9, Dawson 98

E Asia

Osmorhiza, sensu Wood 97

Frankenia, sensu Wahlen 987

Mancoa, sensu Al-Shehbaz comm. pers.

Tropidocarpum, sensu Al-Shehbaz 00

Agoseris, sensu Wood 97

Larrea, sensu Hunziker et al. 97

Figures 3.4 Examples of the antitropical floristic element 78

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 90s (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.

79

Austrocedrus

Embothrium

Fonkia

Grammosperma

Hamadryas

Laureliopsis

Melosperma

Menonvillea

Monttea

Onuris

Pilgerodendron

Saxe-gothaea

Figure 3.5 South-temperate floristic element ((distribution from monographs and checklists) 80

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 element


This 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 Fernndez (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

Balsamocarpon

Bridgesia

Calopappus

Cissarobryon

Conanthera

Copiapoa

Cyphocarpus

Desmaria

Dinemagonum

Dinemandra

Epipetrum

Ercilla

Fascicularia

Francoa

Gethyum

Gomortega

Guynesomia

Gymnachne

Gypothamnium

Hollermayera Homalocarpus

8

JF

Huidobria

Hymenoglossum

Ivania

Jubaea

Lapageria

Latua

Legrandia

Leontochir

Leptocarpha

Leucocoryne

Leunisia

Marticorenia

Metharme

JF

Microphyes

Miersia

Miqueliopuntia

Moscharia

Neuontobothrys

Notanthera

8

JF

Ochagavia

Oxyphyllum

Peumus

Phrodus

Pintoa

Pitavia

Placea

Pleocarphus

Podanthus

Reicheella

Sarmienta

Scyphanthus

Speea

Tecophilaea

Tetilla

Traubia

Trevoa

Valdivia

Vestia

Zephyra

Figure 3.6 Endemic floristic element (continues)

84

3.7 Cosmopolitan floristic element


There 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).

Apium (Apiaceae), sensu Meusel et al. 978

Bidens (Asteraceae), sensu Meusel & Jger 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

Figure 3.7 Cosmopolitan floristic element 8

8

4 Biogeographic analysis

87

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

88

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 1. Pantropical 2. Australasiatic 3. Neotropical 4. Antitropical 5. South-temperate 6. Endemic 7. Cosmopolitan Average (S) -, -4, -9, -, -4, -, -,0

18

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 Schmithsen (9) to consider this region as a transition zone with different converging elements.

24

neotropical pantropical/ endemic antitropical cosmopolitan

30

36

australasiatic south-temperate

42

48

After grouping the Chilean plants in seven floristic 54 elements (Kontingente), Reiche (907, p. 9) proposed that the neotropical and antitropical (Reiches Californian) elements occur in Chile due to Figure 4.1 Average range for floristic elements 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

89

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, Reiches 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


Villagrn & 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) South Pacific tropical track (25) Circum-austral track (3) Panbiogeographic nodes

Wide Neotropical track (64) Andean (113) and South-Amazonian (39) tracks Wide Antitropical track (84) Pacific antitropical track (56) Circum-Pacific track (9)

Figure 4. Generalized tracks for the Chilean plant disjunct ditributions (in parenthesis the number of genera composing the tracks) 90

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 97a,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 99). c) long-distance dispersal, probably by migrating birds; from the 90s 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 90s 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 realm


As 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 47S (figure 2.17); b) Takhtajans (1978) newest view setting this line at around 23S (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 Morrones (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 ~47S 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 (Muoz 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) Coxs (2001) vision neglects or minimizes the australasiatic element in the American flora and therefore do not leave opportunity for comparisons. d) In defense of Morrones (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, 100 90 80 several genera are restricted to temperate 30 Chile and are absent in the northern part, but they reappear in the northern Andes, Podocarpus like Rumohra, Citronella, Prumnopitys, Weinmannia Nertera; some of them even reaching their Drimys/Centropogon 20 highest level of species richness in the northern Andes: e.g. Fuchsia, Weinmannia, and Uncinia. Gunnera So the question remains: which of the Escalllonia classifications fit better to the Chilean flora? With the analysis of the floristic elements in the latitudinal profile we can attempt to Figure 4. Northward extent of spread of Gondwana genera in tropical America (redrawn from Webster 99) answer the question. Since the latitudinal
9
Desfontainia/
10

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), Biobo (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. Rodrguez 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.

Islas Desventuradas Isla de Pascua Juan Fernndez

ANT COQ BIO MAG

Jaccard similarity index was applied to the data Figure 4.4 Chilean continental regions included set (Zunino & Zullini 00, Cox & Moore 00, in the similarity analysis and location of Chilean Pacific islands. 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 ANT COQ BIO MAG Regions Antofagasta Coquimbo Biobo Magallanes Area (km) .049 40.80 7.0 .0 N native vascular plant genera  47 4  Source Marticorena et al. 998a Marticorena et al. 00 CONC, R. Rodrguez pers. comm. Henrquez et al. 99

Table 4. Floristic similarity and geographic distance different regions in Chile: Antofagasta (ANT), Biobo (BIO), Coquimbo (COQ) and Magallanes (MAG). Geographic distance have been calculated as the latitudinal difference between the geographic centroid of each region.
Similarity Jaccard 0,4 0,9 0, 0,4 0,7 0,

Compared regions ANT/COQ ANT/BIO ANT/MAG COQ/BIO COQ/MAG BIO/MAG

Shared genera 44 74 97  0 78

Distance (km) 777 7 7 70 40 700

9

Jaccard similarity

0.6

0.8

1.0

COQ/BIO ANT/COQ ANT/BIO BIO/MAG COQ/MAG ANT/MAG

0.0

0.2

0.4

2000

4000

6000

Figure 8000 4. Similarity versus geographic 10000 distance between Chilean regions.

Distance [km]

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 ANT COQ BIO MAG
40,00

pantropical n 8 9 4  % 8,89 8,9 9,09 4,8

australasiatic n 8  4 8 % ,4 4,8 8,87 ,4

neotropical n  4 00 9 % , , , ,

antitropical n 4 7 7 44 % 4,9 ,08 ,8 7,

southtemperate n 7 4  4 % ,40 9, ,04 ,7

endemic n  8 7  % 4,7 8,7 ,84 0,40

cosmopolitan n 8 7  84 % ,0 ,77 , ,47

30,00

20,00

10,00

1. 2. 3. 4. 5. 6. 7.

Pantropical Australasiatic Neotropical Antitropical South-temperate Endemic Cosmopolitan

0,00

ANT

COQ Regions

BIO

MAG

Figure 4.6 Floristic elements present in four regional floras (percentage) 94

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 (80) 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 scaledependant, 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 Muoz-Schick & Moreira-Muoz (000).
50

40
N endemic genera

30

20

10

0 1 2 3 4 5 6 7 8 9 10+ N species per genus

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

9

-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

Fig. 4.8 Latitudinal ranges for the 7 Chilean endemic genera. Small distribution ranges on top, large towards the buttom. 9

Notanthera Francoa Ercilla Hymenoglossum

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. 17-18 The endemic genus with the widest latitudinal 19-20 21-22 distribution in Chile is Leucocoryne with a 23-24 distribution range of 9 latitudinal degrees from 25-26 27-28  to 40 S.
29-30 31-32

Latitude (S)

33-34 35-36 37-38 39-40 41-42 43-44 45-46 47-48 49-50 51-52 53-54 55-56 0 10 20 30 40

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

N genera

Fig. 4.9 Endemic genera at each latitude degree.

97

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.

Figure 4.0 Geographic ranges of endemic Chilean genera

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

Figure 4. Generic richness for each  x  km cell

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
98

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 40 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 (31-38S) = a central coastal floristic block; and a group of central Chilean Andean cells (33-39S) = a central Andean floristic block. A third group of southern cells including Chilo Island (44S) 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 Chilo 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
99

Majority rule tree (from 5 trees, cut 50) Hypothetical_area Aan Bco Ban Cco Can Dan Ean Fan Gin Gan Han Ian Kan AEco AEan AFco AFan AGan AHco AHan AIco AIan AJco AJin AJan AKco AKin AKan 100 ALco ALan AMco AMan Jan 100 Lan Eco Fco 60 Gco Hco 60 Ico Jco Dco Man 60 Nan 60 Oan 60 100 Pan 60 Kco Lco 100 Mco 80 Nco AAan ABco ACco ADco ADin ADan 80 AGco Zan 80 ABan 100 ACan Wan 100 AAco 100 Yco Yan 100 Zco 100 Xan 100 100 100 100

Xco

Vco Wco 80 80

Van 80

Tan 100

Uan 100

60 100

San

Uco 100 100

Qan Ran

Tco 100

Sco 100

Rco 100

Qco

Oco Pco

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. 00

Central coastal floristic block Central Andean floristic block Southern floristic block

Fig 4.4 Floristic blocks based on terminal cells from figure 4.13.

(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 77S (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 4S) seems to be very recent (Villa-Martnez et al. 00). This suggests, for Chile as well as for the European Mediterranean region (Verd 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
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 methods0. 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, Domnguez 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 elements


The 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 & Villagrn 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 relationship Chilean 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 breakup of Gondwana (Villagrn & 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 & Villagrn 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!)

Nemuaron (N Caledonia) Atherosperma (Australia / Tasmania) Laurelia sempervirens (Chile endemic) Dryadodaphne (E Australia / N Guinea) Daphnandra (E Australia) Doryphora (E Australia) Laureliopsis (Chile / Argentina) Laurelia novae-zelandiae (NZ) Gomortega (Chile endemic)

Source: Renner 2004

Figure 4. Phylogenetic and geographic relationships of Gomortega and Atherospermataceae

Ripogonum (Australia/N. Guinea/NZ) Lapageria (endemic Chile) Philesia (Chile/Argentina)

Source: Vinnersten & Bremer 2001

Figure 4. Phylogenetic and geographic relationships of Philesiaceae 0

Neotropical relationship Chilean 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 (Per/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 relationships The 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.
04

Table 4. Australasiatic relationships of South-temperate genera


Taxon Pilgerodendron (Cupressaceae) Lebetanthus Huanaca (Apiaceae) Philesia (+ endemic Lapageria) subfamily Coronantheroideae Phylogenetic relationships Genus from southern Chile/Argentina, closely related to Libocedrus (New Zealand); some authors propose to unite them in one genus. Epacridaceous genus from (MAG/Argentina) is closely related to Prionotes (Tasmania) Patagonian genus close related to New Zealand Stilbocarpa the only members of the South American Philesiaceae, are supposedly splitted from the AustralianNew Zealand Ripogonum at 47 +- 8,4 mya. 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. Source Gadek et al. 000, but see Farjon et al. 004 Crayn et al. 998 Andersson et al. 00 Vinnersten & Bremer 00 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 AustralianNew 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 Fernndez, 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.

Figure 4.7 Isla de Pascua (from Skottsberg 909).

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 (90-9) 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 Zizkas 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
0

41% 49%

1. Pantropical 2. Australasiatic 3. Neotropical 4. Antitropical 5. South-temperate 6. Endemic 7. Cosmopolitan

0% 0%

3%0%

7%

Figure 4.8 Floristic elements Isla de Pascua

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 (90-9). 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.

Figure 4.9 Isla San Flix. [www.cordell.org/


SFX/SFX_pages/SFX_Main.html]

Figure 4.0. Isla San Ambrosio (after Kuschel 9)

4.5.2 Islas Desventuradas, San Flix (2617 S / 800) and San Ambrosio (S / 79W) This relatively small volcanic archipelago is located approximately 80 km off the Chilean coast (figure 4.4.). Desventuradas Islands consist of the two main islands San Flix (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. km. The topography is very rugged, with peak elevations of 9 m asl on Isla San Flix, 479 m a.s.l. on Isla San Ambrosio.

07

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 (97, 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 Fernndez at the genus level. Furthermore, Marticorena (990) reports a level of endemism of 0,% at the species level, the highest for a Chilean region.

0% 5% 11%

11% 52% 0%

1. 2. 3. 4. 5. 6. 7.

Pantropical Australasiatic Neotropical Antitropical South-temperate Endemic Cosmopolitan

21%

Figure 4. Floristic elements Islas Desventuradas

The natural history of the archipele has been analysed by Skottsberg (97), coming to a similar conclusion that for the Juan Fernndez 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.
08

Noteworthy, in his thesis about the biogeography of Brassicaceae, Brggemann (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. Brggemann (000) mentions as example the absence of Lepidium from Juan Fernndez, 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.

Figure 4. Isla Alejandro Selkirk (Masafuera)

Figure 4.. Isla Robinson Crusoe (Masatierra) (from Skottsberg 90-9)

4.5.3 Archipilago de Juan Fernndez 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 / 8047W) (also known as Isla Ms Afuera), 80 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 Ms 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 Fernndez (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 90-9, Muoz Pizarro 99, Stuessy et al. 984b, Marticorena et al. 998b,
09

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 islands native flora at a competitive disadvantage (Dirnbck 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 Fernndez 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 Brggens Tierra de Juan Fernndez, an old submerged landmass west of today South America (Brggen 90, Skottsberg 9, p. 94; see figure 5.12). Skottsbergs 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
0

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 Fernndez 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).

17% 28%

11%

14% 4%

14%

1. Pantropical 2. Australasiatic 3. Neotropical 4. Antitropical 5. South-temperate 6. Endemic 7. Cosmopolitan

12%

Figure 4.4 Floristic elements Juan Fernndez

Furthermore, including Juan Fernndez (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 Fernndez
Regions ANT/COQ ANT/BIO ANT/MAG COQ/BIO COQ/MAG BIO/MAG JF/MAG JF/BIO JF/COQ JF/ANT SIMIL DIST 0,47 777 0,9 7 0, 7 0,4 70 0,7 40 0,4 700 0,18  0, 79 0, 89 0,08 49

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:



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 040 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 Fernndez Ridge, that controls the tectonic and geological evolution of the southern Andes at -4S since the Tertiary (Yez et al. 2001). The seafloor age assigned by Mller et al. (1997) to the seafloor offshore Chile between 8S and 40S, based on magnetic anomalies and relative plate reconstructions, ranges from 20 to 48 mya. Juan Fernndez 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 (Muoz Cristi 94, Caas 9, Illies 97). Brggen (90,
1.0 Jaccard similarity 0.6 0.8

COQ/BIO ANT/COQ

0.4

ANT/BIO

BIO/MAG COQ/MAG JF/MAG ANT/MAG

0.2

JF/BIO JF/COQ

0.0

JF/ANT

2000

4000

6000 Distance [km]

8000

10000

Figure 4. Floristic similarity between different regions in Chile: Antofagasta (ANT), Biobo (BIO), Coquimbo (COQ) and Magallanes (MAG). The Jaccard similarity index as related to geographical distance within Chile (squares), and between Juan Fernndez 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 Fernndez 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 Fernndez Land occurred in Late Tertiary, and that this land was not at all at the same location of todays Juan Fernndez 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 Fernndez islands (Juan Fernndez Land) could be located at the same longitude (80W), but at 48S 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 Wikstrm 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 PlioPleistocene (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 Figure 4. Possible position of the hypotheaized of long-distance dispersal Crawford et al. Tierra de Juan Fernndez, according to floristic 00, p. 89). similarity at the genus level (see figure 4.25).


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 Brggen (90) 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 98b). 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).

4

 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 

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 Earths surface


Plate tectonics is the current paradigm in geology (and related sciences like biogeography), after the general acceptance of Alfred Wegeners Kontinentalverschiebungs theory. The Earths 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 grabentype 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. Villagrn & Hinojosa 997, McLoughlin 00, Sanmartn & 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 landmassess 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.).
7

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


Major separation events (W Gondwana / E Gondwana) Age (mya) 800 Period Possible cause 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) Opening South Atlantic Ocean, due to the emplacement of Plume-related ParanaEtendecka continental flood basalts in Brazil and Namibia (7-7 Mya). Final break-up of Africa and S Am was completed only at 80 mya) 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). Source

Early to Late Jurassic

Scotese & McKerrow 990, Storey 99, Reeves & de Wit 000 Turner et al. 994, Jones 987, Barker et al. 99, Grunow et al 99, MacLoughlin 00 Lawver & Gahagan 998

Africa South Am separation

90

Early Cretaceous

West AntarcticaS Am

-8

Palaeogene/ Oligocene

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). 8

00 mya

80 mya

0 mya

4 mya

0 mya Figure . (continued)

0 mya

9

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).

Glossopteris fossil leafs (www.paleontology.unibonn.de/glossopteris.htm)

Middle Permian (c. 260 Ma)


Subtropical desert Mid-latitude desert Cool temperate Cold temperate Glossopteris palaeoflora

Figure . Distribution of the Permian Glossopteris flora in Gondwana (after McLoughlin 00). Palaeo-biomes after Willis & McElwain 00

Late Triassic (c. 200 mya)


Tropical summerwet Subtropical desert Winterwet Warm temperate Cool temperate Dicroidium palaeoflora

Dicroidium leafs (www.rosssea.info/geology. html) 0

Figure . Distribution of the Triassic Dicroidium flora in Gondwana (after McLoughlin 2001). Palaeobiomes after Willis & McElwain 00

Table 5.2 Gondwanan floras: five major phases of development before the appearance of angiosperms (sensu McLoughlin 2001)
Composition Free-sporing, herbaceous forms occupying moist habitat (Coastal areas, river flats, delta swamps, lake margins) = Rhynopsida, Zosterophyllopsida, Trimerophytes and lycopod fossils. Palaeoenvironmental conditions

Floras

. Cosmopolitan early land plant floras Herbaceous to shrub-sized lycophytes attributed to Haplostigma, Leclercqia, Archaeosigillaria and Protolepidodendron. Emergence of first arborescent plants: Lepidodendron and progymnosperms. Seed-fern floras progressively replaced lycophyte floras. Ginkgoales, conifers, Cycadales, Cordaites. Glossopteris is a group of extinct gymnosperms. Appears in the fossil record at about the end of the Carboniferous in W Gondwana.

Age (mya) ~4090

Period

Silurian Early Devonian

90

Middle Devonian

. Arborescent lycopod and seed-fern floras in a cooling climate

70

Late Devonian

0

Carboniferous

Fossils of Zosterophyllum and the lycopod Baragwanathia present in both hemispheres suggest little floristic provincialism (McLoughlin 00), but several authors recognize at least five biogeographical units (see Willis & McElwain 00) General cooling climate, with at least four periods of southern hemisphere glaciation. W Gondwana rotated into polar latitudes. Parts of central and norther S Am were affected by glaciation. E Gondwana remained in middle latitudes. Strong latitudinal gradient permitted the initial development of a distinctive S hemisphere flora (Meyen 1987) and the first expression of intra-Gondwanan floristic provincialism. Early Permian Gondwana locked in deep glaciation. Middle to Late Permian warmer climatic conditions favoured expansion of Glossopteris, that disappear from all continents along with ca. 80% of the worlds biota during the Permian-Triassic extinction.. The interior of Pangea was hot and dry. Warm Temperate climates extended to the Poles. Possibly one of the hottest times in Earth history. There was no ice at either North or South Poles. Warm temperate conditions extended towards the poles. Conditions generally warm and humid, local coal deposits, lack of glaciation. Parts of S Am and S Afr experienced arid conditions. The cosmopolitan spread of conifers and the biota in general is interpreted as a consequence of more equable global climate and a lack of physical barriers to plant migration (=range expansion).


More diverse floras dominated by Dicroidium (see also Herbst et al. 00) (Corystospermales), voltzialnean conifers, ginkgophytes, peltasperms, putative gnetales, bennettitaleans, pentoxylaleans and cycadophytes, plus many lycophytes and osmundaceaen, gleicheniacean, dicksoniaceaem dipteridacean and marattiacean ferns. Major radiation of the conifers. Conifer- and benettitalean- dominated communities replaced Dicroidiumdominated floras across the S hemisphere. Plant groups that have their origins in the Triassic reached more importance in the Jurassic: caytonialeans, bennettitaleans, pentoxylaleans and pachypterid seed-ferns. Also marattaicean, matoniacean, osmnundacean, discksoniaceae and dipteridacean ferns, equisetaleans, herbaceous lycophytes, and cycadaleans. Most genera persist into the earliest Cretaceous. Araucarian, podocarp, cheirolepidacean and taxodiacean conifers, caytonialeans, bennettitaleasn and ginkgoaleans dominated broad tracts of both N and S hemisphere.

. The glossopterid flora

00

Permian

4. The Dicroidium flora

00

Triassic

. Jurassic pteridospermconifer floras

004

Jurassic

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 & Villagrn (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 & Villagrn (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).

Late Cretaceous (c. 70 mya)


Warm temperate Tropical summerwet Subtropical desert Mid-latitude desert Cool temperate Tropical palaeoflora Nothofagus expansion

Early Eocene (c. 50 mya)


Warm temperate Tropical everwet Cool temperate Subtropical summerwet

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 .) 

Subtropical desert

Early Oligocene (c. 30 mya)


Tropical everwet Subtropical summerwet Warm / cool temperate Cool / cold temperate

D
Cold temperate / arctic Glacial Tropical palaeoflora Antarctic palaeoflora Mixed palaeoflora

Miocene (c. 10 mya)


Tropical everwet Subtropical summerwet Winterwet

Warm temperate Cool temperate Arctic Glacial

Pliocene (5 - 2 mya)
Tropical everwet Subtropical summerwet Desert (hyperarid) Desert (arid)

Subtropical summerwet Cool temperate Arctic Glacial Andean uplift

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

Table 5.3 Cretaceous to Neogene evolution of the floras of South America, mainly from Romero (1993), Hinojosa & Villagrn (1997), Troncoso & Romero (1998), Willis & McElwain (00), Hinojosa (00).
Composition Floras dominated by a diversity of conifer and pteridosperm groups. Gymnospermae (Brachyphyllum and Ptilophyllum), Gingkoales, Cycadopsida, Coniferopsida (Araucariaceae, Podocarpaceae, Cheirolepidaceae). Possibly angiosperm fossil pollen (Clavatipollenites and Liliacidites) from the Berriasian (limit Jurassic/ Cretaceous) Big change!: replacement of gymnosperms by the angiosperms as dominant. Lauracea, Sterculiaceae, Bignoniaceae, Monimiaceae (extant genera Laurelia, Peumus). Schinopsis = tropical forest and a more xerofitic community. Palaeoenvironmental conditions Mild icehouse world. Cool temperate forests covered the polar regions.

Palaeofloras

Age (mya)

Period

. Primitive angiosperms

4-00

Early Cretaceous

. Neotropical (without Nothofagus)

99-8

Late Cretaceous (Cenomanian to Coniacian)

8- Neotropical flora with marginal presence of Nothofagus. Campanian first appearance of Nothofagus in Antarctica. Maastrichtian first appearance of Nothofagus in the fossil record from Central Chile and Tierra del Fuego. In spite of its marginal presence, it is the peak of northern expansion of Nothofagus in S Am, reaching 0S. Tropical forests with palms and mangroves, Myrtaceae, Lauraceae, Proteaceae with marginal presence of Nothofagus. Azonal presence of gymnosperms (Cheirolepidaceae, Araucariaceae, Podocarpaceae, Zamianceae. High presence of Nothofagidites in Tierra del Fuego. A number of families recorded in South America have a pantropical to subtropical distribution: Anacardiaceae, Annonaceae, Araceae, Arecaceae, Combretacae, Myricaceae, Olacaceae, Rhizophoraceae, Rutaceae, Sapindacae, Zamiaceae, some of them which are best represented in S Am: Bombacaceae, Loranthaceae, Melastomataceae. Also some Old world families, like Pandanaceae, Casuarinaceae, Caprifoliaceae. Australasiatic genera (Eucryphia, Embothrium, Drimys), and subtropical Sapindaceae.

CampanianMaastrichtian

4

(with Nothofagus) sensu Troncoso & Romero 998)

Six global biomes are recognized at the end of the Cretaceous (Willis & McElwain 00), five of them to be found in S Am (figure 5.4 A) Warm and tropical conditions dominated in W Gondwana.

. Tropical Gondwanic (sensu Hinojosa 00)

-

Palaeocene

At the end of the Palaeocene, the planet heated up in one of the most rapid (in geologic terms) and extreme global warming events recorded in geologic history, called the PalaeoceneEocene Thermal Maximum.

Table 5.3 (continued) Cretaceous to Neogene evolution of the floras of South America, mainly from Romero (1993), Hinojosa & Villagrn (1997), Troncoso & Romero (998), Willis & McElwain (00), Hinojosa (00).

4. Subtropical Gondwanic (sensu Hinojosa 00)

-40

Eocene

New families like Araliaceae, Guttiferae, Meliaceae, Sterculiaceae, Euphorbiaceae, Bignoniaceae, Malpighiaceae, Brassicacae, Solanaceae, Asteraceae, Chenopodiaceae, and Ericaceae.

Dominance of a subtropical climate, more warm than today but cooler than at the Paleocene-Eocene. The Eocene global climate was perhaps the most homogeneous of the Cenozoic; polar regions were much warmer than today; temperate forests extended right to the poles (figure 5.4 B). General cooling. Separation of Australia from Antarctica c. 7 mya, origin of the circumantarctic ocean circulation and the E Antarctic glaciation. Global mid-Miocene warm climatic optimum. Well-defined climatic belts stretched from pole to equator (figure 5.4 D)

. Mixed (sensu Hinojosa 00) Mixture of the tropical elements with antarctic elements (figure 5.4 C). Subantarctic flora prevails till today in southernmost SAm and Tierra del Fuego. Reduction of former important Sapindaceae and Lauraceae, extinction of Moraceae, Annonaceae, Dilleniaceae, Malpighiaceae, Vochysiaceae, Tiliaceae, Sterculiaceae, Sapotaceae, and Styracaceae. Australantarctic cold element is replaced by more mesic element. Development of Mediterranean-type sclerophyllous vegetation in Central Chile. First record (Cerro Los Litres, 8) of extant sclerophyllous taxa from Central Chile: Beilschmiedia, Peumus, Myrceugenia, and Cryptocarya, together with some taxa that correspond today to the subtropical forests of the E Andes = Athyana and Choclospermum. Fragmentation of former continue subtropical forests. Origin of disjunctions between central Chile and NE Brazil.

40-

Oligocene



. Subtropical Neogene

-

Miocene

7. Subtropical Neogene (more xerophytic)

-

Pliocene

Increasing aridity in subtropical S Am, as a consequence of the separation of Antarctica from S Am, the glaciation of W Antarctica, the establishment of the Humboldt current, and the final uplift of the Andes and its increasing rain shadow effect (figure 5.4. E).

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 presentday 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 & Villagrn 1997, 2005; Troncoso & Romero 1998) (figure 5.4).

5.2 Oceanic transgressions over the continental surface


Riccardi (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


area was again downwarped. A marine back-arc basin extended in a NNW direction, from about 40S to the latitude of Copiap 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 CretaceousEarly 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 & Villagrn (997) considers this wide transgression having affected only E South America. Martnez-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).

30

36

42

48

54

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

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

7

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 Earths surface by different 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 palaeobotany (Ghosh et al. 00). Analysing palaeobotanical data, Gregory-Wodzicki (000) concluded 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 (404S) suggest an OligoceneMiocene 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 Villagrn & 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
8

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 subtropicaltropical 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 intraAndean 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 Chilo with Subantarctic Evergreen Forest was increasingly replaced by parkland under progressive cooling after 47,000 [4]C yr BP (Heusser

9

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 4,40 [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 Aisn, Magallanes, insular and continental Chilo, and the central and southern Andes (Villagrn & 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).
36

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, Villagrn et al. 004).

42

Campos de Hielo Norte and Sur are the last remnants of the Pleistocene glaciations, the biggest 48 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 54 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 Figure .7 Maximal extension of the last of plants, while the size of possible refuges would cycle of the Llanquihue glaciation (after be considerably enlarged if debris-covered glaciers Denton et al. 999, Heusser 00). 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
0

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 sciencesfaces 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)



dont like the alleged size or placement of a pre-Cenozoic ocean, just wait a while. It will change (McCarthy 00 b). Land bridges The idea of former land bridges in the site of todays 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 todays 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)



Figure . Land bridges as proposed by Leon Croizat (98)

Pacifica continent As discussed in section 4.5.3, the idea of a Pacifica continent West of South America is an old idea. During the 80s, 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 5.12 Former land areas in the present Pacific and Indian Oceans (from Dickins et al. 1992, 1996)

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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 earth The 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 (97) 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, 00a). 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, McCarthy 00). See also http://www.expandingearth.org/ 

Bonus track: modeling expanding earth, Paola pregnant on ..004 (top); same on .0.007 (bottom)



 Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography

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

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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 paradigm


The program of molecular dating of phylogenies has becoming very popular due to the availability of better resolved phylogenies in combination with new methods for estimating divergence times (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. Wikstrm 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];

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. 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. Wikstrm 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 (00b) 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, Headss 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
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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 Pulqurio & 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 Pulqurio & 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 (Pulqurio & 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 earlymid 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).

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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. Magalln & 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 (Wikstrm et al. 00). The most basal Chilean clades in the angiosperm tree are the following (proposed ages by Wikstrm 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 [49 Mya] Oligocene.
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Jurassic

Early Cretaceous

Late Cretaceous

Paleogene

Neogene

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160

140

120

100

80

60

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Malpighiales Oxalidales Celastrales Fabales Fagales Cucurbitales Rosales Zygophyllales


eurosids eurosids I

Myrtales Sapindales Malvales Brassicales


Tapiscia Aphloia Ixerba Crossomatales

eurosids II

Geraniales Vitaceae
eudicots

Saxifragales Berberidopsidales Dipsacales


euasterids II

Eremosyce

Escallonia Apiales Asterales


Aquifoliales

asterids

Lamiales
euasterids I

Solanales Gentianales
Oncotheca
Garryales

Ericales Cornales Caryphyllales


Dilleniaceae

Santalales Gunnerales
Trochodendraceae Buxaceae Sabiaceae

Proteales Ranunculales Ceratophyllum Monocots Magnoliales Winterales Laurales Piperales


Chlorantales Austrobaileyaceae Illiciaceae Schisandraceae Nymphaeaceae Amborellaceae
eumagnoliids

magnoliids

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100

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

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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 CretaceousPaleogene 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 (MidCretaceous) (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 (Wikstrm et al. 00) or at 0 mya (Anderson et al. 00), Proteales at 44-0 mya (Wikstrm 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 (Wikstrm 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, Wikstrm 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 Wikstrm 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).

44

6.3 Vicariance v/s dispersal in the Chilean Flora


As 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 (Muoz 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 longdistance 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).
Htten die Wandervgel die ihnen oft zugeschriebene Bedeutung fr Verbreitung der Pflanzen, so wrden die Zugstraen der Vgel als deren Fcalstraen sich floristisch ebenso darstellen, wie etwa die prhistorischen Handelstraen aus den Fundstcken 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.

4

Table . Evolutionary processes (dispersal v/s vicariance) proposed for several Chilean taxa
Families or groups Apocynaceae Apiaceae Genera or group Oreomyrrhis Evolutionary process Gondwanan origin for the family Origin in Eurasia and subsequently dispersal to North America and southern Pacific Rim. Gondwanan vicariance between Australasia-South America and LDD to Hawaii Gondwanan origin Colonization Aus from N Am Vicariance, widespread ancestor Dispersal between Australasia South America Vicariance Dispersal from NW Africa across the Atlantic Ocean for the origin of the South American taxa Initial diversification at 100140 mya, probably in West Gondwana, early entry into Antarctica, and long-distance dispersal to New Zealand and New Caledonia South America-Old World disjunct distribution due to Cretaceous vicariance three independent dispersals from North to South American dispersal from South America to Australasia, or vice versa LDD via migrating birds explains the amphitropical disjunction between South American R. philippiana and North American R. curvisiliqua. Probably long-distance dispersal from western North America to South America by birds in the Pleistocene Gondwanan ancestry LDD Autralasia to South America Laurasian in origin.with following dispersals Dispersals between California and Central Chile Origin in North hemisphere and dispersals to the South and the Pacific Dispersal from South America into Aus+NZ less than ,7 mya Gondwanic origin, Coronantheroid Gesneriaceae (Asteranthera, Mitraria, Sarmienta) relicts Early vicariance followed by recent dispersals Late Cretaceous vicariance Sources Potgieter & Albert 00 Chung et al. 00

Araliaceae

Pseudopanax

Mitchell & Wagstaff 000

Araucariaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae

Microseris Microseris Abrotanella Abrotanella Hypochaeris

Setoguchi et al. 998 Vijverberg et al. 999 Grehan 007 Swenson & Bremer 997, Wagstaff et al. 00 Heads 999 Tremetsberger et al. 00

Atherospermataceae

Laurelia

Renner et al. 000

Berberidaceae Boraginaceae Brassicaceae Brassicaceae

Berberis Tiquilia Cardamine Rorippa

Kim et al. 004 Moore et al. 00 Bleeker et al. 00a Bleeker et al. 00b

Brassicaceae Cunoniaceae Cyperaceae Ericaceae Fabaceae Fabaceae Gentianaceae Gesneriaceae

Lepidium Oreobolus Hoffmannseggia Sophora Gentianella Coronantheroid

Mummenhoff et al. 00 Bradford & Barnes 00 Chacn et al. 00 Kron & Luteyn 00 Simpson et al. 00 Hurr et al. 999, Pea et al. 000 Von Hagen & Kadereit 00 Burtt 998

Gunneraceae Gunneraceae

Gunnera Gunnera

Wanntrop & Wanntrop 00 Fuller & Hickey 00

47

Table . (continuation)


Lauraceae Lycopodiaceae Monimiaceae Beilschmiedia/ Cryptocarya Peumus Gondwanan vicariance (diverged from ancestor about 90+- 0 mya. Vicariance, Permian origin of the genera Vicariance at 7 mya, former continuous range from Chile across Antarctica and the Kerguelen plateau to Madagascar Gondwanan vicariance Gondwanan vicariance vicariant main massings of the four subgenera compatible with allopatric differentiation and no substantial dispersal Nothofagaceae Plantaginaceae Plantaginaceae Nothofagus Hebe Ourisia Vicariance Am-Aus subgen Fuscospora, dispersal in Lophozonia at least two instances of transoceanic LDD from Australasia to South America. origin in the central Chilean Andes followed by consecutive dispersal to the southern Andes and dispersal(s) to the northern Andes and Australasia Gondwanan vicariance Dispersal California to South America American origin and dispersals into Australia Primula Caltha Gondwanan; Mid-Cretaceous divergence between major groups Ancestral haplotype gave origin to S Am, Euro and N Am lineages. 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. Gondwanic origin followed by dispersals Probably Gondwanan relict Vicariance Africa as ancestral area followed by longdistance dispersal into the Pacific LDD New Zealand South America migration from tropical Gondwana to Antarctica and Australia throughout South America dispersals between South America and southern Africa Chanderbali et al. 00 Wikstrm & Kenrick 00 Renner 004

Monocots (major groups) Nothofagaceae

Nothofagus

Bremer & Janssen 00 Swenson et. al 00, Heads 00

Knapp et al. 00, Cook & Crisp 00 Godley 97, Wagstaff & Garnock-Jones 998 Meudt & Simpson 00

Poales Polemoniaceae Portulacaceae Proteaceae Primulaceae Ranunculaceae

Gilia

Bremer 00 Morrell et al. 000 Applequist & Wallace 00 Hoot & Douglas 998 Guggisberg et al. 00 Schuettpelz & Hoot 004

Restionaceae Rhamnaceae Rubiaceae Rubiaceae Tetrachondraceae Winteraceae

Apodasmia Discaria Coprosma Coprosma Tetrachondra -

Linder et al. 00 Richardson et al. 000 Heads 99 Anderson et al. 00 Wagstaff et al. 000 Barreda & Archangelsky 00 Beier et al. 004

Zygophyllaceae

Fagonia

48

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. Sanmartn & Ronquist 004). Recently Sanmartn et al. (007) tested the directional dispersal in the Southern Hemisphere using eventbased 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 ZealandSouth 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 (Sanmartn 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 hiptesis de relaciones entre reas formuladas por la geologa no tienen validez intrnseca superior a las que se formulen a partir de datos biolgicos, es decir, de los cladogramas de reas. A partir de este principio, de por s totalmente aceptable, algunos autores concluyen que las hiptesis geolgicas no pueden utilizarse ni para corroborar ni para refutar las hiptesis biogeogrficas, en contradiccin patente con uno de los principios bsicos del cladovicariancismo: la idea del paralelismo entre la evolucin 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?
49

In the words of Heads (00b): 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. Thats 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 00b). 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
0

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 navet. However, I suspect that in the final analysis it will be the geologist who is proved nave 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. Its easy to see why it causes alarm, because it obviously cant be applied to fossils. Its 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 Earths species distributed?, and How have species distributions changed over time? Heads find equally myopic speciescentrism in Hubbells (00) Unified Neutral Theory of Biodiversity and Biogeography (Heads 00a, 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 4S. 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 groups 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 Simpsons 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 Fernndez as one of the best scenarios for


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 speciesrichness 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. Dickermans (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. An 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

4

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 Fernndez, 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 earths surface, much in the sense of Willis 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, Sssenguth, Pachosky, Rosa, Schrter, 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 NatureYet 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 (90) 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 Nairs 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


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 00b). 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 4040 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).

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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).
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6.6.2 Angiosperms origin(s) The origin of flowering plants is considered by some to be the Holy Grail (Miller 2006), or Darwins 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 090 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 8040 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. Wikstrm et al. (00) recognize the calibrated phylogeny here is a working hypothesis and should be viewed as such. In Bremers opinion (00): The analysis by Wikstrm 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
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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 Crepets 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).
0

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 transitionalcombinational 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

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

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 (98) and Berry (90) 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 99). Others like Golenkin (97, 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

suggested an origin of the angiosperms during the Permian as the aftermath of the so called Permo-Carboniferous Glaciations (Croizat 90, 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 cant paleobotanists be more honest and say that they dont 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.



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7 Back to Postmodern Biogeography



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

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 postEnlightenment 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. W.D. Matthew 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. ). Matthews 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 Simpsons book Evolution and Geography (box 7.). A discussion on Matthews biogeographic program has been done by Nelson & Ladiges (00) (see also an early review by Barbour 9). A critique is to
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Box 7. George Gaylord Simpson

G.G. 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
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Figure 7.1 Griffith Taylors map: Zones of migration showing the evolution of the races (1921).

varieties of the human race occupied the tropical zones That Taylors 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 cooltemperate climate [where] they reach their highest physical, mental and social attainments (Matthew 9, p. 4). Livingstone continues noting the reprint of Taylors map in Huntingtons 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 Figure 7. Matthews map: migration and the distribution of the human races (9). should ever rise very high
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in the scale of civilisation (Huntington 94, 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 Livingstones opinion: the cartography accompanied a discourse of climates 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 Mller (99). Lets 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 Haeckels courses at Jena during 89. Haeckel is known as one of the most convinced defender of Darwins ideas (Livingstone 99, p. 98). Ratzels work was also greatly influenced by the naturalist, explorer and evolutionary theorist Moritz Wagner. He was an early devotee of Darwins theory, but he believed that Darwin had failed to appreciate the significance of migration and geographical isolation in the processes of speciation (Wagner 88). 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 Ratzels new anthropogeographie Ratzels Anthtropogeographie can best be read as an attempt to situate the new science of human geography within the naturalistic framework of Wagners 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 Mller 99 and his claims against Stoddart 98). Lets 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 biogeography8:
Aber wenn man nun die wirkliche Verbreitung der Lebewesen ansieht, kann man doch allen diesen Hilfsmitteln der passiven Wanderung nicht die groe Wirksamkeit zusprechen, die seit Darwin und Wallace so viele ihnen zugeschrieben haben

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Da sich Lnder und Meere auf der Erde ununterbrochen verschoben haben und noch heute unter unseren Augen sich verschieben, so verndert sich also bestndig der Lebensraum fr die wasserlebenden und ebenso die landlebenden Organismen9. 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, da man weder nach Punkten (Ursprung) noch nach Linien (Wegen), sondern nach Rumen oder Gebieten zu fragen hat.

Ratzels (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 Friedrich Ratzel 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). Dawkins is graphically described as an ultradarwinist 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

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 largescale 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 Gieryns (999) Cultural Boundaries of Science, or Kohlers (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 Clements famous field ecology quadrats might be the reflection of Nebraskas 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 believecultural 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 Cornets 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 angiospermlike 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 (99) 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).
7

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 Maihuenioideae Opuntioideae Cactoideae Trichocereeae Notocacteae Browningieae Pachycereeae Tribe Genera Maihuenia Cumulopuntia, Maihueniopsis, Miqueliopuntia, Pterocactus, Tunilla Echinopsis, Haageocereus,Oreocereus Austrocactus, Copiapoa, Eriosyce, Eulychnia, Neowerdermannia Browningia Corryocactus

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 (99) 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 cant 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 99). 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

74

Box 7. (continuation)

Notocacteae
Neowerdermannia Copiapoa Eriosyce Eulychnia Austrocactus

Browningieae

$ !

Maihuenioideae
$ Maihuenia

! Browningia

Pachycereae
$ Corryocactus

$ $ $

Trichocereeae
Echinopsis Haageocereus Oreocereus

Opuntioideae
Tunilla Cumulopuntia # Miqueliopuntia Maihueniopsis Pterocactus # # #

7

Box 7. (continuation) 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.
!
" $ $ !

18

Opuntioideae
! Cumulopuntia " Maihueniopsis # Miqueliopuntia $ Tunilla $ Pterocactus ! !
"

! $ "!
" "

"

Notocacteae
24

$ $ # # #

18

$ ! $! #

18

Trichocereeae
24 !

#
! !

! !
24

#" " # #! "


! !

! Copiapoa $ Eriosyce # Eulychnia $ Neowerdermannia ! Austrocactus

# # # # # #
!

Echinopsis $ Haageocereus # Oreocereus

30

30

! ! ! ! ! ! ! ! !

30

36

!
$

36

36

42

42

42

$
48

!
48
48

54

54

54

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.

7

Box 7. (continuation) 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.

Other Opuntioideae + Pereskioideae + Cactoideae

basal Opuntioideae

Maihuenia

Biogeographical scenario for the evolution of Cactaceae (compare with Fig.  in Edwards et al. 00)

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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 waywhether Postmodernism is nonsense or truththe 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 multifactorial 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 Hengevelds 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.
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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 evodevo, 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, Garca 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. Its a dance through morphospace, the space of the forms of organisms (Goodwin in Brockman 99, p. 97)0

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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 Horaces 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 00a, 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. Lets take Reason & Goodwins positive view, together with Gyerin pragmatic one: may the best science win (Gieryn 1999, title of chapter ).

Figure 7. Valparaso (photo courtesy M. Richter)

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 dont 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 ones beliefs and ones scientific evidence may be one of the most difficult aspects of evolutionary studies (Grande 1994, p. 79).

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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, southtemperate, 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.

8

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 Fernndez 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 hemispheres 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 Fernndez 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.

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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 (Fernndez-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 Raups 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 Wettsteins work can be seen as the precursor of the modern concept of allopatric speciation.


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.)
4

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)


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).
7

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

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.
9

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/~geo7/ crap.html]
0

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 Muoz Pizarro (9) considered also many naturalized taxa.



8

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


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
7 8

Croizat apparently took the term from Van Steenis (Croizat 9) Due to human pressure, Santalum fernandezianum is all but extinct from Juan Fernndez (Islands).

9

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).
0 

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

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


4

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

7

It is a pity that Mller (99) in his excellent revision of Ratzels program does not explicit analyze this great piece: Der Lebensraum: eine biogeographische Studie (Ratzel 90).
8 9

Note that this was written  years before the proposal of continental drift by Alfred Wegener. Also available at http://www.edge.org/rd_culture/bios/goodwin.html

0

87

References
Aagesen 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 Ramrez (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 92,77-748. 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. An C & MJ Sanderson (00) Missing the forest for the trees: phylogenetic compression and its implications for inferring complex evolutionary histories. Systematic Biology, 54, 47. 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, 994. Applequist WL & RS Wallace (00) Phylogeny of the portulacaceous cohort based on ndhF sequence data. Systematic Botany, 26, 4049. Arldt T (98) Die Entwicklung der Kontinente und ihrer Lebewelt (d. ed.) Berlin. Arroyo MTK, C Castor, C Marticorena, M Muoz Schick, L Cavieres, O Matthei, FA Squeo, M Grosjean & R Rodrguez (1998). The flora of Llullaillaco National Park located in the transitional winter-summer rainfall area of the northern Chilean Andes. Gayana Botnica 55, 9-0. Arroyo MTK, O Matthei, C Marticorena, M Muoz Schick, F Perez & AM Humaa (2000) The vascular plant flora of the Bellotos del Melado National Reserve, VII Region. Chile: a documented checklist. Gayana Botnica 57, 7-9. Arroyo MTK, C Marticorena, O Matthei, M Muoz Schick & P Pliscoff (2002) Analysis of the contribution and efficiency of the Santuario de la Naturaleza Yerba Loca, 33 S in protecting the regional vascular plant flora (Metropolitan and Fifth regions of Chile). Revista Chilena de Historia Natural, 75, 77-79. Arroyo MTK, O Matthei, M Muoz-Schick, J Armesto, P Pliscoff, F Perez & C Marticorena (00) Flora de cuatro reservas nacionales en la Cordillera de la Costa, - S, VII Regin, Chile y su papel en la proteccin de la biodiversidad regional. Biodiversidad y Ecologa 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, 977.

88

Axelrod DI (9) How old are the angiosperms? American Journal of Science, 259, 447-49. Backeberg C (94) Cactaceae. Jahrbcher der DKG, Teil , Juni 94: , Cactaceae Lindley; Systematische bersicht (Neubearbeitung) mit Beschreibungsschlssel. Bader FJW (90) Die Verbreitung borealer und subantarktischer Holzgewchse in den Gebirgen des Tropengrtels. Nova Acta Leopoldina, 23, -44. Barbour M (99) Patterns of genetic similarity between Larrea divaricata of North and South America. American Midland Naturalist, 81, 47. Barbour T (9) Some Remarks upon Matthews Climate and Evolution. Annals of the New York Academy of Sciences, 27, -0. [http://www.wku.edu/~smithch/biogeog/ BARB9.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 kologie sdamerikanischer Hochgebirgspflanzen. Physische Geographie (Zrich), 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, 48. Beloussov VV (98) 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, 47. 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, -.

89

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 (00a) Phylogeny and biogeography of Southern Hemisphere high-mountain Cardamine species (Brassicaceae). Australian Systematic Botany, 15, 78. Bleeker W, C Weber-Sparenberg & H Hurka (00b) 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, 48. 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, 7097. 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, 988-990. 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, 887. Britton NL & JN Rose (999) 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 Mller (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, 879. Brown RW (9) Palm-like plants from the Dolores Formatioon (Triassic), southwestern Colorado. U.S. Geol. Surv. Prof. Pap. 274, 0-09. Brggemann H (000) Molekulare Systematik und Biogeographie der Gatttung Lepidium L. und Verwandter Sippen (Brassicaceae). Dissertation im Fachbereich Biologie/Chemie der Universitt Osnabrck. Brggen J (90) Fundamentos de la Geologa de Chile. Inst. Geogr. Militar, Santiago, Chile.

90

Bunge W (9) Theoretical Geography. Lund Studies in Geography, Gleerup, Lund. Burckhardt C (1902) Traces gologiques dun 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. 4-48, Oxford University Press. Buxbaum F (99) 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 fitogeogrfico de la Repblica Argentina. Revista del Museo Eva Pern, Botnica 8, 878. Cabrera AL (9) The genus Lagenophora (Compositae). Blumea, 14, 8-08. Cabrera AL & A Willink (97) Biogeografa de Amrica Latina. Monografa . Serie de Biologa. Secretara General de la Organizacin 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. Caas J (9) Tectnica y sedimentacin de la Formacin 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) Sonderbnde des Naturwissenschaftlichen Vereins in Hamburg, 7, 747, Verlag P. Parey, Hamburg/Berlin. Castro SA, A Camousseight, M Muoz-Schick & FM Jaksic (00) Rodulfo Amando Philippi, el naturalista de mayor aporte al conocimiento taxonmico de la diversidad biolgica 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-. Chacn J, S Madrin, 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, 044. 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) ber die Verbreitung der Pflanzen der alpinen Region der europischen Alpenkette. Denkschr. Schweiz. Nat. Ges. vol. 22.

9

Christaller W (98) 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, 04-07. Collier A (994) Critical Realism: An Introduction to Roy Bhaskars 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, 7-74. Cowling RM, PW Rundel, BB Lamont, MTK Arroyo & M Arianoutsou (99) Plant diversity in Mediterraneanclimate 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, 779. Crane PR (99) The fossil history of the gnetales. International Journal of Plant Sciences, 157, (Suppl. S), S0-S7. 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 Fernndez Archipelago, Chile. Plant Species Biology, 16, 89.

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, 8700. Crepet WL (998) The Abominable Mystery. Science, 282, -4 Crepet WL (000) Progress in understanding angiosperm history, success, and relationships: Darwins abominably perplexing phenomenon. PNAS, 97, 99-94. 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-Juent (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 (98) 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 (90) 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) Biogeografa analtica y sinttica (Panbiogeografa) de las Amricas. Biblioteca de la Academia de Ciencias Fsicas, Matemticas 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 conservacin, restauracin y desarrollo de las islas Juan Fernndez, Chile. Revista Chilena de Historia Natural, 74, 899-90. 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 les de Robinson: Trsor vivant des mers du Sud, entre lgende et ralit. Nathan, Paris. Danton Ph & Ch Perrier (2003) Liste de la flore vasculaire de lle Robinson Crusoe, archipel Juan Fernndez, Chili. Le Journal de Botanique de la Socit Botanique de France, 24, 7-78. Darwin Ch (89) The Voyage of the Beagle. John Murray, London. Darwin Ch (89) 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) Darwins abominable mystery: Insights from a supertree of the angiosperms. PNAS, 101, 904-909.

9

Davis CC, WR Anderson & MJ Donoghue (00) Phylogeny of Malpighiaceae: evidence from chloroplast ndhF and trnL-F nucleotide sequences. American Journal of Botany, 88, 8084. 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. 98. Las tribus y generos de Escrofulariaceas representados en Austro-Amrica y su distribucin geogrfica. Rev. Mus. La Plata (Bot.), 59, 0-8. de Candolle, Alphonse (8) Introduction ltude de la Botanique I, II. Librairie Encyclopdique de Roret, Paris. de Candolle, Alphonse (8) Gographie botanique raisonne, . vols. Paris. de Candolle, Augustin-Pyramus (80) 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, 87. 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 Schlchter & 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 (98) Esquisse cologique du Chili. Biologie del Amrique 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) Bioclimatologa de Chile. Vicerrectora Acadmica, Universidad Catlica de Chile, Santiago. Dickerman AW (998) Generalizing phylogenetic parsimony from the tree to the forest. Systematic Biology, 47, 444 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 Gschen, Gschensche Verlagshandlung, Leipzig. Dillon MO, M Nakazawa & S Leiva Gonzlez (00) The Lomas Formations of Coastal Peru: Composition and Biogeographic History. El Nio 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. Dirnbck 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, 09.

94

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. Domnguez MC, S Roig-Juent, 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, 77. 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 gegenwrtigen Verbreitungsverhltnisse der Pflanzen: Ein Beitrag zur vergleichenden Erdkunde. Ergnzungsheft 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 Handbcher (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 Gonzlez Lpez & J Juez-Larr (00) OligoceneMiocene 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, 47444. Ebach MC & CJ Humphries (00) Ontology of biogeography. Journal of Biogeography, 30, 999. Ebach MC & DF Goujet (2006) The first biogeographical map. Journal of Biogeography, 33, 779. 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, 89. 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, 47. 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 C8 O Bonds in Paleosol Carbonates. Science, 314, 70c. 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, 40. Engels E-M (00) Ethik in den Biowissenschaften. Ethisch-Philosophisches Grundlagenstudium 2. Ein Projektbuch. (ed by M Maring), pp. -, Mnster, LIT Verlag. Engler A (879) Versuch einer Entwicklungsgeschichte der Pflanzenwelt, insbesondere der Florengebiete seit der Tertirperiode. Vol 1. Die Extratropischen Gebiete der Nrdlichen Hemisphre. Verlag von W. Engelmann, Leipzig. Engler A (88) Versuch einer Entwicklungsgeschichte der Pflanzenwelt, insbesondere der Florengebiete seit der Tertirperiode. Vol. 2. Die Extratropischen Gebiete der Sdlichen Hemisphre 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 Beitrge zum Gedchtnis der hundertjhrigen Wiederkehr des Antritts von Alexander von Humboldts Reise nach Amerika. (ed. by Gesellschaft fr Erdkunde zu Berlin), pp. -47. Evenstar L, A Hartley, C Rice, F Stuart, A Mather & G Chong (00) Miocene-Pliocene climate change in the PeruChile 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, highlatitude floodplain, Alexander Island, Antarctica. Journal of the Geological Society, 158: 709-74. 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, 47-4. Fernndez-Armesto F (997) Truth -a history-. Bantam Press, London. Ferr 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 Grninger, 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, 474. Frey W & R Lsch (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.

9

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) Resea botnica 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, 044-07. Gajardo R (994) La vegetacin natural de Chile. Clasificacin y distribucin geogrfica. 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. 474. Royal Botanic Gardens, Kew. Garca Azkonobieta T (2005) Evolucin, desarrollo y (auto)organizacin. Un estudio sobre los principios filosficos de la evo-devo. Tesis doctoral, Universidad del Pas 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 Bltenpflanzen im zentralen Nordbayern.  vols. IHW-Verlag Eching. Gauckler K (90) Das sdlich-kontinentale Element in der Flora von Bayern mit besonderer Bercksichtigung des Frnkischen Stufenlandes. Abhandl. Naturhist. Ges. Nrnberg, 24, -0. Gay C (8) La Isla de Juan Fernndez. Reprint 974, Museo Nacional de Historia Natural (Santiago) Serie Educativa, 9, -. Gay C (8484) Historia Fsica y Poltica de Chile segn documentos adquiridos en esta Repblica durante doce aos 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, 4-44. 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 (97) 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.

97

Goodwin B (994) How the Leopard Changed Its Spots: The Evolution of Complexity. Charles Scribners 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, 090. Grehan JR (00) Panbiogeography from tracks to ocean basins: evolving perspectives. Journal of Biogeography, 28, 4-49. 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, 49499. Griffith MP (2004b) Early cactus evolution: the postmodern view. Haseltonia, 10, -. Griffiths G (1974) On the Foundations of Biological Systematics. Acta Biotheoretica, 34, 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, 79794. 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 Bltenpflanzen in Mitteleuropa. Hoppea, Denkschr. Regensb. Bot Ges., 66, -9. Haeupler H & P Schnfelder (eds.) (988) Atlas der Farn- und Bltenpflanzen 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, 9-40. 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.

98

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, 4-44. 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, 840. 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 (00a) The history and philosophy of panbiogeography. Regionalizacin Biogeogrfica en Iberoamrica y Tpicos Afines (ed. by J Llorente & JJ Morrone), pp. 7-, Universidad Nacional Autnoma de Mxico. Heads M (00b) 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, 007. 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 (90) Grundzge 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. Henrquez JM, E Pisano & C Marticorena (1995) Catlogo de la flora vascular de Magallanes (XII Regin), 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 Puqun formations (Chile). Ameghiniana, 37, 8-9. Herbst R, A Troncoso & J Muoz (2005) Las Tafofloras Trisicas de la Regin de Los Lagos, Xma Regin, 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.

99

Hershkovitz MA (2006) Ribosomal and chloroplast DNA evidence for diversification of western American Portulacaceae in the Andean region. Gayana Botnica, , -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 Universitt Regensburg, Band 7 - Evolutionstheorie und ihre Evolution, Regensburg. Hettner A (97) 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 (~40S). 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 climticos y vegetacionales inferidos a partir de paleofloras Cenozoicas del sur de Sudamrica. Revista Geolgica de Chile, 32, 9-. Hinojosa LF & C Villagrn (997) Historia de los bosques del sur de Sudamrica, I: antecedentes paleobotnicos, geolgicos y climticos del Terciario del cono sur de Amrica. Revista Chilena de Historia Natural, 70, -9. Hinojosa LF & C Villagrn (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 Villagrn (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 Flix (Archipilago de las Desventuradas, Chile). Gayana Botnica, 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 students 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, 478. Hooker JD (844-80) The botany of the Antarctic voyage of H. M. Discovery ships Erebus and Terror, in the years 18391843. I Flora Antarctica, II Flora Nova-Zelandiae, III Flora Tasmaniae, L. Reeve, London.

00

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, 4-44. 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. 49-448, 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 Sudamrica. 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, 04-09. 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. Hultn E (90) Atlas of the Distribution of Vascular Plants in NW-Europe. Stockholm. Hultn E (94) The Circumpolar Plants I Vascular Cryptogams, Conifers, Monocotyledons. Stockholm. Hultn E (97) The Circumpolar Plants II Dicotyledons. Stockholm. Humboldt A von (80) Essai sur la gographie des plantes, accompagn dun tableau physique des rgions quinoxiales. 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 Scribners 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 sdlichen Chile. Geologische Rundschau, 57, 8--0. Irmscher E (1922) Pflanzenverbreitung und Entwicklung der Kontinente. I Teil. Studien zur genetische Pflanzengeographie. Mitteilungen aus dem Institut fr allgemeine Botanik in Hamburg, 5, 7-. Irmscher E (1929) Pflanzenverbreitung und Entwicklung der Kontinente. II Teil. Weitere Beitrge zur genetische Pflanzengeographie unter besonderer Bercksichtigung der Laubmoose. Mitteilungen aus dem Institut fr 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 Chaaral 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 Flix 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 Fernndez. Santiago, Chile. Jokat W, T Boebel, M Knog & U Meyer (00) Timing and geometry of early Gondwana breakup. Journal of Geophysical Research, 08, N B9, 48. Jones EJW (987) Fracture zones in the equatorial Atlantic and the breakup of western Pangea. Geology, 15,  . Jrgensen PM & S Len-Yez (eds.) (999) Catalogue of the vascular plants of Ecuador. Monographs in Systematic Botany, Missouri Botanical Garden, 75, iviii, 8.

Kamp PJJ (1980) Pacifica and New Zealand: proposed eastern elements in Gondwanalands 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 Applies to 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 Stckler, 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 LabField Border in Biology. University of Chicago Press, Chicago.

0

Kppen W (884) Die Wrmezonen der Erde, nach der Dauer der heissen, gemssigten and kaltern Zeit and nach der Wirkung der Wrme auf die organische Welt betrachtet. Meteorologische Zeitschrift, 1, -. Kppen W (1930) Die Klimagebiete nach Kppens Klassifikation. Klimakunde von Sdamerika (ed. by K Knoch), vol. , part G, pp. 4-9. Handbuch der Klimatologie (ed. by W Kppen & 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 Verhltnisse und Pfalzenverbreitung. Ark. Bot., 4, 4-49. 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, 90999. 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, 779. Lindemann B (97) Kettengebirge, kontinentale Zerspaltung und Erdexpansion. Jena. Linder CR & LH Rieseberg (004) Reconstructing patterns of reticulate evolution in plants. American Journal of Botany, 91, 700708. Linder HP (00) On areas of endemism, with an example from the African Restionaceae. Systematic Biology, 50, 899. 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, 8870.

0

Livingstone DN (99) The Geographical Tradition: Episodes in the History of a Contested Enterprise. Blackwell, Oxford & Cambridge. Livingstone DN (994) Climates 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. Lpez RP (00) Phytogeographical relations of the Andean dry valleys of Bolivia. Journal of Biogeography, 30, 98. Lu A-M & Y-C Tang (00) Consideration on some viewpoints in researches of the origin of angiosperms. Acta Phytotaxonomica Sinica, 43, 40-40. Luebert F & R Gajardo (000) Vegetacin de los Andes ridos del norte de Chile. Lazaroa 21, -0. Luebert F & R Gajardo (00) Vegetacin alto-andina de Parinacota (norte de Chile) y una sinopsis de la vegetacin de la Puna meridional. Phytocoenologia 35, 79-8. Luebert F & P Pliscoff (00) Sinopsis bioclimtica y vegetacional de Chile. Editorial Universitaria, Santiago. Luebert F, M Muoz-Schick & A Moreira-Muoz (00) Vegetacin y Flora de La Campana. Parque Nacional La Campana: Origen de una Reserva de la Biosfera en Chile Central. (ed. by S Elrtegui & A Moreira-Muoz), pp. -9, Taller La Era, Santiago.

Mabberley DJ (997) The Plant-Book (nd ed.). Cambridge University Press, Cambridge MacArthur RH & EO Wilson (97) 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. Magalln S & MJ Sanderson (00) Relationships among seed plants inferred from highly conserved genes: Sorting conflicting phylogenetic signals among ancient lineages. American Journal of Botany, 89,9900. Maldonado A, JL Betancourt, C Latorre & C Villagrn (00) Pollen analyses from a 0 000-yr rodent midden series in the southern Atacama Desert ( degrees 0S). Journal of Quaternary Science, 20, 49-07. Mann G (980) Ernst Haeckel und der Darwinismus: Popularisierung, Propaganda, und Ideologisierung. Medizinhistorisches Journal, 15, 9-8. Marquet PA, M Fernndez, 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) Contribucin a la estadstica de la flora vascular de Chile. Gayana Botnica, 47, 8-. Marticorena C (99) Bibliografa Botnica Taxonmica de Chile. Monographs in Systematic Botany vol. 4, Missouri Botanical Garden. Marticorena C (99) Historia de la exploracin botnica en Chile. Flora de Chile, vol. 1 (ed. by C Marticorena & R Rodrguez), pp. -, Universidad de Concepcin.

04

Marticorena C (997) La presencia de Maireana brevifolia (R.Br.) Paul G. Wilson (Chenopodiaceae), en Chile continental. Gayana Botnica, 54, 9-94. Marticorena C & R Rodrguez (eds.) (99-00) Flora de Chile. 2 vol. Universidad de Concepcin, Concepcin. Marticorena C, O Matthei, R Rodrguez, MTK Arroyo, M Muoz-Schick, FA Squeo & G Arancio (998a) Catlogo de la Flora Vascular de la Segunda Regin (Regin de Antofagasta), Chile. Gayana Botnica, 55, -8. Marticorena C, TF Stuessy & CM Baeza (1998b) Catalogue of the vascular flora of the Robinson Crusoe or Juan Fernndez Islands, Chile. Gayana Botnica, 55, 87-. Marticorena C, FA Squeo, G Arancio & M Muoz-Schick (2001) Catlogo de la flora vascular de la IV Regin de Coquimbo. Libro Rojo de la Flora Nativa y de los Sitios Prioritarios para su Conservacin: Regin de Coquimbo (ed. by FA Squeo, G Arancio & JR Gutirrez), 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 - . Martnez-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 (99) 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 (94) bersicht ber die Florenreiche und Florengebiete der Erde. A. Englers Syllabus der Pflanzenfamilien. 12 ed. vol. 2. Angiospermen. (ed. by H. Melchior), pp. -9, Gerbrder 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, 79740. 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, 700.

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. Einfhrung in die Lehre von der Verbreitung der Gewchse mit besonderer Bercksichtigung der mitteleuropischen Flora.  vols. Verlag Borntraeger, Berlin-Zehlendorf. Meusel H & E Jger (eds.) (99) Vergleichende Chorologie der Zentraleuropischen Flora, vol. 3, Jena. Meusel H, E Jger & E Weinert (eds.) (9) Vergleichende Chorologie der Zentraleuropischen Flora, vol. 1, Jena. Meusel H, E Jger, S Rauschert & E Weinert (eds.) (978) Vergleichende Chorologie der Zentraleuropischen 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) Evolucin de la serie Microphyllae (Adesmia, Fabaceae) en la Cordillera de los Andes: una perspectiva biogeogrfica. Revista Chilena de Historia Natural, 79, 89-404. Milbradt J (97) Nordische Einstrahlungen in der Flora und Vegetation von Nordbayern, dargestellt an ausgewhlten Beispielen. Hoppea, Denkschr. Regensb. Bot. Ges., 35, -0. Miller H (1970) Das Problem des hypothetischen Pazifischen Kontinentes gesehen von der chilenischen Pazifikkste. International Journal of Earth Sciences, 59, 97-98. 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, 409-44. 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 (80) Saggio sulla storia naturale del Chile. (d revised edition), Bologna.

0

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 80s. 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 Rser (eds.) (998) Eduard Friedrich Poeppig 1798-1868. Gelehrter und Naturforscher in Sdamerika anlsslich seines 200. Geburtstages. Universitt Leipzig. Morley RJ (00) Why are there so many primitive angiosperms in the rain forests of Asia-Australasia? Faunal and floral migrations and evolution in SE Asia-Australasia (ed. by I Metcalfe, JMB Smith, M Morwood & I Davidson), pp. 899. 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, 48-44. Morrone JJ (00) Biogeografa de Amrica Latina y el Caribe. M&TManuales & Tesis SEA, vol. . Zaragoza, 48 pp. Morrone JJ (00) Biogeographical regions under track and cladistic scrutiny. Journal of Biogeography, 29, 49-. Morrone JJ (2003) Quin 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, 884. 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, 47494. Morton AG (98) History of Botanical Science. Academic Press, London. Mller GH (99) Friedrich Ratzel (1844-1904): Naturwissenschaftler, Geograph, Gelehrter: Neue Studien zu Leben und Werk und sein Konzept der Allgemeinen Biogeographie. Verlag fr Geschichte der Naturwissenschaften und der Technik, Stuttgart. Mller RD, WR Roest, J-Y Royer, LM Gahagan & JG Sclater (1997) Digital isochrones of the worlds 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, 00. Muoz J, AM Felicsimo, F Cabezas, AR Burgaz & I Martnez (004) Wind as a long-distance dispersal vehicle in the Southern Hemisphere. Science, 304, 4447. Muoz Cristi J (94) Rasgos generales de la constitucin geolgica 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. Muoz Pizarro C (944) El itinerario de don Claudio Gay. Boletn Museo Nacional de Historia Natural, 22, 7-44.

07

Muoz Pizarro C (99) Sinopsis de la Flora Chilena. (nd edition 9), Ediciones de la Universidad de Chile. Muoz Pizarro C (9) Flores Silvestres de Chile. Mnchen. Muoz Pizarro C (99) El Archipilago de Juan Fernndez y la conservacin de sus recursos naturales renovables. Reprint 974, Museo Nacional de Historia Natural (Santiago) Serie Educativa, 9, 7-47. Muoz Pizarro C (97) Chile: plantas en extincin. Santiago. Muoz Pizarro C & E Pisano (1947) Estudio de la vegetacin y flora de los Parques Nacionales Fray Jorge y Talinay. Agricultura Tcnica (Santiago), 7, 7-90. Muoz-Schick M (980) Flora del Parque Nacional Puyehue. Ed. Universitaria, Santiago. Muoz-Schick M & A Moreira-Muoz (000) Los Gneros Endmicos de Monocotiledneas de Chile Continental. Museo Nacional de Historia Natural, Chile. Online version: http://www.mnhn.cl/apuntes/botanica/Principalbot.htm Muoz-Schick M & A Moreira-Muoz (00) El Herbario Nacional y la Conservacin de la Flora Chilena. Museo Nacional de Historia Natural, Chile. Online version: http://www.mnhn.cl/botanica/Herbario/index.html Muoz-Schick M, R Pinto, A Mesa & A Moreira-Muoz (00) Oasis de neblina en los cerros costeros del sur de Iquique, regin de Tarapac, Chile, durante el evento El Nio 997998. Revista Chilena de Historia Natural, 74, 8940. 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. 779. 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, 89409. 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 (98b) 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, 79-740. 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, -.

08

Ochsenius, C. (999) The Neotropical Aridity. CCOS PCSS Geowissenschaftliche Beitrge, Baden-Wrttemberg, 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, 47-44. 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 AndeanPatagonian forest. Journal of Biogeography, 29, 778. Pea 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 Flix. 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) Fitogeografa de Fuego-Patagonia chilena I. Las comunidades vegetales entre las latitudes  y  S. Anales Instituto de la Patagonia, 8, -0. Pisano E (1981) Bosquejo fitogeogrfico de Fuego-Patagonia. Anales Instituto de la Patagonia, 12, 9-7. Pisano E (98) Comunidades vegetales en el sector Norte de la Pennsula Muoz Gamero (ltima Esperanza, Magallanes). Anales Instituto de la Patagonia, 14, 8-0. Pliscoff P (00) Priorizacin de reas para fortalecer la conservacin de la flora arbrea nativa en la zona mediterrnea de Chile. MScThesis. Universidad de Chile, Santiago. Poole I, AMW Mennega & D Cantrill (00) Valdivian ecosystems in the Late Cretaceous and Early Tertiary of Antartica: further evidence from myrtaceous and eucryphiaceous fossil wood. Review of Palaeobotany and Palynology, 124, 97. Posada D & Crandall KA (00) Intraspecific gene genealogies: trees grafting into networks. Trends in Ecology and Evolution, 16, 74. 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.

09

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. Pulqurio 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, 94. 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 bioclimtica de Chile Central. Revista Geogrfica de Valparaso, 5, -8. Quintanilla V (1983) Biogeografa. Coleccin Geografa de Chile. Instituto Geogrfico Militar, Santiago. Quintanilla V (1985) Carta fitogeogrfica de Chile mediterrneo. Contribuciones Cientficas y Tecnolgicas, Area Geociencias (U. de Santiago), 70, -0. Quintanilla V (1988) Fitogeografa y cartografa de la vegetacin de Chile rido. Contribuciones Cientficas y Tecnolgicas, 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. Lauppsche Buchhandlung, Tbingen. 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, 44. Raven PH & DI Axelrod (974) Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden, 61, 97. 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, 780. Reiche K (89-9) Flora de Chile, 6 vols. Santiago. Reiche K (1905) La distribucin geogrfica de las Compuestas de la flora de Chile. Anales del Museo Nacional de Chile, Seccin Botnica, 17, -4. Reiche K (907) Grundzge der Pflanzenverbreitung in Chile. Die Vegetation der Erde: Sammlung Pflanzengeographischer Monographien, Vol. VIII. (ed. by A Engler & O Drude) Verlag von V. Engelmann, Leipzig.

0

Renner SS (004) Variation in diversity among Laurales, Early Cretaceous to Present. Biol. Skr. 55, 44-48. 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 clsico de la fitogeografa de Argentina: un anlisis crtico. 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, 094. Richter M (99) Klimakologische Merkmale der Kstenkordillere in der Region Antofagasta (Nordchile). Geokodynamik, 16, 8-. Richter M (997) Allgemeine Pflanzengeographie. Teubner Studienbcher der Geographie, Stuttgart. Richter M, T Fickert & F Grninger (2004) Pflanzen auf schuttbedeckten Gletschern wandernde Kuriositten. Geoko, 25, -. Riddle BR (00) Is biogeography emerging from its identity crisis? Journal of Biogeography, 32, 88. Rivas-Martnez S & G Navarro (000). Figure I.4, in G Navarro & M Maldonado (00) Geografa Ecolgica de Bolivia: Vegetacin y Ambientes Acuticos. Ed. Centro de Ecologa Simn I. Patio, 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, 747. 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. 4748. Chapman & Hall, London. Rosen BR (99) Empiricism and the biogeographical black box: concepts and methods in marine palaeobiogeography. Palaeogeography, Palaeoclimatology, Palaeoecology, 92, 70. Rothmaler W (9) Allgemeine Taxonomie und Chorologie der Pflanzen. Grundzge 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, 484. 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 vegetacin del Parque Nacional Pan de Azcar en el Desierto de Atacama. Gayana Botnica, 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 Fernndez 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, 009. Sanderson MJ (00) Molecular data from 7 proteins do not support a Precambrian origin of land plants. American Journal of Botany, 90, 949. 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 Wikstrm & K Bremer (004) Molecular Evidence On Plant Divergence Times. American Journal of Botany, 91, . Sanmartn I & F Ronquist (004) Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology, 53, 4. Sanmartn 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, 984. 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, 48-40. Schmithsen J (9) Die rumliche Ordnung der chilenischen Vegetation. Bonner Geographische Abhandlungen, 17, 8. Schmithsen J (98) Allgemeine Vegetationsgeographie. d ed., De Gruyter, Berlin. Schneider H, E Schuettpelz, KM Pryer, R Cranfill, S Magalln & R Lupia (2004) Ferns diversified in the shadow of angiosperms. Nature, 428, 7. Schnfelder P & A Bresinsky (990) Verbreitungsatlas der Farn- und Bltenpflanzen Bayerns. Ulmer Verlag. Schouw JF (8) Grundzge einer allgemeinen Pflanzengeographie (+ Atlas). Reimer, Berlin. Schroeder F-G (998) Lehrbuch der Pflanzengeographie. Quelle & Meyer, Wiesbaden. Schrter C (1913) Genetische Pflanzengeographie (Epiontologie). Handwrterbuch 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 (98) Phytochorologie als Wissenschaft, am Beispiele der vorderasiatischen Flora. Feddes Rep. Beiheft 00, Dahlem, Berlin. Sclater PL (88) 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, 52,004.



Sempere T, A Hartley & P Roperch (00) Comment on Rapid Uplift of the Altiplano Revealed Through C-8O Bonds in Paleosol Carbonates. Science, 314, 70b. 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. 474, 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 superpramo. Flora, 202, 0. Skottsberg C (90) Vegetationsbilder aus Feuerland, von den Falkland-Inseln und von Sdgeorgien. Karsten und Schenck Vegetationsbilder, 4 (-4),  lm. Skottsberg C (90 a) Botanische Ergebnisse der schwedischen Expedition nach Patagonien unde dem Feuerlande 9071909. I. bersicht ber die wichtigsten Pflanzenformationen Sdamerikas S von 41, ihre geographische Verbreitung und Beziehungen zum Klima. Kungl. Svenska Vetenskapakademiens Handlingar, 46, (), -8 +  map. Skottsberg C (90 b) Vegetationsbilder von den Juan Fernndez Insel. Karsten und Schenck Vegetationsbilder, 8 (), 7-. Skottsberg C (9) Botanische Ergebnisse der schwedischen Expedition nach Patagonien und dem Feuerlande 907909. V. Die Vegetationsverhltnisse lngs der Cordillera de los Andes sdlich von 4 Grad sdlicher Breite. Ein Beitrag zur Kenntnis der Vegetation in Chilo, West-Patagonien, dem andinen Patagonien und Feuerland. Kungl. Svenska Vetenskapsakademien Handlingar, 56 () -,  lm. Stockholm, Almqvist & Wiksell. Skottsberg C (90-9) (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 poblacin botnica y zoolgica de las islas chilenas del Pacfico. Imprenta Universitaria, Santiago. Translated by G. Looser from the original French: Le peuplement des les pacifiques du Chili. Contribution ltude du peuplement zoologique et botanique des les du Pacifique (ed. by L Berland, J Berlioz, EH Bryan et al.) (94), pp. 780. Socit de Biogographie, Paris. Skottsberg C (97) Die Flora der Desventuradas-Inseln (San Flix und San Ambrosio) nach Sammlungen F. Johows. Gteborgs Kungl. Vetensk.-Vitterh.-Samh. Handl., ser, , B, 5, -88. Skottsberg C (9) Weitere Beitrge zur Flora der Insel San Ambrosio. Arkiv fr 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. 940, Almqvist & Wiksell, Uppsala.



Skottsberg C (1963) Zur Naturgeschichte der Inseln San Ambrosio (Islas Desventuradas, Chile) 2. Bltenpflanzen. Arkiv fr Botanik, 4, 44-490 (+ tables -). Sluys R (1994) Explanations for biogeographic tracks across the Pacific Ocean: a challenge for paleogeography and historical biogeography. Progress in Physical Geography, 18, 48. 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 Gutirrez & IR Hernndez (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) Evolucin de la Flora Chilena. Flora de Chile vol. 1 (ed. by C Marticorena & R Rodrguez) pp. 8-8, Universidad de Concepcin, Chile. Stuessy TF, KA Foland, JF Sutter, RW Sanders & M Silva (1984a) Botanical and geological significance of PotassiumArgon dates from the Juan Fernandez Islands. Science, , 49. Stuessy TF, RW Sanders & M Silva (1984b) Phytogeography and evolution of the flora of the Juan Fernndez 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.

4

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, 84. Stuessy TF, DJ Crawford, C Marticorena & M Silva O (998a) Isolating mechanisms and modes of speciation in endemic angiosperms of the Juan Fernndez 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 Mllner, 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, 879. Stuessy TF, G Jakubowsky, R Salguero G, M Pfosser, PM Schlter, TFer, B-Y Sun & H Kato (00) Anagenetic evolution in island plants. Journal of Biogeography, 33, 9-. Suessenguth K (98) Neue Ziele der Botanik. Lehmanns Verlag, Mnchen/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 Fernndez Islands, Chile. Pacific Science, 51, -. Swenson U, RS Hill & S McLoughlin (00) Biogeography of Nothofagus supports the sequence of Gondwana breakup. Taxon, 50, 004. Szumik CA, F Cuezzo, P Goloboff & AE Chalup (00) An optimality criterion to determine areas of endemism. Systematic Biology, 51, 808. Szumik C & P Goloboff (004) Areas of endemism: an improved optimality criterion. Systematic Biology, 53, 98977.

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 (99). 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 of Antarctica. Science, 257, 777. 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 gnero para Chile (Islas Desventuradas). Gayana Botnica, 54, -7.



Teillier S, A Hoffmann, F Saavedra & L Pauchard (994) Flora del Parque Nacional El Morado (Regin Metropolitana, Chile). Gayana Botnica, 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, 8970. Thorne RF (00) How many species of seed plants are there? Taxon, 51, -. Torres T & M Rallo (98) Anatoma de troncos fsiles del Cretcico Superior de Pichasca en el norte de Chile. Anais II Congreso Latinoamricano Paleontologa. 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 Geolgica 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. Rwer, Gttingen. Troncoso A & R Herbst (999) Triassic Ginkgoales from northern Chile. Revista Geolgica de Chile, 26, -7 Troncoso A & EJ Romero (1998) Evolucin de las comunidades florsticas en el extremo sur de Sudamrica durante el Cenoftico. Proceedings of the Congreso Latinoamericano de Botnica, No.  (ed. by R Fortunato & N Bacigalupo), pp. 49-7, Monographs in Systematic Botany from the Missouri Botanical Garden. Troncoso A, C Villagrn & M Muoz-Schick (980) Una nueva hiptesis acerca del origen y edad del bosque de Fray Jorge (Coquimbo, Chile). Boletn 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 40Ar9Ar 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) Whats 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 BBiological 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.



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, 99. Varela F (000) El Fenmeno de la Vida. Editorial Dolmen, Santiago de Chile. Veit H (994) Holocene landscape and climate evolution of the Central Andes. Zentralblatt Geologie und Palontologie, 7/8, 887-89. Verd M, P Dvila, P Garca-Fayos, N Flores-Hernndez & A Valiente-Banuet (00) Convergent traits of mediterranean woody plants belong to pre-mediterranean lineages. Biological Journal of the Linnean Society, 78, 447. Vester H (940) Die Areale und Arealtypen der Angiospermen-Familien. Doctoral thesis, Naturwissenschaftlichen Fakultt der Ludwig-Maximilians Universitt zu Mnchen. Akad. Verlagsgesellschaft Leipzig. Villa-Martnez R, C Villagrn & B Jenny (00) The last 700 cal yr B.P. of westerly rainfall in Central Chile inferred from a high-resolution pollen record from Laguna Aculeo (4S). Quaternary Research, 60, 849 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, 4484. Villagrn C (00) Flora y vegetacin del Parque Nacional Chilo: gua de excursion botnica por la Cordillera de Piuchu. Conaf, Chilo. Villagrn C (00) Botnica Indgena de Chile. Revista Chagual (Santiago), 1, 0-. Villagrn C & LF Hinojosa (1997) Historia de los bosques del sur de Sudamrica, II: Anlisis fitogeogrfico. Revista Chilena de Historia Natural, 70, 4-7. Villagrn C, A Len & FA Roig (004) Paleodistribucin del alerce y ciprs de las Guaitecas durante perodos interestadiales de la Glaciacin Llanquihue: provincias de Llanquihue y Chilo, Regin de Los Lagos, Chile. Revista Geolgica de Chile, 31, -. Vinnersten A & K Bremer (00) Age and Biogeography of Major Clades in Liliales. American Journal of Botany, 88, 970. 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. Englers Botanische Jahrbcher, 17, -4. von Ihering H (907) Archhelenis und Archinotis: Gesammelte Beitrge zur Geschichte der Neotropischen Region. W. Engelmann, Leipzig. von Wettstein R (898) Grundzge der geographisch-morphologischen Methode der Pflanzensystematik. Jena, Verlag Gustav Fischer.

7

Wagner M (88) 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, 447. 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 (94) Grundlagen der Pflanzenverbreitung. Einfhrung in die Pflanzengeographie f. Studierende der Hochschulen, 2.Teil Arealkunde. Walter H & H Straka H (970) Arealkunde: Floristisch-historische Geobotanik. d. edition. Series Einfhrung 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, 74. Wangerin W (9) Florenelemente und Arealtypen. Beitrge 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, 979987. 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, 47. 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, 07. 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, 77-7. Wellman CH, PL Osterloff & U Mohiuddin (00) Fragments of the earliest land plants. Nature, 425, 8-8. Wel W & R Lindacher R (994) Beitrge zur Chorologie und Florenstatistik der Kanarischen Inseln. Hoppea, Denkschr. Regensb. Bot. Ges., 55, 84-87.

8

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. Mnchen, 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. Wikstrm 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, 778. Wikstrm 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 Ridleys 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, 78. 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 (90) An introduction to historical plant geography. The Chronica Botanica Company. Waltham, Massachusets.

Yez GA, CR Ranero, R von Huene & J Daz (00) Magnetic anomaly interpretation across the southern central Andes (-4S): The role of the Juan Fernndez Ridge in the late Tertiary evolution of the margin. Journal of Geophysical Research-Solid Earth, 106, -4.

9

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, 00-08. Zimmermann W (99) 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 Rodrguez (999) Anlisis de la biodiversidad en plantas vasculares de la Argentina. Kurtziana, 27, 7-7. Zunino M & A Zullini (00) Biogeografa: la dimensin espacial de la evolucin. Fondo de Cultura Econmica, Mxico. Transl. from the Italian edition, 99.

0

Appendix A: List of Chilean genera, geographic distribution, floristic elements



GENUS FAMILIA Orchidaceae Asteraceae Fabaceae Rosaceae Verbenaceae Asteraceae Asteraceae Asteraceae Euphorbiaceae Fabaceae Pteridaceae Aextoxicaceae Orobanchaceae Asteraceae Asteraceae Poaceae Rosaceae Alismataceae Nyctaginaceae Solanaceae Scrophulariaceae Poaceae Verbenaceae Alstroemeriaceae Amaranthaceae Amaranthaceae Asteraceae Asteraceae Myrtaceae Poaceae Cyperaceae 1 12 2 43 1 60 100 75 30 Am: SW US to Chile & Arg Chile, S Brasil, Per, Arg Tropics & subtrop, S Am, N Am, Aus, 1 sp Caucasus Subtrop & temp, Euro, S Am, N Am, 16 sp Russia, 1 sp. Calif, NW Mexico, Chile Canada, Mexico, SA, West Indies Temp S Am: Arg, Chile Aus, NZ, S Am Canada, USA, Arg, Chile AUS ANT-P ANT-P W-NT 36 16 6 Chile Trop Am Mexico, C Am, Chile, Bol, 2 sp S Afr warm temp & subtrop regions & montane tropics W-NT W-ANT W-NT S-AMZ 3 SW & C US, Mexico to Chile & Arg 10 250 temp N hemis, Middle Asia, montane tropics, S Afr temp N Hemishp, Euro, Middle asia, Australia, S Am ANT-P 220 temp & warm regions, montane tropics, JF, IP 11 9 sp W N Am, Mexico, 2 sp Chile, Arg ANT-P W-ANT W-ANT 290 E USA, C & W S Am, West Indies W-NT 40 Trop, warm C & S Am, USA ANT-P 4 3 4 4 4 7 4 6 3 4 3 3 1 7 4 3 5 2 4 1 Chile, Arg 5 150 Cosmop, esp Neotrops, Madag 7 6 1 1 1 2 24 3 1 1 6 1 4 3 33 2 2 1 2 2 1 1 230 Peru, Chile, Arg, S Brasil, TFuego S-AMZ 3 132 1 Chile 6 1 5 W NA, Mexico, Hawaii, Guatemala, Chile, Arg, Himalayas, E Asia W-ANT 4 1 -36,83 -30,50 -18,20 -21,27 -30,67 -29,00 -25,00 -30,95 -31,00 -18,20 -33,43 -20,17 -18,83 -29,50 -31,50 -20,50 -20,33 -18,50 -33,23 -22,08 -17,58 -35,33 -35,40 -27,00 2 Central Chile 6 2 -31,62 30 Tropics Afr, Am, Madagascar, 1 1 -18,23 6 Arid S Am, Arg, Chile, Bol AND 3 3 -18,42 -30,00 -18,23 -45,50 -55,00 -36,83 -54,83 -45,40 -43,67 -30,00 -36,67 -56,00 -56,00 -18,23 -40,00 -23,50 -32,00 -36,67 -53,67 -30,50 -54,00 -28,50 -40,35 -30,65 -42,72 -46,83 -43,00 -55,00 100 S Afr (1), New Guinea, Aus, Tasmania, NZ, subantarctic Is, Polynesia, Hawaii, California to SA, JF 4 19 -22,00 -56,00 1200 Tropics, Subtrop, esp Afr y Aus 1 1 -27,25 -37,90 -32,58 -39,00 -24,21 -18,23 -38,56 -45,92 -33,67 -36,52 -33,33 -37,17 -29,50 -30,83 -43,48 -43,50 -18,22 -36,72 -21,83 -25,42 -33,08 -42,58 -25,50 -37,17 -23,50 -36,79 -26,37 -30,15 -41,08 -39,20 -41,00 20 New Guinea, Aus, Tasmania, NZ, S Am (TFuego), JF, Falkland Is. SPT 2 4 -43,00 -55,75 -49,38 27 Andes Colombia to Argentina AND 3 1 -17,58 -24,50 -21,04 6,92 12,75 10,65 34,00 11,58 0,00 13,88 18,17 6,33 36,63 24,13 13,00 1,00 11,67 25,05 25,00 0,03 6,57 3,33 13,17 7,17 22,17 10,00 33,67 10,00 7,12 8,57 25,13 11,50 7,60 28,00 N SP DISTRIB TRACK ELEM SP CHILE N_MAX S_MAX MittelW Range

CLASE A

ORDEN

Aa

Monocots

Asparagales

Abrotanella

Asterids

Asterales

Acacia

Rosids

Fabales

Acaena

Rosids

Rosales

Acantholippia

Asterids

Lamiales

Achyrocline

Asterids

Asterales

Acrisione

Asterids

Asterales

Adenocaulon

Asterids

Asterales

Adenopeltis

Rosids

Malpighiales

Adesmia

Rosids

Fabales

Adiantum

Filicopsida

Pteridales

Aextoxicon

Eudicots

Berberidopsidales

Agalinis

Asterids

Lamiales



Ageratina

Asterids

Asterales

Agoseris

Asterids

Asterales

Agrostis

Monocots

Poales

Alchemilla

Rosids

Rosales

Alisma

Monocots

Alismatales

Allionia

Eudicots

Caryophyllales

Alona

Asterids

Solanales

Alonsoa

Asterids

Lamiales

Alopecurus

Monocots

Poales

Aloysia

Asterids

Lamiales

Alstroemeria

Monocots

Liliales

Alternanthera

Eudicots

Caryophyllales

Amaranthus

Eudicots

Caryophyllales

Amblyopappus

Asterids

Asterales

Ambrosia

Asterids

Asterales

Amomyrtus

Rosids

Myrtales

Amphibromus

Monocots

Poales

Amphiscirpus

Monocots

Poales

Amsinckia Boraginaceae Myrsinaceae Fabaceae Poaceae Malvaceae Primulaceae Ranunculaceae Phytolaccaceae Asteraceae Poaceae Poaceae Santalaceae Rosaceae Asteraceae Apiaceae Restionaceae Corsiaceae Vivianiaceae Araucariaceae Caryophyllaceae Papaveraceae Bignoniaceae Asteraceae Poaceae Aristolochiaceae Elaeocarpaceae Santalaceae Plumbaginaceae Asteraceae Oleandraceae Aspleniaceae Asteliaceae Asteraceae 25 250 650 15 390 90 10 Temp S Am, Brazil SAm Andes to Tfuego, Euro, Russia temp Eurasia & W N Am, N + S Afr, S Am New Guinea, NZ, Juan Fernandez, Madag Subcosmop, Madag, JF Masc, New Guinea, Aus, NZ, Polinesia, Chile temp Eurasia, S Afr, Madag, Am, Hawaii SPT W-ANT 5 300 290 20 Colombia, Ecuador, Peru, Chile N Am, Medit, temp Asia, tropics & subtropics trop, subtrop, some in temp regions E Aus, Tasman, NZ, Arg, Chile AUS S-AMZ W-ANT W-ANT SPT 12 28 N & S AM, WI, Hawaii S Peru, N C Chile & Arg, A. uspallatensis S Bol 160 America, temp Eurasia, N Afr 18 1 Chile New Guinea, Aus, New Caledonia, S Chile, SE Brasil SPT W-ANT W-NT AND AND 1 Chile, Arg 10 SE Asia, Malesia, New Guinea, Aus, Tasm, NZ, Chile, Arg SPT 27 Euro, Medit, N & S Afr, Madag, W & E Asia, Malesia, Aus, N, C & S Am, JF, IP 7 2 5 6 2 4 3 3 3 7 1 2 3 4 4 2 7 2 4 5 Andes S Bolivia, N Chile, NW Arg AND 3 20 Am, Euro, Medit, Ethiop, Australia, Caucasus 7 3 1 8 1 1 1 1 7 4 11 1 3 2 1 3 1 2 1 7 1 4 -18,00 -40,17 -30,65 -55,92 -56,00 -56,00 -36,96 -48,08 -43,33 8 W trop SA: S Mexico, Guatem, Costa Rica, Colom, Venez, Ecuad, Peru, Bol, Bras, Chile, Cuba, Haiti, P.Rico W-NT 3 1 20 temp & warm Eurasia, N Afr, montane tropics 7 7 -36,50 -37,67 -33,40 -18,82 -19,80 -36,83 -33,00 -30,17 -37,50 -18,00 -18,48 -20,82 -29,50 -18,83 -27,50 -30,83 -31,00 -30,67 -17,75 1 Andes Chile Peru AND 3 1 -18,38 50 Arctic & temp Euras, Andes S Am W-ANT 4 1 -31,00 2 Chile 6 2 -24,83 -35,97 -53,13 -21,50 -56,00 -42,37 -37,17 -27,78 -54,93 -51,00 -50,00 -30,83 -40,05 -55,00 -35,35 -34,47 -36,45 -37,83 -33,48 -44,00 -54,50 -56,00 -53,00 90 Euras, Sumatra, S & E Afr, NZ, A multifida disj N Am to Chile W-ANT 4 6 -30,83 -53,15 150 N Hemisphere, China, few sp SA (Tfuego) W-ANT 4 1 -53,17 -53,17 -53,17 -41,99 -30,40 -42,07 -19,94 -46,25 -40,02 -35,28 -23,30 -37,37 -43,92 -41,50 -30,50 -38,78 -36,50 -26,92 -27,64 -32,98 -28,33 -30,49 -37,42 -42,75 -43,33 -35,38 4 Chile, Peru, Bol AND 3 1 -28,58 -36,90 -32,74 11 Arg, Bol, Chile, Ecuador, Peru AND 3 4 -19,83 -25,50 -22,67 5,67 8,32 0,00 22,32 11,13 22,13 3,12 19,50 4,70 3,77 8,97 35,13 14,17 17,00 0,67 2,55 37,00 16,87 13,65 6,95 19,00 5,98 13,17 23,50 25,33 35,25 0,00 37,92 15,83 25,35 15 Andes Chile Arg 5 5 -30,47 -52,67 -41,57 22,20 20 Euro, Russia, N & S Afr, Madag, W Asia, Himalayas, N Aus, 2 sp. S Am 7 2 -25,00 -55,00 -40,00 30,00 15 W US, W temp S Am ANT-P 4 2 -22,25 -53,00 -37,63 30,75

Asterids

Unplaced

Anagallis

Asterids

Ericales

Anarthrophyllum

Rosids

Fabales

Anatherostipa

Monocots

Poales

Andeimalva

Rosids

Malvales

Androsace

Asterids

Ericales

Anemone

Eudicots

Ranunculales

Anisomeria

Eudicots

Caryophyllales

Antennaria

Asterids

Asterales

Anthochloa

Monocots

Poales

Anthoxanthum

Monocots

Poales

Antidaphne

Eudicots

Santalales

Aphanes

Rosids

Rosales

Aphyllocladus

Asterids

Asterales

Apium

Asterids

Apiales



Apodasmia

Monocots

Poales

Arachnitis

Monocots

Liliales

Araeoandra

Rosids

Geraniales

Araucaria

Pinopsida

Pinales

Arenaria

Eudicots

Caryophyllales

Argemone

Eudicots

Ranunculales

Argylia

Asterids

Lamiales

Aristeguietia

Asterids

Asterales

Aristida

Monocots

Poales

Aristolochia

Magnoliids

Piperales

Aristotelia

Rosids

Oxalidales

Arjona

Eudicots

Santalales

Armeria

Eudicots

Caryophyllales

Artemisia

Asterids

Asterales

Arthropteris

Filicopsida

Davalliales

Asplenium

Filicopsida

Aspleniales

Astelia

Monocots

Asparagales

Aster

Asterids

Asterales

Asteranthera Gesneriaceae Apiaceae Fabaceae Amaranthaceae Cactaceae Cupressaceae Euphorbiaceae Poaceae Salicaceae Azollaceae Apiaceae Asteraceae Gratiolaceae Asteraceae Plumbaginaceae Ledocarpaceae Fabaceae Ranunculaceae Orobanchaceae Lauraceae Asteraceae Solanaceae Berberidopsidaceae Berberidaceae Asteraceae Orchidaceae Blechnaceae Asteraceae Myrtaceae Loasaceae Urticaceae Nyctaginaceae Apiaceae Alstroemeriaceae Calyceraceae Lardizabalaceae Ophiglossaceae 13 1 35 100 5 35 50 12 Pantrop, JF tropics & subtropics Temp SA Mex to trop Am Andes, Arg, S Brasil Arg, Chile temp, polar, trop montane W-ANT W-NT S-AMZ 17 3 2 California, 1 Chile trop & subtro S Am, WI Brasil, Uruguay, Paraguay, Argentina, Chile 175 Subcosmop, Madag, JF ANT-P AND S-AMZ 8 Temp S Am: Chile, Arg 240 cosmop 470 Euras, N Afr, N & S AM 2 12 S Chile-Arg, Patagonia B. beckleri en Aus: Queensland & New S Wales, B. corallina: Chile AUS W-ANT 1 Andes Arg, Chile 230 pantropics, Mex to S Bras, Chile, Aus, NZ, Afr, Madag 60 Circumboreal, Eur, Medit, Afr, Andes 3 Arg, Chile AND W-ANT 1 Chile 6 3 4 1 5 5 2 4 7 5 7 4 3 3 1 1 5 3 3 5 4 8 Peru, Bol, Chile, Arg AND 3 1 Chile 6 1 3 1 3 3 2 1 4 1 47 7 4 10 1 1 3 1 2 2 3 8 1 1 -20,50 -49,15 -18,00 -36,00 -35,83 -50,50 -37,00 -55,52 -39,00 -54,00 -42,70 -55,67 -28,75 -52,33 -28,50 -45,00 -39,27 -53,08 10 SW US, Mex, Chile ANT-P 4 1 60 Trop & Subtrop, Hawaii, Madag, Aus 1 1 400 AM, esp S AM W-NT 3 43 -17,58 -20,00 -21,27 -23,50 -18,00 -27,37 -29,82 -18,00 -32,55 -34,83 -46,58 -37,13 -25,02 -17,58 -25,00 -30,67 -33,42 -32,50 -33,75 70 Andes to temp S Am, Malvinas, Antarctic Islands AUS 2 14 -18,00 6 trop & warm, Euro, Asia, Madag, Tristan da Cunha, Hawaii 7 1 -18,18 -42,55 -55,50 -56,00 -30,50 -36,62 -30,00 -30,00 -29,83 -34,25 -36,50 -36,67 -40,75 -53,67 -37,67 -55,97 -37,17 -40,50 -55,97 -36,60 -41,30 -45,50 10 8 temp Chile-Arg, s subtrop NW Arg, Bol, SE Bra, Uruguay, JF S-AMZ 3 8 -30,00 -46,00 110 pantrops, IP 1 1 -38,00 -30,37 -36,75 -36,79 -25,25 -28,94 -26,75 -24,00 -28,60 -32,03 -27,25 -34,61 -37,79 -50,13 -37,40 -40,49 -27,38 -32,75 -43,32 -35,01 -36,90 -39,63 1 Chile 6 1 -33,83 -34,10 -33,97 1 Arg, Chile 5 1 -32,65 -40,22 -36,43 5 Arg, Chile 5 2 -33,78 -46,50 -40,14 12,72 7,57 0,27 0,00 16,00 24,37 37,50 38,42 10,50 15,35 6,50 12,00 2,47 4,43 18,50 4,12 5,92 7,08 0,53 30,95 19,58 15,50 25,30 3,18 8,80 11,75 0,00 16,50 6,37 21,00 18,00 6,87 5,17 250 cosmop, Eurasia, Aus, ID 7 20 -18,00 -53,33 -35,67 35,33 2000 N Am, Andes, temp Eurasia, montane trop Afr W-ANT 4 46 -17,58 -53,50 -35,54 35,92 8 Mexico to Patagonia W-NT 3 4 -25,20 -37,83 -31,52 12,63 1 Chile, Arg 5 1 -35,60 -47,00 -41,30 11,40

Asterids

Lamiales

Asteriscium

Asterids

Apiales

Astragalus

Rosids

Fabales

Atriplex

Eudicots

Caryophyllales

Austrocactus

Eudicots

Caryophyllales

Austrocedrus

Pinopsida

Pinales

Avellanita

Rosids

Malpighiales

Axonopus

Monocots

Poales

Azara

Rosids

Malpighiales

Azolla

Filicopsida

Salviniales

Azorella

Asterids

Apiales

Baccharis

Asterids

Asterales

Bacopa

Asterids

Lamiales

Bahia

Asterids

Asterales

Bakerolimon

Eudicots

Caryophyllales

Balbisia

Rosids

Geraniales

Balsamocarpon

Rosids

Fabales

4

Barneoudia

Eudicots

Ranunculales

Bartsia

Asterids

Lamiales

Beilschmiedia

Magnoliids

Laurales

Belloa

Asterids

Asterales

Benthamiella

Asterids

Solanales

Berberidopsis

Eudicots

Berberidopsidales

Berberis

Eudicots

Ranunculales

Bidens

Asterids

Asterales

Bipinnula

Monocots

Asparagales

Blechnum

Filicopsida

Blechnales

Blennosperma

Asterids

Asterales

Blepharocalyx

Rosids

Myrtales

Blumenbachia

Asterids

Cornales

Boehmeria

Rosids

Rosales

Boerhavia

Eudicots

Caryophyllales

Bolax

Asterids

Apiales

Bomarea

Monocots

Liliales

Boopis

Asterids

Asterales

Boquila

Eudicots

Ranunculales

Botrychium

Filicopsida

Ophioglossales

Bouteloua Poaceae Apiaceae Asteraceae Orchidaceae Sapindaceae Poaceae Poaceae Cactaceae Scrophulariaceae Cyperaceae Zygophyllaceae Fabaceae Loasaceae Poaceae Portulacaceae Calceolariaceae Cunoniaceae Fabaceae Plantaginaceae Asteraceae Poaceae Ranunculaceae Calyceraceae Iridaceae Convolvulaceae Onagraceae Bignoniaceae Brassicaceae Caryophyllaceae Cyperaceae Cyperaceae Aizoaceae Orobanchaceae Poaceae Asteraceae 4 500 200 30 15 2000 cosmop, JF trop & S Afr, Madag, japan, New Guinea, Aus, Tasm, NZ, temp S Am S Afr, Aus, Tasman, NZ, Calif, Chile Euras, E N Am, C & S Am temp Eurasia, N Am, Chile Eurasia, N China, N & tro Afr, Aus, NZ, N Am W-ANT W-ANT 6 W N Am to Chile 200 1 Arg, Chile: C. valdivianum Cosmop, temp N hemis & montane tropics, JF W-ANT ANT-P 62 W N Am, Mex, S Am temp 25 15 Trop & Subtrop Am Cosmop, temp & trop hemis N & S, JF ANT-P 17 Temp S Am, Brazil 10 Arctic & temp N Hemis, NZ, temp S AM 1 Arg, Bras, Chile, Uruguay 1 Chile S-AMZ W-ANT S-AMZ W-NT 17 Cosmop 200 C & S Am, Afr, Madag, trop Asia 11 Philippines, Sulawesi, Moluccas, New Guinea, trp Aus, S Chile Arg SPT 2 1 7 6 3 4 3 3 7 4 5 4 4 7 1 4 4 4 7 300 AM from Mex to Tfuego W-NT 3 13 Am W USA to Chile W-NT 3 11 50 1 1 5 1 1 3 11 1 2 1 1 20 3 55 1 1 4 2 7 230 temp & subtrop reg & montane tropics 1 24 56 Peru, Bol, Arg, Chile, 1 sp Urug, Bras, Ecuad S-AMZ 3 13 150 Pantropics, subtropics, Arg, Madag, S Afr, Arabia 1 2 -19,47 -17,58 -18,33 -18,00 -18,17 -35,33 -29,00 -25,00 -32,50 -34,40 -29,88 -22,00 -31,00 -32,88 -27,37 -35,83 -30,00 -18,00 -21,00 -40,50 -23,50 -31,50 -18,33 -22,00 8 S Am: Venezuela Colombia disjunct to Chile-Arg, Brazil AND 3 1 -27,33 90 tropics & subtropics 1 1 -37,33 90 E Asia, tro & subtrop AM, Afr 1 3 -27,28 -42,10 -39,67 -30,60 -30,67 -38,40 -55,00 -54,50 -55,00 -49,42 -30,50 -56,00 -35,50 -37,83 -55,97 -40,00 -36,67 -40,00 -43,17 -52,25 -56,00 -37,83 -56,00 -56,00 -40,00 -39,27 -55,63 -37,80 11 Peru, Bol, Chile AND 3 1 -18,67 -19,17 150 Eurasia, N & S Afr, montane trop Asia, Am, JF W-ANT 4 11 -18,20 -55,00 5 S US & subtrop S Am ANT-P 4 2 -31,67 -40,00 -35,83 -36,60 -18,92 -34,69 -38,50 -28,97 -25,07 -27,99 -36,67 -36,25 -36,58 -42,38 -29,75 -40,50 -34,00 -36,12 -42,93 -31,00 -33,83 -36,44 -35,27 -44,04 -43,00 -27,92 -38,50 -48,25 -31,75 -35,38 -36,98 -29,90 1 Chile 6 1 -28,57 -32,88 -30,73 22 trop Am esp. Brasil S-AMZ 3 1 -30,67 -43,62 -37,14 12,95 4,32 8,33 36,80 0,50 14,82 2,33 3,27 11,20 20,82 36,67 36,50 36,83 14,08 1,50 31,00 3,00 3,43 26,08 18,00 5,67 7,12 15,80 16,42 26,00 19,83 35,00 15,50 16,50 7,77 37,30 15,80 3 Temp SA, Chile-Arg 5 1 -30,83 -30,95 -30,89 0,12 15 N & S Am, B. incana S US W-NT 3 6 -18,17 -52,00 -35,08 33,83 40 Am from Canada to Argentina W-NT 3 1 -18,37 -24,00 -21,18 5,63

Monocots

Poales

Bowlesia

Asterids

Apiales

Brachyclados

Asterids

Asterales

Brachystele

Monocots

Asparagales

Bridgesia

Rosids

Sapindales

Bromidium

Monocots

Poales

Bromus

Monocots

Poales

Browningia

Eudicots

Caryophyllales

Buddleja

Asterids

Lamiales

Bulbostylis

Monocots

Poales

Bulnesia

Rosids

Zygophyllales

Caesalpinia

Rosids

Fabales

Caiophora

Asterids

Cornales

Calamagrostis

Monocots

Poales

Calandrinia

Eudicots

Caryophyllales

Calceolaria

Asterids

Lamiales

Caldcluvia

Rosids

Oxalidales



Calliandra

Rosids

Fabales

Callitriche

Asterids

Lamiales

Calopappus

Asterids

Asterales

Calotheca

Monocots

Poales

Caltha

Eudicots

Ranunculales

Calycera

Asterids

Asterales

Calydorea

Monocots

Asparagales

Calystegia

Asterids

Solanales

Camissonia

Rosids

Myrtales

Campsidium

Asterids

Lamiales

Cardamine

Rosids

Brassicales

Cardionema

Eudicots

Caryophyllales

Carex

Monocots

Poales

Carpha

Monocots

Poales

Carpobrotus

Eudicots

Caryophyllales

Castilleja

Asterids

Lamiales

Catabrosa

Monocots

Poales

Centaurea

Asterids

Asterales

Centaurium Gentianaceae Asteraceae Apiaceae Asteraceae Caryophyllaceae Ceratophyllaceae Solanaceae Asteraceae Asteraceae Poaceae Pteridaceae Amaranthaceae Asteraceae Asteraceae Asteraceae Asteraceae Euphorbiaceae Orchidaceae Poaceae Polygonaceae Saxifragaceae Asteraceae Poaceae Gentianaceae Apiaceae Vivianiaceae Vitaceae Portulacaceae Icacinaceae Onagraceae Cleomaceae Linaceae Orchidaceae Rhamnaceae Euphorbiaceae Polemoniaceae 5 5 15 3 1 275 42 20 49 200 tropics incl Australia N Am, Calif, Arizona, N Mxico & S Am, Peru, Bol, Arg, Chile Malesia, E Aus, New Cale, Fiji, Samoa, C & trop S Am N Am spec. California, 1 sp Chile/Arg tropics & subtropics, spec. Am temp S Am S trop & temp S Am, Brazil subtrop & temp S Am, Brazil, JF temp S Am, S Brasil W N Am, Bol to Patagonia S-AMZ S-AMZ S-AMZ ANT-P ANT-P SPT ANT-P 1 Chile 3 2 1 Euro & Medit, 1 Calif, W S Am Arg, Bol, Brasil, Chile, Paraguay, Uruguay 110 C & S Am, WI, JF 23 Andes S Am, Patagonia 64 N & temp S Am, Euro, N Afr, Asia, 50 arid & semiarid W Am, Chile 55 Cosmop ANT-P W-ANT AND W-NT W-ANT S-AMZ 50 Andes Chile Per, Arg, Brazil S-AMZ 20 Mexico 2, Peru, S Bras, Arg, Chile ANT-P 7 S Andes Chile, Argentina 5 4 3 7 4 4 3 3 4 3 6 1 4 2 4 1 5 3 3 3 4 3 S Andes Chile, Argentina 5 2 2 4 29 1 24 2 5 13 1 1 1 1 18 1 1 1 1 1 2 4 2 5 S Brasil, Bol, Paraguay, Urug, Chile, N C Arg, Falkland S-AMZ 3 3 9 S Peru, Bol, Chile, Arg AND 3 3 125 cosmop, JF, ID 7 12 150 Cosmop, esp Neotrops & S Afr, Madag 7 9 -18,00 -18,00 -17,72 -33,03 -52,62 -36,00 -22,50 -30,50 -18,47 -18,00 -52,00 -17,72 -30,67 -25,00 -20,82 -33,00 -30,00 -18,58 -30,67 -29,67 -20,22 -34,88 -35,00 -30,25 -23,50 -32,90 6 Central & S Am, USA ANT-P 4 1 -32,13 55 trop & subtrop Am, S USA W-NT 3 1 -33,03 44 S Peru, Bol, Chile, Arg AND 3 37 -17,58 -40,25 -36,67 -43,83 -46,55 -54,00 -23,58 -43,00 -53,25 -56,00 -38,00 -55,33 -18,47 -37,22 -56,00 -35,75 -46,67 -38,00 -40,00 -37,80 -43,00 -37,93 -38,75 -40,33 -26,33 -40,32 -56,00 -46,50 -46,67 -53,00 175 trop Am, esp Brasil & Andes W-NT 3 1 -18,75 -40,33 20 cosmop 7 1 -33,42 -36,82 -35,12 -29,54 -28,92 -34,85 -37,98 -32,28 -36,00 -20,65 -38,02 -52,93 -46,00 -30,25 -42,92 -18,47 -27,61 -54,00 -26,73 -38,67 -31,50 -30,41 -35,40 -36,50 -28,26 -34,71 -35,00 -23,28 -37,60 -45,50 -38,38 -35,08 -42,95 100 cosmop 7 2 -23,00 -33,33 -28,17 6 Madag, Mascarene Is, Asia, Aus, NZ, New Cale, Tahiti, Chile 1 1 -30,00 -41,83 -35,92 11,83 10,33 3,40 21,58 22,67 3,63 11,70 28,55 36,00 5,87 9,97 0,63 20,00 15,50 24,83 0,00 19,22 4,00 18,03 16,00 13,00 19,18 4,80 13,00 19,35 8,08 10,67 6,12 5,43 21,00 16,25 23,17 20,10 40 Pantrop to Chile, NZ, JF 1 1 -35,12 -43,00 -39,06 7,88 2 JF 6 2 0,00 60 Cosmop exc trop & S Afr, N hemisp 1 ext. to Aus, 1 to Chile 7 1 -25,00 -41,50 -33,25 16,50

Asterids

Gentianales

Centaurodendron

Asterids

Asterales

Centella

Asterids

Apiales

Centipeda

Asterids

Asterales

Cerastium

Eudicots

Caryophyllales

Ceratophyllum

Ceratophyllales

Cestrum

Asterids

Solanales

Chaetanthera

Asterids

Asterales

Chaptalia

Asterids

Asterales

Chascolytrum

Monocots

Poales

Cheilanthes

Filicopsida

Pteridales

Chenopodium

Eudicots

Caryophyllales

Chersodoma

Asterids

Asterales

Chevreulia

Asterids

Asterales

Chiliophyllum

Asterids

Asterales

Chiliotrichum

Asterids

Asterales

Chiropetalum

Rosids

Malpighiales



Chloraea

Monocots

Asparagales

Chloris

Monocots

Poales

Chorizanthe

Eudicots

Caryophyllales

Chrysosplenium

Eudicots

Saxifragales

Chuquiraga

Asterids

Asterales

Chusquea

Monocots

Poales

Cicendia

Asterids

Gentianales

Ciclospermum

Asterids

Apiales

Cissarobryon

Rosids

Geraniales

Cissus

Eudicots

Vitales

Cistanthe

Eudicots

Caryophyllales

Citronella

Asterids

Unplaced

Clarkia

Rosids

Myrtales

Cleome

Rosids

Brassicales

Cliococca

Rosids

Malpighiales

Codonorchis

Monocots

Asparagales

Colletia

Rosids

Rosales

Colliguaja

Rosids

Malpighiales

Collomia

Asterids

Ericales

Colobanthus Caryophyllaceae Solanaceae Tecophilaeaceae Convolvulaceae Asteraceae Cactaceae Rubiaceae Boraginaceae Asteraceae Coriariaceae Caryophyllaceae Cactaceae Poaceae Malvaceae Asteraceae Crassulaceae Brassicaceae Convolvulaceae Elaeocarpaceae Malvaceae Euphorbiaceae Rubiaceae Boraginaceae Lauraceae Pteridaceae Asteraceae Lamiaceae Cactaceae Convolvulaceae Poaceae Boraginaceae Cyperaceae Campanulaceae Dryopteridaceae Fabaceae 12 165 3 400 75 9 155 cosmop tropics & subtropics temp & subtrop reg trop, subtrop, temp regions, JF Chile temp & warm regions, montane tropics Am, Canada to Arg W-ANT W-NT 20 1 JF Bol, S Peru, N Chile AND 2 Ecuador, Peru, Chile, Arg 2 temperate N & S 230 100 W US, W temp S Am tropics & subtropics, Madag, S Afr, S India, Ceylon, Malaysia, Philippines, N Australia W-ANT AND 7 Chile, Arg 1250 tropics & subtropics AND ANT-P 75 Chile, Arg, Bol, Per, ID 5 Bol, N Arg, C Chile, Brasil 5 trop & subtrop S-AMZ AND 7 Andes Colombia to Chile AND 200 cosmop, Euro, A & S Afr, Madag, Aus, Am 7 3 1 3 3 1 3 4 1 4 3 6 3 7 1 7 7 6 4 3 55 S Afr, NZ, New Guinea, Aus, S Am 1 7 Chile, Arg 5 7 1 7 1 1 2 19 1 7 39 1 1 1 1 2 10 5 1 26 3 1 3 -18,00 -19,25 -18,47 -32,95 -20,00 -27,18 -18,00 -18,00 -33,00 -40,35 -33,43 -39,98 -43,00 -32,35 -55,00 -25,40 -25,50 -29,80 -25,95 -36,47 -31,50 -29,77 -36,50 -21,70 25 S Am, 4 NZ, 1 New Guinea AUS 2 6 12 Bol, S Peru, N Chile AND 3 1 13 S Am Andes Colom to Chile, Euro, Medit, Afr, Madag 7 3 -30,00 -19,75 -18,47 -30,67 -20,25 -18,00 -17,58 -20,45 -32,23 -18,17 -24,90 -22,08 -18,83 -30,33 -31,30 -17,72 15 Mex to Chile, W Medit, Himal to Japan, New Guinea, NZ. C ruscifolia disj NZ 7 1 -35,02 114 N & S Am W-NT 3 1 -18,17 -18,17 -43,83 -37,00 -20,00 -56,00 -42,53 -22,00 -56,00 -18,50 -35,33 -42,77 -35,63 -25,33 -33,83 -44,50 -40,25 -37,40 -24,50 320 trop & subtropics 1 1 -27,82 -31,83 90 E Malesia, Aus, NZ, Pacific, JF, 12 sp Hawaii SPT 2 2 -29,83 -18,17 -39,43 -33,50 -19,88 -37,23 -36,60 -21,13 -37,00 -18,04 -27,89 -37,50 -26,90 -25,12 -27,96 -31,67 -35,29 -34,35 -21,11 25 N Chile 6 25 -22,00 -31,00 -26,50 60 tropics & subtropics 1 27 -17,58 -55,00 -36,29 150 cosmop esp. temp, trop & subtrops 7 5 -25,00 -40,00 -32,50 5 Chile 6 5 -22,08 -37,95 -30,02 15,87 15,00 37,42 9,00 0,00 4,02 0,00 8,82 7,00 0,25 37,53 11,87 1,75 38,00 0,92 14,88 10,53 17,47 0,43 11,75 25,67 9,92 6,10 6,78 0,00 15,00 21,10 14,97 7,03 23,00 5,17 37,00 7,40 2 Andes Chile Arg 5 2 -37,00 -44,00 -40,50 7,00 20 S Pacific, Aus, NZ, temp S Am, Ecuador SPT 2 3 -17,58 -56,00 -36,79 38,42

Eudicots

Caryophyllales

Combera

Asterids

Solanales

Conanthera

Monocots

Asparagales

Convolvulus

Asterids

Solanales

Conyza

Asterids

Asterales

Copiapoa

Eudicots

Caryophyllales

Coprosma

Asterids

Gentianales

Cordia

Asterids

Unplaced

Coreopsis

Asterids

Asterales

Coriaria

Rosids

Cucurbitales

Corrigiola

Eudicots

Caryophyllales

Corryocactus

Eudicots

Caryophyllales

Cortaderia

Monocots

Poales

Corynabutilon

Rosids

Malvales

Cotula

Asterids

Asterales

Crassula

Eudicots

Saxifragales

7

Cremolobus

Rosids

Brassicales

Cressa

Asterids

Solanales

Crinodendron

Rosids

Oxalidales

Cristaria

Rosids

Malvales

Croton

Rosids

Malpighiales

Cruckshanksia

Asterids

Gentianales

Cryptantha

Asterids

Unplaced

Cryptocarya

Magnoliids

Laurales

Cryptogramma

Filicopsida

Pteridales

Cuatrecasasiella

Asterids

Asterales

Cuminia

Asterids

Lamiales

Cumulopuntia

Eudicots

Caryophyllales

Cuscuta

Asterids

Solanales

Cynodon

Monocots

Poales

Cynoglossum

Asterids

Unplaced

Cyperus

Monocots

Poales

Cyphocarpus

Asterids

Asterales

Cystopteris

Filicopsida

Dryopteridales

Dalea

Rosids

Fabales

Danthonia Poaceae Asteraceae Apiaceae Davalliaceae Asteraceae Dennstaedtiaceae Poaceae Brassicaceae Desfontainiaceae Loranthaceae Bromeliaceae Poaceae Convolvulaceae Dicksoniaceae Acanthaceae Poaceae Malpighiaceae Malpighiaceae Dioscoreaceae Verbenaceae Dryopteridaceae Apocynaceae Asteraceae Apiaceae Rhamnaceae Poaceae Juncaceae Poaceae Sapindaceae Apiaceae Stylidiaceae Asteraceae Blechnaceae Campanulaceae Brassicaceae Thymelaeaceae 1 11 350 12 2 2 4 Peru Chile Tasman, NZ, subantarctic S Am Andes Chile Arg Australia, NZ, Pacific islands, Hawaii, Easter Island, N Guinea W N Am, 1 Chile Arctic & temp n Hemis, Asia, montane C & S Am temp S Am 68 9 3 Andes S Am Am from Canada to Argentina, 1 sp Australia tropics & subtrop, espec Australia AND AUS AND SPT ANT-P W-ANT 16 Peru, Chile, Bol to California 8 Aus, NZ, S Am 3 temp S Am AUS ANT-P AND C-PAC 90 4 Chile, Arg trop Andes from Colombia to Bol & N Chile, 1 sp Costa Rica AND 300 cosmop 4 temperate S Am, Arg, Chile 400 trop, subtrop, warm-temp regions 1 Chile 6 1 5 7 5 3 5 2 4 3 4 1 3 2 3 2 4 4 5 1 Chile 6 1 Peru, Bol, Arg, Chile AND 3 200 tropics & subtropics 1 1 1 1 1 41 1 1 4 3 1 4 1 2 3 1 2 1 2 1 1 7 1 -31,50 -18,00 -46,33 -38,00 -55,00 -56,00 -34,75 -36,50 -51,17 -33,35 -17,58 -31,97 -28,72 -17,58 -18,00 -18,47 -31,00 -22,00 -37,80 -21,50 -47,00 -23,00 -34,47 -55,00 -18,20 -21,00 -46,50 -31,17 -25,43 -56,00 -30,67 -40,18 -20,29 -33,22 -41,86 -17,89 -19,50 -32,48 -31,08 -23,72 -46,90 -26,08 20 trop Am, Malesia, New Guinea, Aus, New Caled, NZ SPT 2 2 -24,50 -18,00 -27,50 -21,50 -23,50 -31,00 -25,50 -18,20 -31,50 -27,42 -42,00 -43,83 -25,00 -18,10 -29,50 -24,46 -32,75 -37,42 7 tropics & subtropics, JF 1 1 58 Toda America W-NT 3 1 -36,00 -25,00 13 Peru, Bol, SW Bra, N Chile, N Arg S-AMZ 3 1 -25,00 1 Chile 6 1 -35,60 -40,18 -26,17 -38,00 -40,83 1 Costa Rica to Cape Horn AND 3 1 -35,33 -53,00 40 S Am, N Am, Canary Is. W-ANT 4 14 -18,20 -53,15 35 temp & cool, espec N Euro, montane tropics W-ANT 4 16 -28,80 -56,00 45 trop to warm temp, Madag 1 1 -32,45 -35,85 -34,15 -42,40 -35,68 -44,17 -37,89 -25,58 -37,00 -32,92 11 JF 6 11 90 W Medit, Himal, Japan, Aus, Tahiti, Afr, Madag, IP 1 1 0,00 0,00 3,40 27,20 34,95 17,67 4,58 1,17 0,00 15,83 0,00 1,00 0,20 4,00 5,92 18,50 12,83 0,00 13,65 5,42 2,50 26,28 0,62 3,00 28,03 0,17 3,43 18,20 9,17 0,00 6,50 37,00 9,67 22 Euro, Medit, SW & S Asia, trop Afr. Aus, NZ, & Am 7 2 -25,38 -46,30 -35,84 20,92 40 Chile, Andes, SE Brazil S-AMZ 3 2 -32,88 -42,47 -37,68 9,58 20 N & S Am, Euro, Caucasus W-ANT 4 8 -32,00 -42,62 -37,31 10,62

Monocots

Poales

Dasyphyllum

Asterids

Asterales

Daucus

Asterids

Apiales

Davallia

Filicopsida

Davalliales

Dendroseris

Asterids

Asterales

Dennstaedtia

Filicopsida

Dennstaedtiales

Deschampsia

Monocots

Poales

Descurainia

Rosids

Brassicales

Desfontainia

Asterids

Unplaced

Desmaria

Eudicots

Santalales

Deuterocohnia

Monocots

Poales

Dichanthelium

Monocots

Poales

Dichondra

Asterids

Solanales

Dicksonia

Filicopsida

Dennstaedtiales

Dicliptera

Asterids

Lamiales

Dielsiochloa

Monocots

Poales

Dinemagonum

Rosids

Malpighiales

8

Dinemandra

Rosids

Malpighiales

Dioscorea

Monocots

Dioscoreales

Diostea

Asterids

Lamiales

Diplazium

Filicopsida

Dryopteridales

Diplolepis

Asterids

Gentianales

Diplostephium

Asterids

Asterales

Diposis

Asterids

Apiales

Discaria

Rosids

Rosales

Dissanthelium

Monocots

Poales

Distichia

Monocots

Poales

Distichlis

Monocots

Poales

Dodonaea

Rosids

Sapindales

Domeykoa

Asterids

Apiales

Donatia

Asterids

Asterales

Doniophyton

Asterids

Asterales

Doodia

Filicopsida

Blechnales

Downingia

Asterids

Asterales

Draba

Rosids

Brassicales

Drapetes

Rosids

Malvales

Drimys Winteraceae Droseraceae Caryophyllaceae Dryopteridaceae Solanaceae Euphorbiaceae Bignoniaceae Cactaceae Lomariopsidaceae Elatinaceae Cyperaceae Hydrocharitaceae Poaceae Apocynaceae Proteaceae Ericaceae Asteraceae Poaceae Ephedraceae Onagraceae Dioscoreaceae Equisetaceae Poaceae Phytolaccaceae Asteraceae Apiaceae Brassicaceae Asteraceae Asteraceae Cactaceae Geraniaceae Fabaceae Apiaceae Escalloniaceae Cunoniaceae Brassicaceae 6 6 40 240 4 75 Cosmop SW US, Mex, 1 sp Chile trop & temp, Eurasia, Am, Afr, Malesia, Aus, JF Andes S Am, SE Brasil, JF SE Aus, Tasm, Chile Andes Ecuador to Arg AND AUS AND ANT-P 35 200 Chile, Arg, S Peru 1 Tierra del Fuego Eurasia, N & S Am, Aus, JF W-ANT AND 2 Andes Chile Peru 9 Andes Chile Peru 10 N & S Am 2 Chile W-NT AND AND 300 temp & trop, ID 30 almost cosmop, Madag 3 Chile 164 temp Euro, N Am, arctic, montane trop 60 Eurasia, AM W-ANT W-ANT 35 temp Eurasia, Am, trop & subtrop 15 SW USA, Mexico, Chile, Peru, Galapagos ANT-P 4 7 4 4 6 7 7 6 3 3 3 5 4 3 7 4 7 3 2 3 3 N temp & Arctic, Europa, S Andes, Falkland, JF W-ANT 4 1 Chile y Argentina, 1 spp. 5 1 1 1 1 7 12 3 2 6 2 1 1 1 1 12 28 1 1 10 13 2 2 1 Chile, Arg 5 1 150 temp & subtrop, montane tropics 7 6 9 AM temp W-NT 3 1 -19,25 -22,00 -33,45 -35,25 -32,83 -19,28 -19,87 -18,17 -18,00 -25,00 -18,40 -18,17 -33,50 -23,53 -21,50 -19,80 -48,83 -29,00 -18,42 -32,75 -25,00 -25,20 -18,50 -36,08 -18,23 200 cosmop, JF 7 19 -17,67 12 temp, trop & subtrop regions W-ANT 4 1 -34,80 400 pantrops & subtrop, Madag, IP 1 3 -39,83 -43,83 -39,63 -55,00 -36,50 -53,00 -43,83 -55,67 -56,00 -30,67 -23,00 -53,00 -55,50 -37,67 -44,35 -38,00 -44,58 -25,92 -27,07 -19,80 -53,50 -56,00 -36,72 -33,38 -30,50 -40,60 -53,00 -45,50 -22,95 100 S Am AND 3 9 -18,50 -35,00 5 Peru, Chile, Arg AND 3 1 -32,93 -40,25 -36,59 -26,75 -41,83 -37,22 -36,33 -27,88 -37,50 -38,64 -45,46 -44,42 -24,98 -21,43 -35,58 -36,75 -31,33 -31,38 -28,08 -39,04 -24,73 -24,28 -19,80 -51,17 -42,50 -27,57 -33,07 -27,75 -32,90 -35,75 -40,79 -20,59 1 Andes Costa Rica, Ecuador, Chile, Arg, JF AND 3 1 -30,67 -55,00 -42,83 5 Andes Colombia to Argentina AND 3 1 -18,20 -18,83 -18,52 0,63 24,33 7,32 16,50 4,00 4,83 37,33 17,25 31,00 10,38 20,42 23,17 11,38 3,13 34,83 37,50 12,67 25,95 19,83 11,08 2,38 5,57 0,00 4,67 27,00 18,30 0,63 5,50 15,40 34,50 9,42 4,72 150 cosmop 7 1 0,00 50 pantrop 1 3 -22,00 -25,40 -23,70 3,40 115 cosmop, espec Aus, Tasm & NZ 7 1 -37,83 -56,00 -46,92 18,17 11 Mexico, C & S Am, JF W-NT 3 1 -30,33 -55,97 -43,15 25,63

Magnoliids

Canellales

Drosera

Eudicots

Caryophyllales

Drymaria

Eudicots

Caryophyllales

Dryopteris

Filicopsida

Dryopteridales

Dunalia

Asterids

Solanales

Dysopsis

Rosids

Malpighiales

Eccremocarpus

Asterids

Lamiales

Echinopsis

Eudicots

Caryophyllales

Elaphoglossum

Filicopsida

Blechnales

Elatine

Rosids

Malpighiales

Eleocharis

Monocots

Poales

Elodea

Monocots

Alismatales

Elymus

Monocots

Poales

Elytropus

Asterids

Gentianales

Embothrium

Eudicots

Proteales

Empetrum

Asterids

Ericales

Encelia

Asterids

Asterales

9

Enneapogon

Monocots

Poales

Ephedra

Gnetopsida

Ephedrales

Epilobium

Rosids

Myrtales

Epipetrum

Monocots

Dioscoreales

Equisetum

Sphenopsida

Equisetales

Eragrostis

Monocots

Poales

Ercilla

Eudicots

Caryophyllales

Erechtites

Asterids

Asterales

Eremocharis

Asterids

Apiales

Eremodraba

Rosids

Brassicales

Eriachaenium

Asterids

Asterales

Erigeron

Asterids

Asterales

Eriosyce

Eudicots

Caryophyllales

Erodium

Rosids

Geraniales

Errazurizia

Rosids

Fabales

Eryngium

Asterids

Apiales

Escallonia

Asterids

Unplaced

Eucryphia

Rosids

Oxalidales

Eudema

Rosids

Brassicales

Eulychnia Cactaceae Euphorbiaceae Orobanchaceae Convolvulaceae Solanaceae Solanaceae Asteraceae Zygophyllaceae Bromeliaceae Poaceae Cupressaceae Asteraceae Asteraceae Gratiolaceae Rosaceae Francoaceae Frankeniaceae Onagraceae Malvaceae Centrolepidaceae Asteraceae Rubiaceae Calyceraceae Asteraceae Asteraceae Ericaceae Orchidaceae Onagraceae Gentianaceae Gentianaceae Fabaceae Geraniaceae Alliaceae Rosaceae Proteaceae Polemoniaceae 3 25 50 2 400 Chile Arctic & temp regions New Guinea, Aus, Chile, Arg W N Am & S Am SPT ANT-P 3 256 360 9 15 Temp S Am, JF temp W N Am & W S Am Euro, NW Afr, Asia, E Aus, Am Eurasia, NW Afr, N, C & S Am, Aus, NZ S Am from Colombia & Venez to Arg, Brazil cosmop, espec tem regs & tropic montane AND ANT-P W-ANT 140 1 S Arg, Chile E & SE Asia, Malesia, Aus, NZ, N, C, S Am, WI, JF C-PAC 40 N & S Am, Brazil, JF 6 temp S Am W-NT 350 cosmop, JF 15 temp & subtrop C & S Am 3 New Guinea, NZ, Tasmania, TFuego, Falkland 14 Mexico, C Andes, Galapagos, ID 105 C & S Am, NZ, Tahiti AUS W-NT SPT W-NT 50 temp Euro, Afr, Aus, Am, ID 1 Chile 6 7 2 3 2 3 7 5 3 5 4 5 4 4 7 3 7 6 7 2 4 15 N & S Am, temp Euras W-ANT 4 1 Chile, Arg 5 1 1 1 4 2 3 1 2 17 6 22 1 11 12 2 6 3 1 5 2 4 1 4 32 SW USA, Mexico, Peru, Chile, Arg ANT-P 4 1 21 W N Am, Mexico, S Am, 1 sp Aus C-PAC 4 1 1 Chile Arg 5 1 -40,17 -19,00 -29,45 -40,75 -35,00 -32,23 -20,63 -28,50 -18,30 -39,88 -17,58 -18,00 -36,83 -17,58 -37,48 -30,67 -30,67 -28,50 -18,00 -21,00 -18,47 -17,67 -29,50 -33,95 -35,40 -18,00 500 cosmop 7 24 -17,67 1 Chile 6 1 -33,58 30 Medit, Euro, SW Asia, NE Afr, SW N Am, Peru, Chile W-ANT 4 2 -18,83 -31,67 -44,00 -56,00 -42,58 -36,60 -33,83 -40,75 -46,43 -43,50 -53,00 -55,00 -30,90 -56,00 -42,62 -56,00 -46,00 -55,25 -37,48 -56,00 -55,33 -37,00 -55,00 -56,00 -31,50 -55,50 -34,25 -55,00 -45,42 -51,00 3 S Brasil, Peru, Bol, Paraguay, Urug, Chile, Arg S-AMZ 3 2 -25,40 -39,82 15 S Peru, Bol, Chile, Arg AND 3 7 -18,20 -45,77 -31,98 -32,61 -25,25 -38,79 -36,83 -41,38 -27,80 -31,64 -40,75 -40,72 -37,87 -36,82 -41,75 -24,60 -47,94 -30,10 -37,00 -41,42 -36,42 -37,48 -43,33 -43,00 -32,75 -36,50 -38,50 -24,98 -36,58 -31,88 -44,48 -40,41 -34,50 6 S Am, Peru, Ecuador, Galapagos, N Chile AND 3 2 -18,38 -24,00 -21,19 100 trops & subtrops 1 1 -35,33 -37,83 -36,58 2,50 5,62 27,57 14,42 12,83 10,42 38,33 2,42 17,60 4,38 0,00 11,43 11,27 32,37 26,50 12,60 16,12 25,03 38,00 9,17 37,67 0,00 25,33 24,67 8,50 37,00 35,00 13,03 37,83 4,75 21,05 10,02 33,00 300 temp N Hemishp, Euro, Middle asia, Australia, NZ, temp S Am, JF W-ANT 4 15 -30,00 -55,00 -42,50 25,00 2000 subcosmop 7 19 -18,47 -52,67 -35,57 34,20 7 Peru, Chile AND 3 6 -18,50 -32,25 -25,38 13,75

Eudicots

Caryophyllales

Euphorbia

Rosids

Malpighiales

Euphrasia

Asterids

Lamiales

Evolvulus

Asterids

Solanales

Exodeconus

Asterids

Solanales

Fabiana

Asterids

Solanales

Facelis

Asterids

Asterales

Fagonia

Rosids

Zygophyllales

Fascicularia

Monocots

Poales

Festuca

Monocots

Poales

Fitzroya

Pinopsida

Pinales

Flaveria

Asterids

Asterales

Flourensia

Asterids

Asterales

Fonkia

Asterids

Lamiales

Fragaria

Rosids

Rosales

Francoa

Rosids

Geraniales

0

Frankenia

Eudicots

Caryophyllales

Fuchsia

Rosids

Myrtales

Fuertesimalva

Rosids

Malvales

Gaimardia

Monocots

Poales

Galinsoga

Asterids

Asterales

Galium

Asterids

Gentianales

Gamocarpha

Asterids

Asterales

Gamochaeta

Asterids

Asterales

Gamochaetopsis

Asterids

Asterales

Gaultheria

Asterids

Ericales

Gavilea

Monocots

Asparagales

Gayophytum

Rosids

Myrtales

Gentiana

Asterids

Gentianales

Gentianella

Asterids

Gentianales

Geoffroea

Rosids

Fabales

Geranium

Rosids

Geraniales

Gethyum

Monocots

Asparagales

Geum

Rosids

Rosales

Gevuina

Eudicots

Proteales

Gilia

Asterids

Ericales

Gilliesia Alliaceae Verbenaceae Gleicheniaceae Molluginaceae Poaceae Fabaceae Asteraceae Asteraceae Gomortegaceae Amaranthaceae Solanaceae Grammitidaceae Brassicaceae Gratiolaceae Bromeliaceae Asteraceae Griseliniaceae Sapindaceae Gunneraceae Asteraceae Asteraceae Poaceae Apiaceae Asteraceae Cactaceae Orchidaceae Haloragaceae Ranunculaceae Asteraceae Anacardiaceae Plantaginaceae Rubiaceae Asteraceae Poaceae Boraginaceae Asteraceae 260 8 50 32 150 80 1 Chile, Peru New Guinea, Aus, Tasm, NZ, temp S Am, Falkland tropics espec Asia & Malesia, JF W N Am, Mexico, C America, Andes temp N Hemis, montane trops, S Afr temp, subtrop, trop regs, arid Peru, Chile, Bol, Arg AND W-NT 75 Am espec Chile 5 Antarctic S Am W-NT AND SPT 27 Aus, N. Caled, NZ, Rapa, JF 600 pantrop & subtrop AUS 20 Peru, Chile 1 N Chile AND 6 Andes Chile Arg 1 Chile AND 1 Chile 27 W N Am & South S Am 40 Am from Mexic to Chile, trop & S Afr, Madag, Malesia, Tasm, NZ, Hawaii, JF ANT-P 3 Chile Arg AND 6 NZ, Chile, Paraguay, S Brasil AUS 2 3 1 4 6 6 3 6 3 1 2 5 3 3 2 1 3 7 7 3 60 W N Am & S Am, Brazil W-NT 3 27 C & S Am, JF W-NT 3 25 temp & subtrop, montane tropics 7 1 2 4 4 1 7 6 1 1 5 1 2 1 2 4 64 1 2 1 7 1 24 2 -50,62 -21,18 -18,75 -46,67 -34,77 -22,00 -36,83 -18,20 -17,58 -54,92 -45,50 -19,25 -56,00 -42,50 -40,33 -36,83 -38,63 -23,87 -52,77 -33,34 -19,00 -51,33 -38,63 -31,17 -36,83 -28,42 -20,73 1 Patagonia 5 1 400 New Guinea, Asia, Am, Aus, Madag, warm & temp, JF 1 3 4 Mexico, Galapagos, temp S Am W-NT 3 1 -25,40 -37,42 -51,00 -34,77 -35,00 -18,42 -24,93 -31,00 -29,83 -21,00 -29,75 -32,88 -23,50 -25,00 -18,33 -28,50 100 trop & subtrop Am, Afr, Indochina, Aus 1 2 -18,00 1 Chile 6 1 -35,62 -37,67 -22,50 -25,40 -55,87 -51,00 -42,50 -43,50 -38,78 -49,20 -35,97 -56,00 -52,50 -31,50 -40,00 -34,55 -26,13 -20,00 -52,00 66 Trop & Subtrop Am, S USA, SE Asia C-PAC 4 1 -30,67 -37,80 100 cosmop 7 30 -17,58 -54,50 20 Eurasia, Aus, N Am & temp S Am W-ANT 4 1 -34,08 -37,58 45 temp & subtrop, montane tropics 7 1 -33,25 -41,67 -37,46 -35,83 -36,04 -34,23 -36,64 -20,25 -25,40 -46,64 -51,00 -38,63 -39,25 -28,60 -37,07 -33,48 -42,92 -36,75 -30,63 -36,44 -29,03 -25,57 -19,17 -40,25 9 pantrop 1 1 -31,50 -37,17 -34,33 110 pantrop, Madag, JF 1 4 -37,23 -55,83 -46,53 18,60 5,67 8,42 3,50 36,92 7,13 2,05 4,50 0,00 18,45 0,00 7,73 8,50 20,37 24,27 4,97 26,17 31,50 1,75 7,12 11,05 1,13 1,67 23,50 0,00 4,30 24,32 0,50 9,33 7,73 18,33 0,00 20,43 6,28 100 disjunct N Am S Am ANT-P 4 6 -21,27 -38,00 -29,63 16,73 3 Chile, Arg 5 2 -32,37 -38,25 -35,31 5,88

Monocots

Asparagales

Glandularia

Asterids

Lamiales

Gleichenia

Filicopsida

Gleicheniales

Glinus

Eudicots

Caryophyllales

Glyceria

Monocots

Poales

Glycyrrhiza

Rosids

Fabales

Gnaphalium

Asterids

Asterales

Gochnatia

Asterids

Asterales

Gomortega

Magnoliids

Laurales

Gomphrena

Eudicots

Caryophyllales

Grabowskia

Asterids

Solanales

Grammitis

Filicopsida

Polypodiales

Grammosperma

Rosids

Brassicales

Gratiola

Asterids

Lamiales

Greigia

Monocots

Poales

Grindelia

Asterids

Asterales

Griselinia

Asterids

Apiales



Guindilia

Rosids

Sapindales

Gunnera

Eudicots

Gunnerales

Gutierrezia

Asterids

Asterales

Guynesomia

Asterids

Asterales

Gymnachne

Monocots

Poales

Gymnophyton

Asterids

Apiales

Gypothamnium

Asterids

Asterales

Haageocereus

Eudicots

Caryophyllales

Habenaria

Monocots

Asparagales

Haloragis

Eudicots

Saxifragales

Hamadryas

Eudicots

Ranunculales

Haplopappus

Asterids

Asterales

Haplorhus

Rosids

Sapindales

Hebe

Asterids

Lamiales

Hedyotis

Asterids

Gentianales

Helenium

Asterids

Asterales

Helictotrichon

Monocots

Poales

Heliotropium

Asterids

Unplaced

Helogyne

Asterids

Asterales

Herbertia Iridaceae Agavaceae Asteraceae Zosteraceae Asteraceae Plantaginaceae Dennstaedtiaceae Fabaceae Brassicaceae Apiaceae Poaceae Apiaceae Loasaceae Lycopodiaceae Violaceae Hydrangeaceae Apiaceae Hymenophyllaceae Hymenophyllaceae Hypericaceae Asteraceae Dennstaedtiaceae Campanulaceae Poaceae Convolvulaceae Polemoniaceae Isoetaceae Brassicaceae Solanaceae Poaceae Calceolariaceae Arecaceae Arecaceae Juncaceae 275 1 1 JF Chile Cosmop, JF, IP 5 NZ, Chile 59 23 1 N Chile temp S Am. S Peru, Bol, Bras, Parag, Chile, Arg, Uru Arg, Bol, Chile, Colombia, CosRica, Ecuador, El Salvador, Guatemala, Mexico, Peru, USA, Uruguay, Venezuela S-AMZ W-NT AUS 150 cosmop, Madag 26 W N Am & Florida, 1 Chile Arg 650 pantrop ANT-P 9 trop, subtrop & temp regions 5 E Aus, NZ, Andes S Am 40 trop & subtrop, Madag AUS 55 temp S Am, Eurasia Medit, N Afr 370 temp regs, montane tropics W-ANT 250 trop & S temp, Euro, Japan, Madag, JF 1 Chile, JF 130 cosmop 7 6 7 7 4 1 2 7 1 4 7 6 3 3 2 6 6 7 29 Himalayas, Asia, Philippines, N, C Am, Andes to Chile W-ANT 4 130 trops & subtrops 1 1 1 8 1 19 2 33 1 1 4 1 1 2 1 9 22 3 1 1 28 -30,50 -17,58 -35,50 -56,00 -33,00 -36,79 -21,00 -37,80 -27,40 -18,18 -17,67 -35,75 -26,93 -53,53 -27,40 -53,83 -53,50 -40,00 -23,97 -45,67 -27,40 -36,01 -35,58 -37,88 350 Subcosmop, Madag 7 1 2 Chile 6 2 4 Patagonia 5 4 -36,60 -19,00 -53,67 -36,50 -33,50 -18,83 -37,60 -30,67 -34,77 -17,58 -30,67 -18,00 -30,60 30 temp & subtrop regs, montane tropics W-ANT 4 11 -18,18 6 Chile 6 6 -25,20 1 Chile 6 1 -38,67 -41,00 -33,75 -55,00 -52,95 -29,50 -54,67 -37,08 -46,67 -51,27 -49,15 -56,00 -42,50 -54,00 -49,15 -54,00 -40,33 28 25 SW US, Chile, Arg ANT-P 4 7 -18,00 -33,45 5 pantrop, Madag 1 1 -42,05 -54,07 1 circumboreal, S Am, Australia W-ANT 4 1 -37,00 -55,00 -46,00 -48,06 -25,73 -39,83 -29,48 -36,59 -44,78 -24,25 -54,17 -36,79 -40,08 -35,05 -43,38 -43,33 -38,63 -35,79 -39,91 -36,00 -35,47 200 temp regions & montane tropics, excl Australasia W-ANT 4 6 -34,65 -55,00 -44,83 1 coastal temp Aus, Tasm, N Chile AUS 2 1 -30,20 -30,30 -30,25 5 SW USA, Mexico, C Am, Peru, Arg, Chile W-NT 3 2 -17,58 -23,00 -20,29 5,42 0,10 20,35 18,00 12,02 15,45 2,33 8,55 36,82 16,35 10,50 1,00 0,58 13,17 32,43 11,55 25,33 7,73 36,42 18,48 36,00 9,73 0,00 5,93 15,73 0,00 35,65 35,83 4,25 0,00 5,00 38,42 8 Brasil, Uru, Parag, Arg, Chile S-AMZ 3 1 -35,33 -37,00 -36,17 1,67 5 temp S Am, Chile, Brazil S-AMZ 3 1 -33,00 -40,00 -36,50 7,00

Monocots

Asparagales

Herreria

Monocots

Asparagales

Heterosperma

Asterids

Asterales

Heterozostera

Monocots

Alismatales

Hieracium

Asterids

Asterales

Hippuris

Asterids

Lamiales

Histiopteris

Filicopsida

Dennstaedtiales

Hoffmannseggia

Rosids

Fabales

Hollermayera

Rosids

Brassicales

Homalocarpus

Asterids

Apiales

Hordeum

Monocots

Poales

Huanaca

Asterids

Apiales

Huidobria

Asterids

Cornales

Huperzia

Lycopsida

Lycopodiales

Hybanthus

Rosids

Malpighiales

Hydrangea

Asterids

Cornales

Hydrocotyle

Asterids

Apiales



Hymenoglossum

Filicopsida

Hymenophyllales

Hymenophyllum

Filicopsida

Hymenophyllales

Hypericum

Rosids

Malpighiales

Hypochaeris

Asterids

Asterales

Hypolepis

Filicopsida

Dennstaedtiales

Hypsela

Asterids

Asterales

Imperata

Monocots

Poales

Ipomoea

Asterids

Solanales

Ipomopsis

Asterids

Ericales

Isoetes

Lycopsida

Isoetales

Ivania

Rosids

Brassicales

Jaborosa

Asterids

Solanales

Jarava

Monocots

Poales

Jovellana

Asterids

Lamiales

Juania

Monocots

Arecales

Jubaea

Monocots

Arecales

Juncus

Monocots

Poales

Junellia Verbenaceae Rosaceae Poaceae Krameriaceae Lamiaceae Cyperaceae Lactoridaceae Asteraceae Verbenaceae Lemnaceae Philesiaceae Lardizabalaceae Apiaceae Zygophyllaceae Polygonaceae Asteraceae Fabaceae Solanaceae Atherospermataceae Atherospermataceae Ericaceae Malvaceae Campanulaceae Myrtaceae Lemnaceae Portulacaceae Alstroemeriaceae Lamiaceae Brassicaceae Santalaceae Asteraceae Podocarpaceae Asteraceae Asteraceae 1 33 3 1 2 180 Cosmop, ID S Peru, Chile Patagonia Chile-Arg. 2 NZ, 1 S Chile/Arg New Guinea, Aus, NZ, Subantarctic Is, S Am Chile AUS SPT AND 40 1 Chile California, Andes, Mexico to S Brasil, Arg, Chile W-NT 1 Chile Arg 14 Cosmop AND 1 Chile 2 1 Calif, 1 Chile 7 Arg, Chile ANT-P 1 Patagonia 1 Chile Arg 2 NZ & Chile 1 Chile AUS 155 temp N hemis, Medit, montane tropics, E Afr, S Am W-ANT 18 SW US, Chile-Arg ANT-P 4 4 6 2 5 5 5 4 6 7 3 6 3 7 3 5 2 2 6 3 California, Chile ANT-P 4 5 SW N Am, S Am ANT-P 4 3 1 1 11 1 1 1 1 1 1 1 3 1 1 3 22 1 1 1 1 1 1 Chile, Argentina AND 3 1 1 Chile and adjacent Argentina 5 1 1 Chile 6 1 -30,67 -32,50 -30,17 -28,80 -27,83 -31,25 -29,47 -40,17 -34,73 -38,37 -43,42 -52,10 -40,00 -35,27 -18,00 -23,00 -27,90 -29,83 -18,00 -36,33 -52,33 -40,17 -40,00 -35,78 1 cosmop 7 1 -55,50 3 Bol, Chile, Arg AND 3 1 -18,17 -23,50 -55,50 -40,83 -39,87 -35,50 -36,75 -35,97 -51,00 -53,00 -43,00 -41,17 -46,67 -56,00 -52,10 -40,00 -36,68 -46,50 -30,25 -28,08 -40,00 -55,00 -43,00 -53,00 -55,00 -56,00 -40,82 15 SE Asia, Malesia, Australia, Tasmania, NZ, Central and South America, JF, Falkland, Tristan da Cunha. SPT 2 3 -33,67 -56,00 1 JF 6 1 -44,83 -20,83 -55,50 -35,75 -36,18 -32,83 -32,78 -31,90 -41,13 -41,23 -41,58 -37,95 -42,52 -49,71 -52,10 -40,00 -35,98 -32,25 -26,63 -27,99 -34,92 -36,50 -39,67 -52,67 -47,58 -48,00 -38,30 40 pantrops 1 1 1 Chile / Arg AND 3 1 -28,72 -30,50 -29,61 18 SW US, Mexico, Ecuador, Chile, Arg, Brazil W-NT 3 2 -17,58 -33,50 -25,54 15,92 1,78 0,00 0,00 22,33 5,33 0,00 10,17 7,37 5,33 7,95 8,13 19,75 23,53 2,83 6,43 8,30 12,58 0,00 0,00 1,42 28,50 7,25 0,18 10,17 37,00 6,67 0,67 14,83 16,00 5,03 45 temp & subtrops, montane trops 7 2 -31,50 -54,83 -43,17 23,33 4 Chile, Peru, Bolivia, Arg AND 3 2 -30,33 -38,67 -34,50 8,33 47 Peru, Bol, Chile, Arg AND 3 31 -18,00 -52,50 -35,25 34,50

Asterids

Lamiales

Kageneckia

Rosids

Rosales

Koeleria

Monocots

Poales

Krameria

Rosids

Zygophyllales

Kurzamra

Asterids

Lamiales

Kyllinga

Monocots

Poales

Lactoris

Magnoliids

Piperales

Lagenophora

Asterids

Asterales

Lampaya

Asterids

Lamiales

Landoltia

Monocots

Alismatales

Lapageria

Monocots

Liliales

Lardizabala

Eudicots

Ranunculales

Laretia

Asterids

Apiales

Larrea

Rosids

Zygophyllales

Lastarriaea

Eudicots

Caryophyllales

Lasthenia

Asterids

Asterales

Lathyrus

Rosids

Fabales



Latua

Asterids

Solanales

Laurelia

Magnoliids

Laurales

Laureliopsis

Magnoliids

Laurales

Lebetanthus

Asterids

Ericales

Lecanophora

Rosids

Malvales

Legenere

Asterids

Asterales

Legrandia

Rosids

Myrtales

Lemna

Monocots

Alismatales

Lenzia

Eudicots

Caryophyllales

Leontochir

Monocots

Liliales

Lepechinia

Asterids

Lamiales

Lepidium

Rosids

Brassicales

Lepidoceras

Eudicots

Santalales

Lepidophyllum

Asterids

Asterales

Lepidothamnus

Pinopsida

Pinales

Leptinella

Asterids

Asterales

Leptocarpha

Asterids

Asterales

Leptochloa Poaceae Poaceae Rubiaceae Saxifragaceae Asteraceae Alliaceae Asteraceae Iridaceae Loranthaceae Juncaginaceae Apiaceae Plumbaginaceae Gratiolaceae Polemoniaceae Linderniaceae Linaceae Verbenaceae Anacardiaceae Plantaginaceae Sapindaceae Loasaceae Campanulaceae Proteaceae Asteraceae Lophosoriaceae Fabaceae Asteraceae Asteraceae Onagraceae Myrtaceae Fabaceae Juncaceae Luzuriagaceae Asteraceae Solanaceae Solanaceae Lycopodiaceae 10 55 80 1 3 75 cosmop, JF 1 NZ, 3 Chile, Arg to T Fuego Desventuradas Islands (Chile) N Am, tem S Am, Eurasia, S Afr, Aus, IP S Am Colombia to Chile, Brazil trop & temp AND AUS 220 4 Chile Arg N & S Am, Medit, montane trops, E Afr W-ANT 81 Cosmop 12 Andes S Am, Chile, Arg, Peru 12 Andes S Am, Chile, Arg, Peru 100 1 trop Am Macaronesia, temp eurasia, Afr, Aus, S Am AND AND 6 Peru, Bol, Chile, Arg 12 Aus, Tas, S Am 330 trops & subtrops AUS AND W-NT 36 Chile, Arg, 1 sp coastal Peru 3 Andean trop S Am 3 1 N Am, 1 S Am, 1 Euro AND AND 4 S Am: Arg, Bol, Paraguay, SE Brazil, Chile. S-AMZ W-ANT 200 Am trop, many in Afr trop 1 3 4 3 3 1 2 3 3 7 3 3 7 5 4 7 2 6 7 3 7 200 temp & subtrops 7 100 trops, subtrops, temp, Madag 7 1 5 2 1 1 1 36 8 3 2 1 1 3 2 4 2 6 6 3 1 9 1 5 -18,00 -18,27 -37,00 -35,33 -28,88 -56,00 -26,67 -23,58 -46,50 35 W N Am, Chile ANT-P 4 1 15 Cosmop, Madag 7 1 250 N temperate, circumboreal, austral W-ANT 4 2 -27,07 -18,00 -31,00 -38,60 -23,00 -23,50 -30,00 -39,67 -29,73 -21,03 -25,22 -30,28 -18,00 -35,22 -24,83 -33,00 -17,58 -28,50 -29,50 -18,00 -30,00 -34,77 13 Am Canada to TFuego, Aus, NZ C-PAC 4 1 -18,25 1 Pacific Am Canada to Chile ANT-P 4 1 -33,22 2 C Peru, E Bol, C Chile, Urug, n Arg, S Brasil S-AMZ 3 1 -30,50 -37,00 -42,05 -53,67 -32,55 -55,00 -37,67 -38,60 -37,58 -30,50 -38,25 -55,00 -34,13 -50,00 -43,83 -50,00 -20,08 -46,42 -40,00 -38,67 -23,50 -40,60 -45,42 -40,00 -56,00 -56,00 10 New Guinea, E Aus, NZ, Andes S Am, JF SPT 2 4 -32,23 -49,15 1 C Chile 6 1 -30,80 -32,75 11 Chile 6 11 -20,83 -40,00 -30,42 -31,78 -40,69 -33,75 -37,63 -35,96 -29,81 -36,50 -34,33 -38,60 -30,29 -27,00 -34,13 -47,33 -31,93 -35,52 -34,53 -40,14 -19,04 -40,82 -32,42 -35,83 -20,54 -34,55 -37,46 -29,00 -43,00 -45,38 46 Peru, Bol, Arg, Chile, Falkland AND 3 43 -17,58 -55,00 -36,29 1 SE USA, Mexico, Chile, Arg Uruguay ANT-P 4 1 -32,57 -40,00 -36,28 7,43 37,42 19,17 1,95 16,92 6,50 8,83 35,42 5,48 37,00 6,67 0,00 14,58 7,00 8,25 15,33 4,40 28,97 18,62 19,72 2,08 11,20 15,17 5,67 5,92 12,10 15,92 22,00 26,00 21,23 0,00 17,33 10,62 19,00 6 SE Asia, NZ, W S Am SPT 2 1 -36,75 -42,47 -39,61 5,72 1 Arg Chile, JF 5 1 -38,00 -40,00 -39,00 2,00 17 tropics & subtrop 1 3 -18,42 -24,00 -21,21 5,58

Monocots

Poales

Leptophyllochloa

Monocots

Poales

Leptostigma

Asterids

Gentianales

Lepuropetalon

Eudicots

Saxifragales

Leucheria

Asterids

Asterales

Leucocoryne

Monocots

Asparagales

Leunisia

Asterids

Asterales

Libertia

Monocots

Asparagales

Ligaria

Eudicots

Santalales

Lilaea

Monocots

Alismatales

Lilaeopsis

Asterids

Apiales

Limonium

Eudicots

Caryophyllales

Limosella

Asterids

Lamiales

Linanthus

Asterids

Ericales

Lindernia

Asterids

Lamiales

Linum

Rosids

Malpighiales

Lippia

Asterids

Lamiales

Lithrea

Rosids

Sapindales

4

Littorella

Asterids

Lamiales

Llagunoa

Rosids

Sapindales

Loasa

Asterids

Cornales

Lobelia

Asterids

Asterales

Lomatia

Eudicots

Proteales

Lophopappus

Asterids

Asterales

Lophosoria

Filicopsida

Dicksoniales

Lotus

Rosids

Fabales

Lucilia

Asterids

Asterales

Luciliocline

Asterids

Asterales

Ludwigia

Rosids

Myrtales

Luma

Rosids

Myrtales

Lupinus

Rosids

Fabales

Luzula

Monocots

Poales

Luzuriaga

Monocots

Liliales

Lycapsus

Asterids

Asterales

Lycium

Asterids

Solanales

Lycopersicon

Asterids

Solanales

Lycopodium

Lycopsida

Lycopodiales

Lysimachia Myrsinaceae Cyperaceae Asteraceae Asteraceae Tropaeolaceae Cactaceae Cactaceae Amaranthaceae Asteraceae Malesherbiaceae Malvaceae Brassicaceae Rosaceae Marsileaceae Juncaceae Asteraceae Iridaceae Brassicaceae Celastraceae Gratiolaceae Poaceae Dryopteridaceae Poaceae Plantaginaceae Oleaceae Brassicaceae Loasaceae Zygophyllaceae Dennstaedtiaceae Caryophyllaceae Asteraceae Asteraceae Polypodiaceae Polemoniaceae Alliaceae Asteraceae 1 430 1 40 15 8 3 Chile Brazil, Arg, Urug, Chile W N Am, 1 Chile, 1 Aus, NZ trop & warm, Madag W N Am & S Am Chile pantrops ANT-P S-AMZ C-PAC 45 1 N Chile paleotrop, 1 Afr, Madag, Japan, NZ 80 Chile, Arg to Canada, WI 26 Chile, Arg W-NT 25 trop & subtrop Am, S Afr 2 Chile, Arg 80 temp regs & montane trops 40 Trop Am, 1 Afr to Mascarenes (no Madag), JF W-ANT 2 JF 10 Am S US to Arg 200 trops & subtrops W-NT 7 Peru, Chile AND 16 Peru, Arg AND 3 3 1 3 6 1 4 5 4 5 3 6 1 6 3 4 1 4 6 1 1 Chile 6 3 NZ, S S Am, Tfuego, Falkland AUS 2 3 1 1 6 4 1 2 1 7 2 1 19 7 1 1 3 1 1 1 1 1 1 -18,00 -31,00 -34,50 -30,50 -35,33 -37,92 -24,25 -33,17 -36,21 -24,67 -31,33 -25,00 -37,50 -36,50 -37,83 -31,08 -33,92 -31,42 -30,67 -25,10 -31,00 -32,05 -22,08 -22,08 -20,08 -46,68 -39,88 -36,50 -32,63 -51,17 -33,33 -20,63 -38,68 -32,49 -33,75 -32,34 -36,63 -27,71 -20,36 70 trop & temp, Madag 7 1 3 S Andes to Chile, JF, Brazil S-AMZ 3 1 7 Mexico, Andes Peru, Bol, Arg, Chile ANT-P 4 2 4 3 sp W USA, Mexico, 1 sp Peru to Uruguay, 1 sp. Medit W-ANT 4 1 -25,00 -18,00 -25,00 -30,63 -37,00 -32,72 -18,17 -22,00 -28,50 -35,33 24 Andes Peru, Chile, Arg AND 3 18 -18,50 16 W N Am, Chile ANT-P 4 2 -22,00 -30,67 -35,00 -33,68 -25,00 -43,67 -32,15 -56,00 -33,60 -23,00 -42,00 -55,00 -35,47 58 Australia, Chile AUS 2 1 -27,68 -27,68 18 Andes Peru, Bol, Chile, Arg AND 3 11 -19,42 -30,00 2 Chile Arg 5 1 -35,55 -37,50 -36,53 -24,71 -27,68 -26,33 -26,75 -29,34 -21,50 -34,33 -31,39 -46,50 -33,16 -20,58 -32,00 -41,75 -35,40 2 Chile, Arg 5 1 -46,53 -46,53 -46,53 10 W N Am, Chile-Arg ANT-P 4 2 -30,00 -55,00 -42,50 0,00 1,95 10,58 0,00 8,67 16,50 8,68 7,00 18,67 1,52 19,00 0,88 4,83 20,00 26,50 0,13 0,00 16,02 14,78 5,50 0,58 29,08 11,25 0,55 0,00 12,83 5,17 12,83 0,00 12,50 4,33 3,42 1 Patagonia & Tfuego 5 1 -38,27 -55,00 -46,63 16,73 25,00 45 trops esp Australia, Hawaii, JF 1 1 0,00 150 Cosmop 7 1 -36,33 -41,67 -39,00 5,33

Asterids

Ericales

Machaerina

Monocots

Poales

Macrachaenium

Asterids

Asterales

Madia

Asterids

Asterales

Magallana

Rosids

Brassicales

Maihuenia

Eudicots

Caryophyllales

Maihueniopsis

Eudicots

Caryophyllales

Maireana

Eudicots

Caryophyllales

Malacothrix

Asterids

Asterales

Malesherbia

Rosids

Malpighiales

Malvella

Rosids

Malvales

Mancoa

Rosids

Brassicales

Margyricarpus

Rosids

Rosales

Marsilea

Filicopsida

Marsileales

Marsippospermum

Monocots

Poales

Marticorenia

Asterids

Asterales

Mastigostyla

Monocots

Asparagales



Mathewsia

Rosids

Brassicales

Maytenus

Eudicots

Celastrales

Mecardonia

Asterids

Lamiales

Megalachne

Monocots

Poales

Megalastrum

Filicopsida

Dryopteridales

Melica

Monocots

Poales

Melosperma

Asterids

Lamiales

Menodora

Asterids

Lamiales

Menonvillea

Rosids

Brassicales

Mentzelia

Asterids

Cornales

Metharme

Rosids

Zygophyllales

Microlepia

Filicopsida

Dennstaedtiales

Microphyes

Eudicots

Caryophyllales

Micropsis

Asterids

Asterales

Microseris

Asterids

Asterales

Microsorum

Filicopsida

Polypodiales

Microsteris

Asterids

Ericales

Miersia

Monocots

Asparagales

Mikania

Asterids

Asterales

Mimulus Phrymaceae Caryophyllaceae Cactaceae Nyctaginaceae Misodendraceae Gesneriaceae Asteraceae Portulacaceae Portulacaceae Plantaginaceae Myricaceae Asteraceae Calyceraceae Polygonaceae Poaceae Apiaceae Poaceae Asteraceae Santalaceae Boraginaceae Ranunculaceae Myrtaceae Myrtaceae Haloragaceae Myrtaceae Boraginaceae Santalaceae Asteraceae Loasaceae Asteraceae Poaceae Calyceraceae Polemoniaceae Cactaceae Rubiaceae 30 2 13 9 79 39 100 7 Bol, Arg, Chile Colombia, Ecuador, Peru, few in Chile, Bolivia, Venezuela and C Am (to S Mexico). Bol, Arg, Chile C & S Am Chile, Arg, Falkland Is. W N Am, Chile, Arg Peru, Bol, Chile, Arg Madag, SE Asia, Malesia, Aus, NZ, Hawaii, S Am, JF ANT-P AND C-AUS 1 temp S Am AND W-NT AND W-NT 45 3 Colombia to Chile, JF, Brazil N Am from California,Mex, C Am to Brasil, WI, Hawaii 60 subcosmop AND W-NT 50 trop Am, Andes, Brazil, WI 40 Chile, Arg, SE Brasil, JF 15 temp regs hemis N & S, M apetalus disj N Am 100 Eurasia, montane trops, S Afr, N Guinea, Aus, NZ 1 Chile, Arg W-ANT W-ANT S-AMZ AND 60 Andes Colombia to Chile, Arg, Parag, SE Brasil, Urug 5 1 W US, 4 SW S Am AND 20 S Andes ANT-P 150 temp & subtrop Am, Asia W-ANT 4 5 4 3 5 4 4 3 3 7 3 3 5 3 3 3 3 5 4 3 2 25 New Guinea, Aus, Tasm, NZ, W S Am SPT 2 8 Per, Chile, Arg, Patagonia AND 3 4 1 3 7 2 23 1 2 2 12 1 2 1 2 1 5 1 25 26 5 1 1 1 2 Chile 6 2 40 Sudam Andes Peru Ecuador, Africa, E Asia, Filipinas y Malasia 1 1 3 Chile 1, Arg. 2 AND 3 1 -25,00 -18,83 -29,72 -17,67 -19,65 -19,50 -20,98 -18,18 -17,58 -31,62 -48,47 -29,80 -30,67 -29,43 -19,50 -36,23 -21,50 -40,33 -23,00 -20,00 -30,00 -17,67 -25,00 -33,25 -18,50 -30,67 18 Chile, Argentina, Bolivia y Per AND 3 18 -19,25 20 temp Eurasia, Medit, montane tropics, Afr, A Aus, A & S Am, 7 1 -29,50 -55,00 -40,75 -31,67 -19,12 -36,50 -55,00 -45,50 -41,00 -45,48 -24,00 -48,92 -55,00 -54,88 -53,38 -46,43 -30,00 -55,00 -56,00 -25,50 -56,00 -53,00 -25,00 -55,67 -50,00 -46,00 -40,53 -18,50 -56,00 4 Peru, Chile AND 3 1 -17,58 -18,17 1 Chile, Arg. 5 1 -30,67 -49,88 8 S Am, south of 33 5 8 -33,00 -56,00 -44,50 -40,28 -17,88 -42,25 -30,00 -28,33 -18,98 -33,11 -36,33 -32,58 -30,25 -33,23 -21,09 -33,25 -43,31 -51,68 -41,59 -38,55 -29,72 -37,25 -46,12 -23,50 -48,17 -38,00 -22,50 -42,83 -33,83 -35,50 -36,89 -18,50 -43,33 54 Am trop, Himal, China W-ANT 4 4 -18,33 -36,00 -27,17 1 Chile 6 1 -27,50 -30,00 -28,75 2,50 17,67 23,00 19,22 0,58 25,50 21,50 6,67 0,28 6,78 37,33 25,85 21,50 24,50 5,82 31,33 23,38 6,42 23,58 15,77 0,57 35,50 19,77 4,00 15,67 30,00 5,00 25,67 32,33 21,00 7,28 0,00 25,33 120 Arctic & temp regions, Mexico, Ethiopia, Himalayas, Chile W-ANT 4 1 -33,00 -36,75 -34,88 3,75 150 Am, S Afr, Madag, E Asia, JF 7 8 -22,00 -45,50 -33,75 23,50

Asterids

Lamiales

Minuartia

Eudicots

Caryophyllales

Miqueliopuntia

Eudicots

Caryophyllales

Mirabilis

Eudicots

Caryophyllales

Misodendrum

Eudicots

Santalales

Mitraria

Asterids

Lamiales

Mniodes

Asterids

Asterales

Montia

Eudicots

Caryophyllales

Montiopsis

Eudicots

Caryophyllales

Monttea

Asterids

Lamiales

Morella

Rosids

Fagales

Moscharia

Asterids

Asterales

Moschopsis

Asterids

Asterales

Muehlenbeckia

Eudicots

Caryophyllales

Muhlenbergia

Monocots

Poales



Mulinum

Asterids

Apiales

Munroa

Monocots

Poales

Mutisia

Asterids

Asterales

Myoschilos

Eudicots

Santalales

Myosotis

Asterids

Unplaced

Myosurus

Eudicots

Ranunculales

Myrceugenia

Rosids

Myrtales

Myrcianthes

Rosids

Myrtales

Myriophyllum

Eudicots

Saxifragales

Myrteola

Rosids

Myrtales

Nama

Asterids

Unplaced

Nanodea

Eudicots

Santalales

Nardophyllum

Asterids

Asterales

Nasa

Asterids

Cornales

Nassauvia

Asterids

Asterales

Nassella

Monocots

Poales

Nastanthus

Asterids

Asterales

Navarretia

Asterids

Ericales

Neowerdermannia

Eudicots

Caryophyllales

Nertera

Asterids

Gentianales

Nesocaryum Boraginaceae Brassicaceae Solanaceae Solanaceae Amaranthaceae Brassicaceae Solanaceae Loranthaceae Nothofagaceae Pteridaceae Alliaceae Asteraceae Malvaceae Bromeliaceae Onagraceae Iridaceae Balanophoraceae Brassicaceae Ophiglossaceae Asteraceae Cyperaceae Cactaceae Apiaceae Rubiaceae Proteaceae Orobanchaceae Poaceae Apiaceae Fabaceae Plantaginaceae Thymelaeaceae Oxalidaceae Juncaceae Asteraceae Polygonaceae 1 7 6 600 2 25 temp S Am Cosmop Andes S Am N Chile W N Am, Chile, Arg ANT-P AND 36 10 2 Arg, Chile N Am, E Asia, temp S Am E & S Afr, temp S Am Tasm, NZ, Andes S Am AUS W-ANT 130 temp, subtrop, N Hemis 9 6 temp E Aus, 3 Andes S Am 1 Chile, Arg, Patagonia AUS 23 7 Andes Peru, Bol, Chile, Arg E Asia, Borneo, N Guinea, SE Aus, NZ, Mex, C Am, Andes, Tfuego, Falkland 15 Malesia, Aus, Tasm, NZ, Hawaii, JF, trop & temp Am, Chile, Tfuego, Falkland 37 trop & subtrop S Am, Brazil 26 Subcosmop, Madag, JF S-AMZ SPT AND SPT 5 Chile, Patagonia 4 Peru, W Brasil, N Arg, Chile AND 3 5 7 3 2 3 2 5 2 7 5 4 1 2 5 7 3 6 4 20 Andean and temp S Am and NW N Am W-NT 3 121 temp & subtrop N C & S Am W-NT 3 14 16 1 5 3 5 1 3 1 1 1 2 2 3 2 13 2 117 1 1 1 4 Chile, JF 6 3 100 Ecuador, Peru, Bol, Chile, Arg AND 3 24 20 Peru, Bol, Chile, Arg, Ecuador, Brazil S-AMZ 3 5 -31,92 -17,72 -31,67 -18,00 -23,50 -18,20 -37,58 -33,10 -17,58 -36,50 -18,17 -52,00 -33,00 -35,83 -18,00 -39,48 -30,83 -29,83 -18,23 -37,80 -18,00 -18,00 -24,83 -24,00 26 Am W-NT 3 6 -29,90 40 warm to trop Am, SW US, Mex, Madag, JF 1 2 -17,58 37 New Guinea, Aus, Tasm, NZ, New Caled, temp S Am SPT 2 10 -32,95 -55,97 -27,28 -40,25 -38,73 -34,67 -38,25 -52,00 -55,00 -22,12 -54,67 -54,83 -36,67 -56,00 -22,50 -54,00 -54,50 -38,25 -33,50 -56,00 -54,93 -40,00 -56,00 -45,93 -56,00 -34,25 -26,15 -31,50 1 Chile temperate, JF 6 1 -32,57 -43,00 77 Peru to Chile, Galapagos AND 3 36 -18,25 -40,17 20 N temp, few S Am W-ANT 4 1 -33,50 -54,83 -44,17 -29,21 -37,78 -44,46 -22,43 -35,08 -35,33 -26,19 -34,96 -35,00 -39,25 -20,16 -46,13 -43,97 -27,13 -46,25 -20,33 -53,00 -43,75 -37,04 -25,75 -47,74 -42,88 -34,92 -37,12 -41,87 -37,00 -26,13 -25,49 -27,75 8 US, Mex, Chile, Arg ANT-P 4 1 -22,50 -24,50 -23,50 20 Mex to Arg, Bras W-NT 3 1 -34,83 -51,00 -42,92 16,17 2,00 21,33 21,92 10,43 23,02 9,70 10,35 6,82 16,95 6,58 34,00 31,50 3,92 17,08 21,73 19,08 19,50 4,33 2,00 21,50 2,42 15,50 16,52 24,10 10,17 37,77 8,13 38,00 16,25 1,32 7,50 67 Am, USA to Chile, Aust, New Caledonia, Namibia, JF 1 10 -18,47 -43,62 -31,04 25,15 2 Chile 6 2 -20,13 -20,13 -20,13 0,00 1 Desventuradas Islands (Chile) 6 1 0,00

Asterids

Unplaced

Neuontobothrys

Rosids

Brassicales

Nicotiana

Asterids

Solanales

Nierembergia

Asterids

Solanales

Nitrophila

Eudicots

Caryophyllales

Noccaea

Rosids

Brassicales

Nolana

Asterids

Solanales

Notanthera

Eudicots

Santalales

Nothofagus

Rosids

Fagales

Notholaena

Filicopsida

Pteridales

Nothoscordum

Monocots

Asparagales

Noticastrum

Asterids

Asterales

Nototriche

Rosids

Malvales

Ochagavia

Monocots

Poales

Oenothera

Rosids

Myrtales

Olsynium

Monocots

Asparagales

Ombrophytum

Eudicots

Santalales

7

Onuris

Rosids

Brassicales

Ophioglossum

Filicopsida

Ophioglossales

Ophryosporus

Asterids

Asterales

Oreobolus

Monocots

Poales

Oreocereus

Eudicots

Caryophyllales

Oreomyrrhis

Asterids

Apiales

Oreopolus

Asterids

Gentianales

Orites

Eudicots

Proteales

Orobanche

Asterids

Lamiales

Ortachne

Monocots

Poales

Osmorhiza

Asterids

Apiales

Otholobium

Rosids

Fabales

Ourisia

Asterids

Lamiales

Ovidia

Rosids

Malvales

Oxalis

Rosids

Oxalidales

Oxychlo

Monocots

Poales

Oxyphyllum

Asterids

Asterales

Oxytheca

Eudicots

Caryophyllales

Oziroe Hyacinthaceae Asteraceae Malvaceae Poaceae Asteraceae Urticaceae Caryophyllaceae Hemerocallidaceae Poaceae Passifloraceae Juncaceae Boraginaceae Pteridaceae Myrsinaceae Poaceae Piperaceae Asteraceae Asteraceae Lauraceae Monimiaceae Boraginaceae Poaceae Philesiaceae Apocynaceae Caryophyllaceae Poaceae Poaceae Solanaceae Amaryllidaceae Stylidiaceae Cyperaceae Phytolaccaceae Asteraceae Urticaceae Cupressaceae Rafflesiaceae 1 25 250 2 25 2 Arg, Chile trop & warm subtrop S Am, Chile, Peru, Arg, Brazil trop & warm excl Australasia Chile Arg Am US to Magallanes, Asia, 2 W Aus S-AMZ 4 3 Chile, Arg Tasm, NZ, temp S Am 1 4 Cosmop Chile (Atacama & Coquimbo) AND AUS 15 temperate Eurasia, America 1 Patagonia Chile-Arg. W-ANT 40 Andes Per, Chile, Bolivia, Argentina 1 Chile, Arg AND 20 temperate Eurasia, America 200 W N Am, S Am 1 Chile ANT-P W-ANT 120 trops & subtrops Am & Asia, Macaronesia 63 SW N Am, 1 Chile Peru ANT-P W-ANT 32 Andes from Colombia to Chile, SE Brazil AND 3 4 4 6 4 4 5 3 5 4 7 6 3 2 5 1 3 1 5 1 1000 trop & warm esp Am, JF, Hawaii 1 115 trop & temp regs W-ANT 4 1 4 23 1 2 1 6 2 1 1 1 1 1 1 3 1 2 1 1 2 1 1 2 1 subtrop S Am, 1 Macaronesia 1 1 35 trop & warm-temp, Madag 1 2 13 W N Am, S Am ANT-P 4 4 1 Andes Chile Arg Bol AND 3 1 -24,00 -23,58 -18,27 -32,50 -20,00 -24,83 -17,58 -20,80 -33,08 -30,67 -18,00 -30,83 -39,83 -18,33 -48,00 -29,50 -19,67 -26,43 -29,67 -45,00 -22,00 -37,13 -34,50 -37,62 -39,90 -18,18 500 trop & subtrop Am, 1 Madag, 20 trop E Asia, Malesia, 3 Aus, NZ 1 1 -31,97 330 temp & trop regs, IP 7 5 -20,50 -42,50 -32,50 -38,00 -35,97 -40,32 -40,00 -33,53 -42,50 -56,00 -29,90 -43,18 -40,50 -54,83 -39,88 -55,50 -22,27 -50,00 -55,17 -40,00 -31,33 -36,62 -56,00 -51,00 -41,88 -36,72 -45,42 -54,25 -36,90 1 Peru, Chile AND 3 1 -22,00 -40,00 100 North temperate, austral W-ANT 4 5 -18,18 -37,80 -27,99 -31,00 -31,50 -32,23 -31,00 -29,78 -29,29 -36,25 -26,77 -33,67 -36,79 -25,35 -38,13 -35,58 -36,42 -35,36 -47,67 -20,30 -49,00 -42,33 -29,83 -28,88 -33,14 -50,50 -36,50 -39,51 -35,61 -41,52 -47,08 -27,54 15 subcosmop, JF, ID 7 2 -20,80 -35,62 -28,21 5 Andes Bol, Chile, Per, Arg AND 3 5 -17,58 -25,00 -21,29 600 temp, subtrop, trop regs 7 2 -33,50 -37,67 -35,58 4,17 7,42 14,82 19,62 18,00 22,00 0,53 14,00 12,38 22,05 7,50 13,53 17,67 38,42 9,10 10,10 9,83 36,83 9,05 15,67 3,93 2,00 25,67 20,33 4,90 6,95 11,00 29,00 4,75 2,22 7,80 14,35 18,72 15 Coastal Peru and Chile AND 3 4 -20,38 -25,20 -22,79 4,82 2 Andes N Chile Arg AND 3 2 -28,00 -35,80 -31,90 7,80 5 Chile, Arg, Per, Bol, Paraguay AND 3 4 -18,40 -36,00 -27,20 17,60

Monocots

Asparagales

Pachylaena

Asterids

Asterales

Palaua

Rosids

Malvales

Panicum

Monocots

Poales

Parastrephia

Asterids

Asterales

Parietaria

Rosids

Rosales

Paronychia

Eudicots

Caryophyllales

Pasithea

Monocots

Asparagales

Paspalum

Monocots

Poales

Passiflora

Rosids

Malpighiales

Patosia

Monocots

Poales

Pectocarya

Asterids

Unplaced

Pellaea

Filicopsida

Pteridales

Pelletiera

Asterids

Ericales

Pennisetum

Monocots

Poales

Peperomia

Magnoliids

Piperales

8

Perezia

Asterids

Asterales

Perityle

Asterids

Asterales

Persea

Magnoliids

Laurales

Peumus

Magnoliids

Laurales

Phacelia

Asterids

Unplaced

Phalaris

Monocots

Poales

Philesia

Monocots

Liliales

Philibertia

Asterids

Gentianales

Philippiella

Eudicots

Caryophyllales

Phleum

Monocots

Poales

Phragmites

Monocots

Poales

Phrodus

Asterids

Solanales

Phycella

Monocots

Asparagales

Phyllachne

Asterids

Asterales

Phylloscirpus

Monocots

Poales

Phytolacca

Eudicots

Caryophyllales

Picrosia

Asterids

Asterales

Pilea

Rosids

Rosales

Pilgerodendron

Pinopsida

Pinales

Pilostyles

Rosids

Malpighiales

Pilularia Marsileaceae Lentibulariaceae Zygophyllaceae Poaceae Rutaceae Verbenaceae Pteridaceae Amaryllidaceae Boraginaceae Plantaginaceae Asteraceae Valerianaceae Asteraceae Polypodiaceae Lythraceae Aspleniaceae Asteraceae Plumbaginaceae Poaceae Poaceae Asteraceae Podocarpaceae Poaceae Polemoniaceae Asteraceae Caryophyllaceae Polygalaceae Polygonaceae Rosaceae Polypodiaceae Poaceae Dryopteridaceae Zygophyllaceae Portulacaceae Potamogetonaceae Sapotaceae 95 200 40 5 160 13 Cosmop Cosmop, Madag, IP S USA, Mexico, S Am trop, subtrop & temp regions Cosmop trop. Am, Afr, Asia, Aus, NZ, New Caled ANT-P 150 Cosmop, JF 15 90 Cosmop Andes Venezuela to Arg, Chile AND 500 subcosmop 16 Cosmop 7 Peru, Chile 40 temp Eurasia, N Am, Chile 1 JF W-ANT AND 100 2 Chile Afr, Madag, Asia, Aus, Tasm, NZ, Mex, C & S Am, WI 4 Arg, Chile, Canad, USA 350 temp & cold & montane trops W-ANT ANT-P 20 trop & warm-temp regs 7 4 4 6 1 6 4 3 7 7 7 3 7 7 7 4 7 7 1 80 Asia, N & S Am, Afr, Aus W-ANT 4 4 Espaa, Marruecos, Aus, Tasm, NZ, Chile, Arg 7 1 1 1 61 1 2 2 1 1 7 1 8 1 2 3 4 5 1 2 6 1 -30,83 -17,58 -30,00 -29,00 -30,00 -17,72 -23,52 -18,00 -31,30 -30,00 -18,33 -21,50 -32,10 -54,00 -34,73 -30,00 -54,00 -46,75 -22,00 -52,88 -46,67 -55,00 -34,28 -28,80 -55,00 -33,03 -42,42 -26,16 -30,00 -41,50 -38,38 -19,86 -38,20 -32,33 -43,15 -32,14 -23,57 -38,25 -32,57 9 S Am, Brazil S-AMZ 3 3 10 trop Am, 1 Afr-Madag, India, Sri Lanka 1 1 1 N Chile 6 1 5 4 W N Am, 1 Chile ANT-P 4 1 -32,50 -29,15 -24,00 -25,00 -32,25 -18,42 -24,75 -18,00 -35,25 -30,28 -35,83 3 Peru, Bol, Arg, Chile AND 3 2 -18,18 270 Cosmop, JF, ID 7 18 -18,17 70 W N Am, South Am, E Asia, Aus C-PAC 4 19 -29,83 -54,00 -56,00 -21,95 -37,33 -33,50 -42,40 -34,42 -45,58 -19,12 -31,93 -56,00 -40,33 -38,25 -50,38 5 Chile 6 5 -31,00 -34,55 16 Am trop desde S US, Afr, Madag 1 1 -18,50 -18,67 -18,58 -32,78 -41,92 -37,08 -20,07 -34,92 -31,33 -33,20 -29,71 -38,92 -18,77 -28,34 -37,00 -37,79 -34,27 -43,11 1 S Am, Chile, Peru, Arg AND 3 1 -18,47 -33,83 -26,15 1 Chile 6 1 -35,33 -37,53 -36,43 30 Am US to Arg, JF W-NT 3 7 -29,50 -40,00 -34,75 10,50 2,20 15,37 0,17 3,55 24,17 37,83 3,77 4,83 4,35 18,40 9,42 13,33 0,70 7,18 38,00 5,08 7,97 14,55 0,00 23,17 17,15 0,00 25,00 16,75 4,28 29,37 28,67 23,70 4,28 10,47 33,50 0,93 1 Chile 6 1 -27,33 -28,88 -28,11 1,55 85 Am, Medit, circumboreal W-ANT 4 2 -35,50 -56,00 -45,75 20,50 6 Euro, Aus, NZ, Ethiopia, W S Am 7 1 -32,15 -40,28 -36,22 8,13

Filicopsida

Marsileales

Pinguicula

Asterids

Lamiales

Pintoa

Rosids

Zygophyllales

Piptochaetium

Monocots

Poales

Pitavia

Rosids

Sapindales

Pitraea

Asterids

Lamiales

Pityrogramma

Filicopsida

Pteridales

Placea

Monocots

Asparagales

Plagiobothrys

Asterids

Unplaced

Plantago

Asterids

Lamiales

Plazia

Asterids

Asterales

Plectritis

Asterids

Dipsacales

Pleocarphus

Asterids

Asterales

Pleopeltis

Filicopsida

Polypodiales

Pleurophora

Rosids

Myrtales

Pleurosorus

Filicopsida

Aspleniales

Pluchea

Asterids

Asterales

9

Plumbago

Eudicots

Caryophyllales

Poa

Monocots

Poales

Podagrostis

Monocots

Poales

Podanthus

Asterids

Asterales

Podocarpus

Pinopsida

Pinales

Podophorus

Monocots

Poales

Polemonium

Asterids

Ericales

Polyachyrus

Asterids

Asterales

Polycarpon

Eudicots

Caryophyllales

Polygala

Rosids

Fabales

Polygonum

Eudicots

Caryophyllales

Polylepis

Rosids

Rosales

Polypodium

Filicopsida

Polypodiales

Polypogon

Monocots

Poales

Polystichum

Filicopsida

Dryopteridales

Porlieria

Rosids

Zygophyllales

Portulaca

Eudicots

Caryophyllales

Potamogeton

Monocots

Alismatales

Pouteria

Asterids

Ericales

Pozoa Apiaceae Campanulaceae Primulaceae Fabaceae Asteraceae Podocarpaceae Araliaceae Asteraceae Psilotaceae Pteridaceae Cactaceae Polygalaceae Poaceae Bromeliaceae Caryophyllaceae Cyperaceae Quillajaceae Santalaceae Ranunculaceae Caryophyllaceae Poaceae Rhamnaceae Solanaceae Rhamnaceae Verbenaceae Amaryllidaceae Poaceae Cyperaceae Grossulariaceae Asteraceae Brassicaceae Juncaceae Poaceae Rosaceae Polygonaceae Dryopteridaceae 7 500 200 2 2 75 cosmop NZ, S S Am, Tfuego, Falkland Arg, Caribe, Chil, Mexico, Peru, USA ,Urug cosmop, JF cosmop circumaustral, J Fernandez, Madag C-AUS AUS W-NT 7 JF 150 250 cosmp N Hemis, N Afr, S Am Canada to Tfuego W-ANT 2 S Brasil, Urug, Chile 30 Chile, Arg, Bol, Bras 2 Chile, Arg, JF S-AMZ S-AMZ 160 N Hemisp, S Afr, S Am 4 Chile Arg 3 Chile, Arg AND W-ANT 1 Chile Arg 1 Chile 500 Cosmop, R acaulis disj NZ, Falkland, R bonariensis disj N Am S Am 25 Andes Bol, Chile, Per, Arg AND 3 7 6 5 5 3 4 5 3 3 7 4 6 7 2 3 7 7 2 3 temp S Am, Arg, Chile, Peru, Brazil S-AMZ 3 70 pantrops 1 1 1 16 17 1 1 2 4 1 1 9 1 1 13 6 6 1 1 2 10 1 -33,60 -46,43 -25,00 -36,90 -30,00 -30,67 -43,83 -56,00 -26,38 -55,67 -55,00 -55,08 -38,72 -51,22 -25,69 -46,28 -42,50 -42,88 -30,00 -18,18 -17,78 -18,00 -35,85 -31,62 -19,10 -35,83 -30,67 -25,00 -33,00 -37,83 -29,83 -38,23 -51,00 -54,92 -22,58 -42,08 -37,80 -30,20 -41,00 -45,42 -40,00 -35,62 -39,23 -56,00 -34,12 -34,59 -36,35 -20,29 -38,97 -34,71 -24,65 -38,42 -38,04 -32,50 -34,31 -38,53 -42,92 17 Andes Peru, Chile, Arg, Bol AND 3 5 190 C & S Am, Brazil W-NT 3 8 80 N temp: Asia, N Am, S temp: S Afr, Aus, S Am 7 8 130 Am SW US to Chile W-NT 3 4 -29,00 -20,35 -22,50 -18,17 9 Patagonia Arg, Chile 5 1 -46,50 280 Cosmop, Madag, JF 7 2 -32,55 2 pantrop, Hawaii, Australia, NZ, Madag, IP 1 1 -49,70 -46,50 -37,83 -56,00 -37,80 -24,73 -41,13 -46,50 -33,42 -38,18 -30,15 -21,45 8 W US, temp S Am ANT-P 4 2 -30,00 -38,00 10 Tasm, NZ, Chile AUS 2 2 -35,33 -52,33 8 Costa Rica to Chile, New Caled, NZ, E Aus AUS 2 1 -35,83 -39,50 -37,67 -43,83 -34,00 3 Bol, Chile, Arg, Peru AND 3 3 -18,00 -40,33 -29,17 44 trop & subtrop Am, Afr, W Asia 1 6 -19,00 -34,33 -26,67 15,33 22,33 3,67 17,00 8,00 0,00 17,15 0,00 8,83 35,65 15,30 6,57 0,00 8,23 32,82 37,13 4,58 6,23 6,18 11,10 5,17 14,75 15,00 2,62 1,40 26,17 0,00 10,23 9,57 1,38 18,77 25,00 24,42 400 N Hemisp, trop Asia, temp S Am W-ANT 4 1 -38,50 -55,83 -47,17 17,33 15 SE Asia tropical, Aus, NZ, Am Austral SPT 2 1 -18,83 -56,00 -37,42 37,17 2 Andes Chile Arg 5 2 -31,50 -38,58 -35,04 7,08

Asterids

Apiales

Pratia

Asterids

Asterales

Primula

Asterids

Ericales

Prosopis

Rosids

Fabales

Proustia

Asterids

Asterales

Prumnopitys

Pinopsida

Pinales

Pseudopanax

Asterids

Apiales

Psilocarphus

Asterids

Asterales

Psilotum

Psilotopsida

Psilotales

Pteris

Filicopsida

Pteridales

Pterocactus

Eudicots

Caryophyllales

Pteromonnina

Rosids

Fabales

Puccinellia

Monocots

Poales

Puya

Monocots

Poales

Pycnophyllum

Eudicots

Caryophyllales

Pycreus

Monocots

Poales

Quillaja

Rosids

Fabales

40

Quinchamalium

Eudicots

Santalales

Ranunculus

Eudicots

Ranunculales

Reicheella

Eudicots

Caryophyllales

Relchela

Monocots

Poales

Retanilla

Rosids

Rosales

Reyesia

Asterids

Solanales

Rhamnus

Rosids

Rosales

Rhaphithamnus

Asterids

Lamiales

Rhodophiala

Monocots

Asparagales

Rhombolytrum

Monocots

Poales

Rhynchospora

Monocots

Poales

Ribes

Eudicots

Saxifragales

Robinsonia

Asterids

Asterales

Rorippa

Rosids

Brassicales

Rostkovia

Monocots

Poales

Rostraria

Monocots

Poales

Rubus

Rosids

Rosales

Rumex

Eudicots

Caryophyllales

Rumohra

Filicopsida

Dryopteridales

Ruppia Ruppiaceae Poaceae Caryophyllaceae Alismataceae Salicaceae Solanaceae Solanaceae Lamiaceae Samolaceae Caryophyllaceae Apiaceae Santalaceae Amaranthaceae Gesneriaceae Lamiaceae Podocarpaceae Saxifragaceae Saxifragaceae Saxifragaceae Anacardiaceae Schizaeceae Solanaceae Apiaceae Brassicaceae Asteraceae Cyperaceae Cyperaceae Lamiaceae Loasaceae Boraginaceae Goodeniaceae Asteraceae Fabaceae Hymenophyllaceae Cucurbitaceae Asteraceae 50 3 1 300 1250 Cosmop pantrops S S Am, JF, Falkland Aus, Tasm, NZ, Hawaii, N & S Am, ID subtro & trop regs C-PAC 2 1 JF Aus, NZ, Chile AUS 2 Chile 360 cosmop 20 100 subcosmop North temperate, austral, JF, IP 6 trop & subtrop Am 10 C N Chile, Arg 13 11 NZ, 1 Aus, 1 S S Am 12 Chile, Arg 30 trop & austral, N Am, Madag AND AUS AND W-NT 27 Mexico to Arg 1 Tfuego W-NT 1 Antarctic S Am 5 5 3 1 3 2 3 3 7 7 7 6 6 2 7 1 5 4 1 350 N temp, Eurasia, W N Am, Andes S Am W-ANT 4 1 Chile, Arg 5 38 temp & subtrop, Medit 7 4 1 1 1 1 9 1 12 1 10 2 4 20 3 2 1 1 218 9 1 1 1 -28,45 -17,58 -18,00 -39,63 -20,17 -32,83 -42,72 -56,00 -40,00 -56,00 -33,40 -42,72 -35,58 -36,79 -29,00 -47,82 -26,78 -37,78 1 Chile 6 1 16 Eurasia, Afr, Aus, Tasm, NZ, N & S Am, JF 7 3 25 Indomalasia to Aus, Hawaii, JF (ex) SPT 2 1 -20,63 -30,60 -18,50 -36,00 -36,00 -50,72 -51,00 -18,00 -39,88 -21,03 -38,58 -26,43 -17,58 -35,83 -17,67 -35,33 -31,00 -54,00 -43,83 -54,00 -45,75 -55,00 -56,00 -54,83 -46,55 -49,15 -41,00 -54,48 -35,33 -31,83 -56,00 -56,00 -53,00 -38,23 -37,32 -37,22 -36,25 -40,88 -45,50 -53,36 -52,92 -32,28 -44,52 -31,02 -46,53 -30,88 -24,71 -45,92 -36,83 -44,17 -34,62 40 subcosmop, exc Aus & NZ 7 2 -30,67 1 Desventuradas Islands (Chile) 6 1 -44,50 -37,58 12 Cosmop, IP 7 3 -33,42 -55,00 900 cosmop 7 4 -22,00 -25,33 2 Chile, Arg 5 2 -27,00 -40,00 15 S Am: Ecuad, Peru, Bol, Bra, Parag, Uru, Chile, Arg AND 3 2 -18,25 -19,87 -19,06 -33,50 -23,67 -44,21 450 Arctic temp, few in trops, temp S Am, 1 S Afr W-ANT 4 1 -18,58 -38,78 -28,68 25 Cosmop 7 1 -36,50 -40,00 -38,25 3,50 20,20 1,62 13,00 3,33 21,58 0,00 13,83 0,00 33,37 13,23 35,50 9,75 19,00 5,28 3,83 28,55 9,27 19,97 15,90 8,90 14,25 20,17 38,33 17,67 7,23 0,00 14,27 38,42 22,00 16,37 13,23 9,88 25 temp N Hemis, E Afr, Asia, N Guinea, Aus, NZ, Andes 7 1 -31,50 -42,50 -37,00 11,00 28 Australia, NZ, Arg, Chile, IP AUS 2 6 -30,67 -51,00 -40,83 20,33 10 cosmop 7 2 -21,00 -55,00 -38,00 34,00

Monocots

Alismatales

Rytidosperma

Monocots

Poales

Sagina

Eudicots

Caryophyllales

Sagittaria

Monocots

Alismatales

Salix

Rosids

Malpighiales

Salpichroa

Asterids

Solanales

Salpiglossis

Asterids

Solanales

Salvia

Asterids

Lamiales

Samolus

Asterids

Ericales

Sanctambrosia

Eudicots

Caryophyllales

Sanicula

Asterids

Apiales

Santalum

Eudicots

Santalales

Sarcocornia

Eudicots

Caryophyllales

Sarmienta

Asterids

Lamiales

Satureja

Asterids

Lamiales

Saxe-gothaea

Pinopsida

Pinales

Saxifraga

Eudicots

Saxifragales

4

Saxifragella

Eudicots

Saxifragales

Saxifragodes

Eudicots

Saxifragales

Schinus

Rosids

Sapindales

Schizaea

Filicopsida

Schizaeales

Schizanthus

Asterids

Solanales

Schizeilema

Asterids

Apiales

Schizopetalon

Rosids

Brassicales

Schkuhria

Asterids

Asterales

Schoenus

Monocots

Poales

Scirpus

Monocots

Poales

Scutellaria

Asterids

Lamiales

Scyphanthus

Asterids

Cornales

Selkirkia

Asterids

Unplaced

Selliera

Asterids

Asterales

Senecio

Asterids

Asterales

Senna

Rosids

Fabales

Serpyllopsis

Filicopsida

Hymenophyllales

Sicyos

Rosids

Cucurbitales

Sigesbeckia

Asterids

Asterales

Silene Caryophyllaceae Brassicaceae Iridaceae Apocynaceae Solanaceae Alliaceae Iridaceae Asteraceae Asteraceae Fabaceae Poaceae Alliaceae Caryophyllaceae Malvaceae Asteraceae Poaceae Lamiaceae Valerianaceae Caryophyllaceae Gratiolaceae Acanthaceae Amaryllidaceae Asteraceae Poaceae Potamogetonaceae Amaranthaceae Asteraceae Iridaceae Fabaceae Malvaceae Asteraceae Bignoniaceae Tecophilaeaceae Myrtaceae Asteraceae Francoaceae 1 1 1 2 Chile Chile, Arg S Am, Brazil Chile S-AMZ 12 60 30 3 Mexico, C Am to S Am 2 sp Mexico, disjunct to Peru, Chile, Arg temp N Hemis & temp S Am, NZ, JF Am tropical Arizona to Chile, 1 S Afr, Aus? 1 Chile, Arg W-NT ANT-P W-ANT 50 N & S Am 100 cosmop, ID W-NT 6 Subcosmop 300 temp N Hemis, montane trops, IP 230 S US to Chile 40 Andes 40 S US to C Chile 45 Trops and Subtrops, Madag W-NT AND W-NT 150 cosmop W-ANT 5 Arg,Chil,Per AND 3 7 4 3 3 3 7 7 7 3 5 3 4 4 1 6 5 3 6 300 trop, subtrop, temp regions exc. Australasia 7 160 Am, Asia, Afr, 1 Euro 7 1 11 1 5 1 1 1 2 1 3 4 3 1 1 10 2 1 2 1 1 1 -18,00 -20,25 -17,58 -40,50 -18,47 -18,17 -33,45 -18,47 -30,00 -35,40 -18,42 -31,50 -53,00 -55,00 -36,60 -56,00 -18,47 -38,78 -56,00 -18,50 -33,00 -53,00 -36,50 -35,42 -35,50 -37,63 -27,09 -48,25 -18,47 -28,48 -44,73 -18,48 -31,50 -44,20 -27,46 -33,46 6 trops 1 1 60 W N Am, Mex, temperate S Am ANT-P 4 2 30 cosmop, JF, ID 7 13 -18,00 -29,45 -18,47 -23,62 -22,00 -17,67 -18,00 -29,00 -30,00 -18,00 -17,58 1 Chile 6 2 -32,92 20 Am, W Euro, NW Afr W-ANT 4 1 -36,67 47 trop & most N temp, JF, IP 7 2 -32,00 -45,42 -37,50 -34,27 -40,00 -37,78 -18,47 -23,62 -44,00 -18,08 -55,83 -38,00 -37,80 -18,17 -25,43 8 S Am, N Am, Aus C-PAC 4 4 -29,95 -41,83 150 Temperate America, Macaronesia, Eurasia W-ANT 4 2 -30,95 -44,00 2 Chile, Arg 5 2 -31,00 -50,00 -40,50 -37,48 -35,89 -38,71 -37,08 -33,59 -29,00 -33,62 -18,47 -23,62 -33,00 -17,88 -36,92 -33,50 -33,90 -18,08 -21,51 2 Chile, Arg AND 3 1 -33,00 -36,83 -34,92 1700 subcosmop, JF, ID, IP 7 51 -18,20 -46,50 -32,35 3,83 19,00 13,05 11,88 13,42 0,83 1,35 22,00 8,33 0,00 0,00 22,00 0,42 37,83 9,00 7,80 0,17 7,85 0,00 35,00 34,75 19,02 15,50 0,00 20,62 22,55 0,03 3,00 17,60 18,08 3,92 3 Brasil, Chile S-AMZ 3 1 -25,20 -28,93 -27,07 3,73 28,30 60 N, C & S Am W-NT 3 11 -24,50 -55,25 -39,88 30,75 90 temp Eurasia, Medit, S Afr, Andes, N Am W-ANT 4 23 -18,20 -55,00 -36,60 36,80 600 temp N hemis, Medit, S Afr 7 8 -18,00 -55,00 -36,50 37,00

Eudicots

Caryophyllales

Sisymbrium

Rosids

Brassicales

Sisyrinchium

Monocots

Asparagales

Skytanthus

Asterids

Gentianales

Solanum

Asterids

Solanales

Solaria

Monocots

Asparagales

Solenomelus

Monocots

Asparagales

Solidago

Asterids

Asterales

Soliva

Asterids

Asterales

Sophora

Rosids

Fabales

Spartina

Monocots

Poales

Speea

Monocots

Asparagales

Spergularia

Eudicots

Caryophyllales

Sphaeralcea

Rosids

Malvales

Spilanthes

Asterids

Asterales

Sporobolus

Monocots

Poales

Stachys

Asterids

Lamiales

Stangea

Asterids

Dipsacales

4

Stellaria

Eudicots

Caryophyllales

Stemodia

Asterids

Lamiales

Stenandrium

Asterids

Lamiales

Stenomesson

Monocots

Asparagales

Stevia

Asterids

Asterales

Stipa

Monocots

Poales

Stuckenia

Monocots

Alismatales

Suaeda

Eudicots

Caryophyllales

Tagetes

Asterids

Asterales

Tapeinia

Monocots

Asparagales

Tara

Rosids

Fabales

Tarasa

Rosids

Malvales

Taraxacum

Asterids

Asterales

Tecoma

Asterids

Lamiales

Tecophilaea

Monocots

Asparagales

Tepualia

Rosids

Myrtales

Tessaria

Asterids

Asterales

Tetilla

Rosids

Geraniales

Tetrachondra Tetrachondraceae Rosaceae Aizoaceae Juncaginaceae Lamiaceae Asteraceae Thelypteridaceae Dicksoniaceae Iridaceae Bromeliaceae Boraginaceae Amaryllidaceae Rhamnaceae Saxifragaceae Asteraceae Hymenophyllaceae Poaceae Laxmanniaceae Fabaceae Juncaginaceae Campanulaceae Poaceae Asteraceae Poaceae Alliaceae Loranthaceae Malvaceae Asteraceae Tropaeolaceae Brassicaceae Cactaceae Apocynaceae Typhaceae Myrtaceae Cyperaceae Verbenaceae 60 1 10 13 6 9 4 50 Mexico to Chile California, Chile Andes Peru, Bol, Chile, Arg Bol, Uru, Chile, Arg Cosmop Am from Mex to Chile, JF Malesia, Pacific, Aus, SE Asia, C & S Am, JF Chile, Arg W-NT SPT AND 50 SW N Am to Chile 70 Pantrop W-NT W-NT ANT-P AND S-AMZ 11 Andes Colombia to Chile 6 Chile, Arg, Uru AND 70 cold & temp, montane tropics, JF 12 C Chile, 1 Patagonia 30 trops & subtrops 8 N Am, Mex, Guatem, Medit 12 N temperate, circumboreal, austral 250 temp & subtrop exc Aus 2 Chile, Arg W-ANT W-ANT W-ANT 7 Arabia, trop Afr, S US, Peru 80 patrop, Madag, JF 1 1 5 4 4 4 1 5 7 5 3 1 3 3 4 3 3 7 3 2 3 22 S Am, 1 W Aus AUS 2 1 Chile & TFuego 5 1 5 1 1 1 8 3 1 1 12 14 12 3 1 1 18 1 2 2 2 2 10 1 -18,55 -22,08 -30,67 -18,00 -26,75 -19,17 -34,17 -30,67 -18,00 -20,93 -44,00 -33,38 -22,00 -37,67 -39,00 -45,38 -56,00 -24,73 -19,74 -33,04 -32,03 -20,00 -32,21 -29,08 -39,78 -43,33 -21,37 1 Chile 6 1 1 Chile 6 1 27 arid N & S Am ANT-P 4 5 -18,18 -31,50 -30,42 -40,50 -18,43 -39,87 -18,47 -25,43 -25,00 -20,33 -25,00 -18,38 -23,00 -25,00 -29,00 -28,50 540 trop & subtrop Am, WI W-NT 3 4 -18,00 35 Mex, Guatemala disjunct to Peru, Chile, Bol, Brazil ANT-P 4 1 -22,00 1 JF 6 1 -25,40 -38,00 -29,00 -34,00 -35,83 -55,50 -37,82 -43,10 -18,47 -38,23 -40,00 -55,00 -40,00 -23,58 -42,50 -56,00 -54,83 -42,50 -23,70 -28,00 -23,59 -32,75 -33,13 -48,00 -28,13 -41,48 -18,47 -31,83 -32,50 -37,67 -32,50 -20,98 -32,75 -40,50 -41,92 -35,50 800 trops & subtrops, Madag 1 1 -18,27 -37,33 1 Desventuradas Islands (Chile) 6 1 -27,80 250 Cosmop 7 2 -25,00 -40,00 -32,50 1 temp S Am, Magallanes 5 1 -43,00 -56,00 -49,50 13,00 15,00 0,00 19,07 0,00 3,40 20,00 10,82 2,50 5,42 15,00 19,38 3,23 0,00 12,80 15,00 34,67 15,00 5,20 19,50 31,00 25,83 14,00 0,00 2,38 21,92 2,72 4,00 10,92 19,83 11,22 25,33 6,73 70 N & S Afr, SE Asia, Aus, Tasm, NZ, Polinesia, Peru, Chile W-ANT 4 10 -21,03 -31,92 -26,48 10,88 8 Arg, Chil, Per AND 3 3 -28,72 -46,17 -37,44 17,45 2 1 NZ, 1 Patagonia, Tfuego AUS 2 1 -49,00 -54,83 -51,92 5,83

Asterids

Lamiales

Tetraglochin

Rosids

Rosales

Tetragonia

Eudicots

Caryophyllales

Tetroncium

Monocots

Alismatales

Teucrium

Asterids

Lamiales

Thamnoseris

Asterids

Asterales

Thelypteris

Filicopsida

Blechnales

Thyrsopteris

Filicopsida

Dennstaedtiales

Tigridia

Monocots

Asparagales

Tillandsia

Monocots

Poales

Tiquilia

Asterids

Unplaced

Traubia

Monocots

Asparagales

Trevoa

Rosids

Rosales

Tribeles

Eudicots

Saxifragales

Trichocline

Asterids

Asterales

Trichomanes

Filicopsida

Hymenophyllales

Trichoneura

Monocots

Poales

4

Trichopetalum

Monocots

Asparagales

Trifolium

Rosids

Fabales

Triglochin

Monocots

Alismatales

Triodanis

Asterids

Asterales

Tripogon

Monocots

Poales

Triptilion

Asterids

Asterales

Trisetum

Monocots

Poales

Tristagma

Monocots

Asparagales

Tristerix

Eudicots

Santalales

Triumfetta

Rosids

Malvales

Trixis

Asterids

Asterales

Tropaeolum

Rosids

Brassicales

Tropidocarpum

Rosids

Brassicales

Tunilla

Eudicots

Caryophyllales

Tweedia

Asterids

Gentianales

Typha

Monocots

Poales

Ugni

Rosids

Myrtales

Uncinia

Monocots

Poales

Urbania

Asterids

Lamiales

Urmenetea Asteraceae Urticaceae Lentibulariaceae Escalloniaceae Valerianaceae Caricaceae Verbenaceae Asteraceae Solanaceae Fabaceae Asteraceae Asteraceae Violaceae Vittariaceae Vivianiaceae Poaceae Campanulaceae Brassicaceae Asteraceae Cunoniaceae Ledocarpaceae Brassicaceae Asteraceae Lemnaceae Lemnaceae Dryopteridaceae Asteraceae Brassicaceae Rosaceae Asteraceae Cyperaceae Potamogetonaceae Rutaceae Tecophilaeaceae Alliaceae 2 1 250 5 cosmop Am, Afr, Asia, JF Chile Chile, Arg AND 3 1 JF Arg, Chile, Bol AND 1 JF 6 S Arg, Chile 21 25 Andes Colombia to N Arg, Chile 10 trop & warm Am, 1 S Afr temp & cool temp Eurasia, Afr, Am, exc Aus W-ANT AND 11 40 Andes Ecuador to Chile cosmop, warm temperate and tropical regions 3 N Chile, Arg 3 Chile, Arg AND AND 160 Madag, Australasia, Mex, C & S Am 100 pantrops, W Australia C-AUS 18 Andes Peru, Chile, Arg, Bol AND 150 Southern Hemisphere, Europe, SE Asia W-ANT 4 3 1 2 5 3 3 7 1 4 3 5 6 6 3 7 1 6 3 25 N & S Am, Euro, N Afr, M Asia 7 6 S Brasil, Chile, Arg S-AMZ 3 3 2 1 7 1 1 1 2 9 1 1 1 5 2 1 1 3 1 2 1 2 -20,82 -30,00 -29,00 -33,83 -24,91 -31,92 -17,58 -22,00 -33,67 -54,00 -25,63 -38,00 70 pantrop 1 1 -28,72 -23,33 -30,00 -30,67 -23,58 -35,33 -30,50 -21,27 -17,58 -36,75 -33,33 -18,22 -18,00 -51,00 -36,83 -37,33 -40,00 -36,90 -27,37 -49,42 -41,73 -26,33 -33,82 -36,75 -36,75 -18,35 -23,58 -51,00 -32,78 -30,33 -35,00 -33,78 -25,48 -42,38 -36,12 -23,80 -25,70 -36,75 -35,04 -18,28 -20,79 -51,00 430 cosmop 7 70 10 Mex to Chile W-NT 3 2 180 trop & subtrop Am, WI W-NT 3 6 -18,18 -18,20 -18,00 140 temp N Hemis, Medit, few trop Afr, S Am, Hawaii W-ANT 4 28 -25,00 1 Chile 6 1 -33,03 -40,00 -55,33 -37,80 -25,00 -56,00 300 trop & subtrop Am, WI W-NT 3 2 -22,00 -29,90 225 temp & trop Am, few OW W-ANT 4 33 -18,00 -40,00 20 Mexico to N Arg & S Brasil W-NT 3 1 -29,67 -33,00 -31,33 -29,00 -25,95 -36,52 -40,17 -27,99 -21,60 -37,00 230 N temp Eurasia, S Afr, Andes W-ANT 4 43 -18,00 -55,00 -36,50 1 Chile 6 1 -39,88 -40,25 -40,07 0,37 37,00 3,33 22,00 7,90 6,97 30,33 19,62 6,80 38,00 0,00 8,12 14,00 10,00 6,23 3,78 14,08 11,23 5,07 16,23 0,00 3,42 0,13 5,58 0,00 0,00 0,00 16,08 32,00 0,00 8,18 3,83 200 cosmop 7 1 -28,72 -43,00 -35,86 14,28 75 subcosmop, temperate 7 10 -17,72 -55,50 -36,61 37,78 1 N Chile, NW Arg AND 3 1 -21,50 -23,18 -22,34 1,68

Asterids

Asterales

Urtica

Rosids

Rosales

Utricularia

Asterids

Lamiales

Valdivia

Asterids

Unplaced

Valeriana

Asterids

Dipsacales

Vasconcellea

Rosids

Brassicales

Verbena

Asterids

Lamiales

Verbesina

Asterids

Asterales

Vestia

Asterids

Solanales

Vicia

Rosids

Fabales

Viguiera

Asterids

Asterales

Villanova

Asterids

Asterales

Viola

Rosids

Malpighiales

Vittaria

Filicopsida

Pteridales

Viviania

Rosids

Geraniales

Vulpia

Monocots

Poales

Wahlenbergia

Asterids

Asterales

44

Weberbauera

Rosids

Brassicales

Wedelia

Asterids

Asterales

Weinmannia

Rosids

Oxalidales

Wendtia

Rosids

Geraniales

Werdermannia

Rosids

Brassicales

Werneria

Asterids

Asterales

Wolffia

Monocots

Alismatales

Wolffiella

Monocots

Alismatales

Woodsia

Filicopsida

Dryopteridales

Xenophyllum

Asterids

Asterales

Xerodraba

Rosids

Brassicales

xMargyracaena

Rosids

Rosales

Yunquea

Asterids

Asterales

Zameioscirpus

Monocots

Poales

Zannichellia

Monocots

Alismatales

Zanthoxylum

Rosids

Sapindales

Zephyra

Monocots

Asparagales

Zoellnerallium

Monocots

Asparagales

Appendix A (continuation): Legend and summary for tracks and elements

TRACKS (follow subsections in the text) 3.2.1 Austral-antarctic track 3.2.2 South Pacific tropical track 3.2.3 Circum-austral track 3.3.1 Wide Neotropical track 3.3.2 Andean track 3.3.3 South Amazonian track 3.4.1 Wide antitropical track 3.4.2 Antitropical disjunct Pacific track 3.4.3 Circum-Pacific track AUS SPT C-AUS W-NT AND S-AMZ W-ANT ANT-P C-PAC

N of genera 31 25 3 64 113 39 84 56 9

ELEMENTS 1. Pantropical 2. Australasiatic 3. Neotropical 4. Antitropical 5. South-temperate 6. Endemic 7. Cosmopolitan TOTAL

N of genera 88 59 216 152 81 83 134 813

4

4

Appendix B: Genera shared by several Chilean regions

47

GEN Aa Abrotanella Acacia Acaena Acantholippia Acrisione Adenocaulon Adenopeltis Adesmia Adiantum Aextoxicon Agalinis Agallis Ageratina Agoseris Agrostis Allionia Alona Alonsoa Alopecurus Aloysia Alstroemeria Alternanthera Amaranthus Amblyopappus Ambrosia Amomyrtus Amphibromus Amsimkia Amsinckia Anagallis Anarthrophyllum Anatherostipa Androsace Anemone Anisomeria Antennaria Anthochloa Anthoxanthum Antidaphne Aphanes Apium Apodasmia Arachnitis Araeoandra Araucaria Arenaria Argemone Argylia Aristeguietia Aristida Aristolochia Aristotelia

Summe 1 2 2 5 2 2 1 2 4 5 2 1 1 3 3 4 1 2 2 3 1 4 1 1 2 3 1 1 1 3 4 2 1 1 3 2 3 1 2 1 1 5 2 2 1 1 4 2 3 2 3 1 2

ANT 1

BIO

COQ

JF 1

MAG 1 1

GEN Laretia Larrea Lastarriaea Lasthenia Lathyrus Laurelia

Summe 1 2 1 3 4 1 1 1 1 3 2 4 1 1 1 1 1 2 1 1 4 3 1 3 2 4 1 4 1 1 3 2 2 1 2 4 3 1 3 2 3 2 2 3 4 2 2 1 3 1 1 1 3

ANT

BIO 1 1

COQ 1 1 1 1 1

JF

MAG

1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1

1 1 1

Laureliopsis Lebetanthus Legrandia Lemna Lenzia Lepidium Lepidoceras Lepidophyllum

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

1 1 1 1 1 1 1

Lepidothamnus Leptinella Leptocarpha Leptophyllochloa Leptostigma

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Lepuropetalon Leucheria Leucocoryne Leunisia Libertia Ligaria Lilaeopsis Limonium Limosella Linanthus Lindernia Linum Lippia Lithrea Littorella Llagunoa Loasa Lobelia locenes Lomatia Lophosoria Lotus Ludwigia Luma Lupinus Luzula Luzuriaga Lycium Lycopersicon Lycopodium Lysimachia Machaerina Macrachaenium Madia

48

Arjona Armeria Artemisia Arthropteris Asplenium Astelia Aster Asteranthera Asteriscium Astragalus Atriplex Austrocedrus Azara Azolla Azorella Baccharis Bacopa Bahia Bakerolimon Balbisia Balsamocarpon Barneoudia Bartsia Beilschmiedia Belloa Benthamiella Berberidopsis Berberis Bidens Bipinnula Blechnum Blennosperma Blepharocalyx Boehmeria Boerhavia Boisduvalia Bolax Bomarea Boopis Boquila Botrychium Bouteloua Bowlesia Brachyclados Brachystele Bridgesia Bromidium Bromus Buddleja Bulbostylis Bulnesia Caesalpinia

2 3 2 1 5 1 3 2 3 4 4 1 3 4 4 4 2 3 2 2 1 1 1 1 2 1 1 5 3 3 4 1 1 1 2 2 1 1 2 1 1 1 4 1 2 1 2 5 2 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1

Maihuenia Maihueniopsis Malacothamnus Malacothrix Malesherbia Malvella Mancoa Margyricarpus Marsilea Marsippospermum Mastigostyla Mathewsia Maytenus

1 2 1 2 2 2 1 4 1 2 1 2 3 1 3 2 2 1 4 2 3 2 3 3 1 1 5 1 1 2 2 3 3 3 2 2 1 1 2 2 4 1 3 3 1 3 3 1 4 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

Megalachne Megalastrum Melica Melosperma Menodora Menonvillea Mentzelia Microphyes Micropsis Microseris Microsteris Miersia

1 1 1 1 1 1 1 1 1

Mikania Mimulus Minuartia Miqueliopuntia Mirabilis Misodendrum Mitraria Montia

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Montiopsis Monttea Moscharia Moschopsis Moschopss Muehlenbeckia Muhlenbergia Mulinum Munroa Mutisia Myoschilos Myosotis Myosurus Myrceugenia Myrcianthes Myriophyllum Myrteola Nama Nanodea

49

Caiophora Calamagrostis Calandrinia Calceolaria Caldcluvia Calliandra Callitriche Calopappus Calotheca Caltha Calycera Calydorea Calystegia Camissonia Campsidium Cardamine Cardionema Carex Carica Carpha Carpobrotus Castilleja Catabrosa Centaurea Centaurium Centaurodendron Centella Centipeda Cerastium Ceratophyllum Cestrum Chaetanthera Chaetotropis Chaptalia Chascolytrum Cheilanthes Chenopodium Chersodoma Chevreulia Chiliophyllum Chiliotrichum Chiropetalum Chloraea Chorizanthe Chrysosplenium Chuquiraga Chusquea Cicendia Ciclospermum Cissarobryon Cissus Cistanthe

3 4 4 4 1 1 4 1 1 3 3 2 2 2 2 4 3 5 1 1 3 2 3 3 3 1 2 2 3 1 3 3 2 1 2 3 5 1 1 1 2 3 3 2 1 2 3 2 1 1 2 3

1 1 1 1

1 1 1 1 1

1 1 1 1 1 1 1 1

Nardophyllum Nassauvia Nassella Nastanthus Nasturtium Navarretia 1 Nertera Nicandra Nicotiana 1 Nierembergia Nitrophila Nolana 1 Notanthera Nothofagus 1 Notholaena Nothoscordum Noticastrum 1 1 1 Nototriche Ochagavia Oenothera Olsynium Ombrophytum 1 Onuris Ophioglossum Ophryosporus 1 Oreobolus Oreocereus Oreomyrrhis 1 Oreopolus Orites Ornithopus Orobanche Ortachne Osmorhiza 1 1 1

3 3 5 3 1 2 4 1 4 1 2 3 2 2 2 2 1 2 3 4 4 1 1 4 2 3 1 1 2 1 1 1 1 3 2 3 1 4 2 1 2 3 1 1 1 1 3 3 3 2 1 3

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1

1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1

1 1

1 1

1 1 1 1

1 1 1 1 1 1

1 1 1 1 1

1 1 1 1

1 1

1 1

1 1 1

1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1

1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1

Otholobium Ourisia Ovidia Oxalis Oxychloe 1 Oxyphyllum Oxytheca Oziroe 1 1 Pachylaena Palaua Panicum Parastrephia 1 Parietaria Paronychia Pasithea Paspalum 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Passiflora Patosia

0

Citronella Clarkia Cleome Cliococca Codonorchis Colletia Colliguaja Collomia Colobanthus Combera Conanthera Convolvulus Conyza Copiapoa Coprosma Cordia Coriaria Coronopus Corrigiola Cortaderia Corynabutilon Cotula Crassula Cressa Crinodendron Cristaria Croton Cruckshanksia Cryptantha Cryptocarya Cryptogramma Cuatrecasasiella Cuminia Cumulopuntia Cuscuta Cyclospermum Cynodon Cynoglossum Cyperus Cyphocarpus Cystopteris Dalea Danthonia Dasyphyllum Daucus Dendroseris Dennstaedtia Deschampsia Descurainia Desfontainia Desmaria Deuterocohnia Dichondra Dicksonia

2 2 1 1 2 3 3 2 4 1 3 3 4 2 1 1 1 2 2 4 2 1 4 2 1 2 1 2 3 2 2 1 1 2 3 2 2 1 4 1 4 1 3 1 3 1 1 3 3 2 1 1 4 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1

1 1

Pectocarya Pellaea Pelletiera Pennisetum 1 Peperomia Perezia Perityle 1 Persea Peumus Phacelia Phalaris Philesia 1 1 Philibertia Philippiella Phleum Phragmites Phrodus Phycella Phyllachne 1 Phylloscirpus Phytolacca Picrosia 1 Pilea Pilgerodendron Pilostyles Pilularia Pinguicola Pinguicula Piptochaetium Pitavia Pitraea Placea 1 Plagiobothrys Plantago Plazia Plectritis Pleocarphus Pleopeltis 1 1 1 1 1 1 1 Pleurophora Pleurosorus Pluchea Plumbago Poa Podagrostis Podanthus Podocarpus Podophorus 1 1 1 Polemonium Polyachyrus Polycarpon Polygala Polygonum 1

3 3 1 2 3 4 2 1 2 4 2 1 1 1 2 3 1 2 1 1 1 1 1 1 3 2 1 1 3 1 2 1 3 5 1 1 1 3 2 2 1 2 4 1 2 2 1 2 2 1 3 2 1 5

1 1 1 1 1 1

1 1 1

1 1 1 1 1 1

1 1 1 1 1 1 1 1

1 1 1

1 1 1 1 1

1 1

1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Polylepis Polypodium



Dicllptera Dielsiochloa Dinemagonum Dinemandra Dioscorea Diostea Diplachne Diplolepis Diplostephium Diposis Discaria Distichia Distichlis Dodonaea Domeykoa Donatia Doniophyton Downingia Draba Drapetes Drimys Drosera Drymaria Dysopsis Eccremocarpus Echinopsis Elaphoglossum Elatine Eleocharis Elodea Elymus Elytropus Embothrium Empetrum Encelia Enneapogon Ephedra Epilobium Epipetrum Equisetum Eragrostis Ercilla Erechtites Eremocharis Eriachaenium Erigeron Eriosyce Errazurizia Eryngium Escallonia Eucryphia Eudema Eulychnia Euphorbia

1 1 1 1 3 2 1 3 1 1 3 1 3 1 1 1 2 2 3 1 4 1 1 4 1 2 1 1 5 1 4 1 2 3 2 1 4 4 3 2 3 1 1 1 1 4 3 2 4 4 1 2 2 4

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Polypogon Polystichum Porlieria Portulaca Potamogeton Pouteria Pozoa Pratia Primula Prosopis Proustia Prumnopitys Pseudopanax Psilocarphus Pteris Pteromonnina Puccinellia Puya Pycnophyllum Quillaja Quinchamalium Raimundochloa Ranunculus Reicheella Relchela Retanilla Reyesia Rhamnus Rhaphithamnus Rhodophiala Ribes Robinsonia Rorippa Rostkovia Rubus Rumex Rumohra Ruppia Rytidosperma Sagittaria Salix Salpiglossis Salvia Samolus Sanicula Santalum Sarcocornia Sarmienta Satureja Saxegothaea Saxifraga Saxifragella Saxifragodes Schinus

4 4 1 1 4 1 2 2 1 2 3 1 2 2 3 2 3 3 1 2 3 2 5 1 1 2 2 1 3 3 3 1 1 1 3 3 4 3 3 1 3 2 1 2 2 1 5 2 4 1 2 1 1 2

1 1

1 1 1

1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1



Euphrasia Evolvulus Exodeconus Fabiana Facelis Fagonia Fascicularia Festuca Flaveria Flourensia Fragaria Francoa Frankenia Fuchsia Fuertesimalva Gaimardia Galinsoga Galium Gamocarpha Gamochaeta Gamochaetopsis Gaultheria Gavilea Gayophytum Gentiana Gentianella Geoffroea Geranium Gethyum Geum Gevuina Gilia Gilliesia Glandularia Gleichenia Glinus Glyceria Glycyrrhiza Gnaphalium Gochnatia Gomortega Gomphrena Grabowskia Grammitis Grammosperma Gratiola Greigia Griselinia Guindilia Gunnera Gutierrezia Guynesomia Gymnachne Gymnophyton

4 1 1 3 3 2 1 4 3 2 1 1 4 3 2 1 3 5 2 5 1 5 4 2 4 4 2 4 1 2 1 3 2 3 3 2 1 1 4 2 1 1 1 3 1 1 2 4 1 4 3 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1

Schizaea Schizanthus Schizeilema

1 3 1 1 2 1 2 5 2 2 1 2 4 3 2 2 1 4 4 4 2 4 1 3 2 2 3 1 4 2 2 1 3 4 2 2 1 4 3 3 1 2 2 1 2 2 1 1 2 2 1 3 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1

1 1 1 1 1 1 1

Schizopetalon Schkuhria Schlnus Schoenus Scirpus Scutellaria Scyphanthus Selkirkia Selliera

1 1 1

1 1 1

1 1 1

Senecio Senna Serpyllopsis Sicyos Sigesbeckia Silene Sisymbrium Sisyrinchium Skytanthus Solanum Solaria Solenomelus Solidago Soliva Sophora Spartina Spergularia Sphacele Sphaeralcea Spirodela Stachys Stellaria

1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

1 1

Stemodia Stenandrium Stevia Stipa Suaeda Tagetes Tapeinia Tarasa Taraxacum

1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1

1 1

Tecophilaea Tepualia Tessaria Tetilla

1 1 1 1 1

Tetrachondra Tetraglochin Tetragonia Tetroncium Teucrium Thelypteris Thlaspi



Gypothamnium Habenaria Haloragis Hamadryas Haplopappus Hebe Hedyotis Helenium Helictotrichon Heliotropium Helogyne Herbertia Herreria Heterosperma Heterozostera Hieracium Hierochloe Hippuris Histiopteris Hoffmannseggia Homalocarpus Hordeum Hornungia Huanaca Huidobria Huperzia Hybanthus Hydrangea Hydrocotyle Hymenoglossum Hymenophyllum Hypericum Hypochaeris Hypolepis Hypsela Imperata Ipomopsis Isoetes Jaborosa Jarava Jovellana Juania Jubaea Juncus Junellia Kageneckia Koeleria Krameria Kurzamra Lactoris Lagenophora Lampaya Lapageria Lardizabala

1 2 1 1 3 1 2 3 1 4 1 1 1 1 1 2 2 2 2 2 2 4 1 2 2 1 1 1 3 3 4 2 4 4 2 2 1 3 4 3 1 1 1 5 3 2 1 2 1 1 3 1 2 2

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Thyrsopteris Tigridia Tillandsia Tiquilia Traubia Trevoa Tribeles Trichocline Trichomanes Trichopetalum Trifolium Triglochin Triodanis Triptilion Trisetum Tristagma Tristerix Tropaeolum Tunilla Tweedia Typha Ugni Uncinia Urbania Urmenetea Urtica Utricularia Valeriana Verbena Verbesina Vestia Vicia Viguiera Villanova Viola Viviania Vulpia Wahlenbergia Weberbauera Weinmannia Wendtia Werdermannia Werneria Wolffia Wolffiella Xenophyllum Xerodraba xMargyracaena Yunquea Zannichellia Zanthoxylum Zephyra Zoellnerallium

1 1 3 1 1 2 1 3 1 2 3 4 1 3 5 3 2 3 1 3 3 2 4 1 1 5 2 4 3 2 1 4 3 1 4 2 3 3 3 2 2 1 2 1 1 1 1 1 1 3 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1

1 1

1 1 1

1 1

1 1 1

1 1 1 1 1

4

Appendix C: Matrix for PAE: Distribution of Endemic Genera



N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

A an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

B co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

B an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

C co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

C an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

D co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 4

D an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2

E co 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4

E an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1

F co 0 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 6

F an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 2

G co 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5

G in 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

G an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

H co 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9



N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

H an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

I co 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 0 1 1 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 14

I an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

J co 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9

J an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2

K co 0 0 1 0 0 0 1 1 0 0 0 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 1 0 1 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 18

K an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

L co 0 0 1 1 0 0 0 1 1 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 15

L an 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5

M co 0 0 1 1 1 0 1 1 1 0 0 1 1 1 0 1 0 0 0 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 20

M an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5

N co 0 1 1 1 1 0 0 0 1 0 0 1 1 1 0 1 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 0 1 0 0 1 0 0 0 0 1 1 0 0 0 1 1 0 1 0 0 1 0 0 1 0 0 0 24

N an 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 7

7

N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

O co 1 1 1 1 0 0 0 0 1 0 0 1 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 1 1 0 1 0 0 1 0 1 1 0 0 1 1 1 0 0 1 0 1 1 1 1 0 0 0 26

O an 0 0 0 1 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 0 1 0 0 0 12

P co 1 1 1 1 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 0 0 1 1 0 1 0 1 1 0 1 0 0 0 1 1 1 0 0 1 1 1 1 1 1 0 1 0 28

P an 0 0 0 1 0 0 0 0 1 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 1 0 0 0 12

Q co 1 1 0 1 0 1 0 0 1 0 1 1 0 0 0 0 0 1 1 1 1 0 0 0 1 0 0 1 0 0 0 1 1 0 0 0 0 1 0 0 0 1 1 0 1 0 1 1 0 1 0 0 0 1 1 1 0 0 0 1 1 1 1 1 0 1 0 31

Q an 1 0 0 1 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 1 0 0 0 0 1 0 0 0 1 0 1 0 0 1 1 1 1 0 1 0 0 0 20

R co 1 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 1 0 1 0 1 1 0 1 0 0 0 1 0 1 0 0 0 0 0 1 1 1 0 0 0 20

R an 1 0 0 1 0 1 0 0 0 1 1 1 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 1 1 0 1 0 1 0 1 0 18

S co 1 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 1 1 0 0 0 0 1 0 1 0 1 1 0 1 0 0 1 0 0 1 0 1 1 0 0 1 0 1 0 1 0 26

S an 1 0 0 1 0 0 0 0 0 1 1 1 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 0 0 17

T co 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 1 0 0 1 0 1 1 0 1 0 0 1 0 0 1 0 1 1 0 0 0 0 1 0 1 0 22

T an 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 14

U co 1 0 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 1 1 1 0 1 0 1 0 0 0 0 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 1 1 0 1 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 21

U an 1 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 1 0 13

V co 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 14

8

N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

V an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 9

W co 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 15

W an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4

X co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 11

X an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 10

Y co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 5

Y an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4

Z co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 8

Z an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2

AA co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 3

AA an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AB co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AB an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

9

N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

AC co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AC an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AD co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AD in 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AD an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AE co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AE an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AF co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AF an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AG co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

AG an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AH co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AH an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AI co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AI an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0

N 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

GENERO Acrisione Adenopeltis Alona Anisomeria Araeoandra Avellanita Bakerolimon Balsamocarpon Bridgesia Calopappus Cissarobryon Conanthera Copiapoa Cyphocarpus Desmaria Dinemagonum Dinemandra Epipetrum Ercilla Fascicularia Francoa Gethyum Gomortega Guynesomia Gymnachne Gypothamnium Hollermayera Homalocarpus Huidobria Hymenoglossum Ivania Jubaea Lapageria Latua Legrandia Leontochir Leptocarpha Leucocoryne Leunisia Marticorenia Metharme Microphyes Miersia Miqueliopuntia Moscharia Neuontobothrys Notanthera Ochagavia Oxyphyllum Peumus Phrodus Pintoa Pitavia Placea Pleocarphus Podanthus Reicheella Sarmienta Scyphanthus Speea Tecophilaea Tetilla Traubia Trevoa Valdivia Vestia Zephyra TOT

AJ co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AJ in 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AJ an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AK co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AK in 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AK an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AL co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AL an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AM co 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

AM an 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

SUM 14 7 10 15 2 2 6 4 9 4 7 19 10 7 8 7 9 9 19 10 19 5 3 3 7 2 2 13 14 18 1 5 11 3 2 2 10 22 3 2 2 18 6 3 9 1 15 12 3 17 10 2 3 7 5 15 2 11 12 4 5 7 4 12 2 12 6





Appendix D: Native Genera in the Chilean Pacific Islands



Islas Desventuradas Atriplex Chenopodium Cristaria Eragrostis Frankenia Fuertesimalva Lepidium Lycapsus Maireana Nesocaryum Parietaria Plantago Sanctambrosia Sicyos Solanum Spergularia Suaeda Tetragonia Thamnoseris

Isla de Pascua Agrostis Apium Asplenium Axonopus Davallia Diplazium Doodia Dryopteris Elaphoglossum Ipomoea Juncus Kyllinga Lycium Microlepia Microsorum Ophioglossum Paspalum Polystichum Psilotum Pycreus Rytidosperma Samolus Scirpus Solanum Sophora Stipa Thelypteris Triumfetta Vittaria

Juan Fernndez Abrotanella Acaena Adiantum Agrostis Apium Arthropteris Asplenium Azara Berberis Blechnum Boehmeria Bromus Calystegia Cardamine Carex Centaurodendron Centella Chenopodium Chusquea Colletia Coprosma Cuminia Cyperus Cystopteris Danthonia Dendroseris Dichondra Dicksonia Drimys Dysopsis Elaphoglossum Eleocharis Empetrum Erigeron Eryngium Escallonia Euphrasia Galium Gamochaeta Gaultheria Gavilea Gleichenia Grammitis Greigia Gunnera Haloragis Hedyotis Histiopteris Hymenoglossum Hymenophyllum Hypolepis Juania Juncus Lactoris Lagenophora

Juan Fernndez (cont.) Leptophyllochloa Libertia Lobelia Lophosoria Luzula Lycopodium Machaerina Margyricarpus Megalachne Megalastrum Mimulus Myrceugenia Myrteola Nassella Nertera Nicotiana Notanthera Notholaena Ochagavia Ophioglossum Oreobolus Parietaria Peperomia Piptochaetium Plantago Pleopeltis Podophorus Polypodium Polypogon Polystichum Pteris Ranunculus Rhaphithamnus Robinsonia Rubus Rumohra Santalum Sarcocornia Scirpus Selkirkia Serpyllopsis Solanum Sophora Spergularia Taraxacum Thyrsopteris Trichomanes Trisetum Ugni Uncinia Urtica Wahlenbergia xMargyracaena Yunquea Zanthoxylum

4

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 pequea flor, mi pequeo roble, y mi pequeo amor. Agradezco tambin a Mlica Muoz, no slo por ser mi madre que ya es mucho, sino por compartir siempre sus inagotables conocimientos botnicos y por haber sido desde que tengo memoria el pilar fundamental de mi formacin naturalista. A travs de ella no puedo dejar de mencionar a mi abuelo Carlos Muoz y su esposa Ruth, quienes an nos iluminan en los momentos de desconcierto. A travs 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 slo 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, Ivn ha enviando imprescindible msica y videos; Simn ha descuidando su propia tesis para conseguir literatura indispensable. Federico Luebert ha revisado la tesis en forma exhaustiva y crtica, y ha sido junto con Patricio Pliscoff siempre partcipe de una discusin enriquecedora. Espero contar con su amistad por muchos aos (viva Berln!) Tambin Sergio Elrtegui ha sido un gran compaero en la distancia, mgico creador de ilustraciones e ilusiones. Agradezco tambin la amistad y la compaa musical de Carlos Ledermann, y el duende de los Mrtires del Comps; sin l hubiese perdo el rumbo hace tiempo. Meinem Doktorvater, Herrn Prof. Dr. Michael Richter, danke ich fr die engagierte Betreuung der Arbeit am Anfang und dafr, mir am Ende die notwendigen Freirume gewhrt zu haben. Frau Prof. Dr. Perdita Pohle und Herrn Prof. Dr. Achim Bruning danke ich herzlich fr die Bereitschaft, bei meinem Rigorosum den Prfungsvorsitz bernommen bzw. mich geprft zu haben. Dank dafr gebhrt natrlich auch Herrn Prof. Dr. Werner Nezadal, meinem Prfer im Nebenfach; ihm verdanke ich auch, viele Pflanzen am Cabo de Gata (Spanien) kennengelernt zu haben. Walter Wel hie mich in der Mitte seiner umfangreichen Bibliothek im Botanischen Garten willkommen; langsam sind viele Bcher auch zu mir gewandert... Er ist in Teilen verantwortlich fr die Ergebnisse meiner Arbeit (zumindest die guten). Herrn Prof. Dr. Tod Stuessy (Wien) danke ich fr die bernahme des Zweitgutachtens. Lieben Dank an Sabine Donner fr die stetige Hilfe in allen brokratischen Fragen und danke allen Mitarbeitern des Instituts fr Geographie fr die freundliche Atmosphre und die Mglichkeit 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 Lingenhhl, Henning Schrder, Tobias Bolch, Frieda Grninger und Tom Fickert. Klaus Geiselhart verdient einen speziellen Dank fr seine grundlegende Erfindung: den Geofanten. Auch Alex Brenning muss ich hier erwhnen, 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 Glck Euch kennenzulernen! Die sehr hilfreiche Englisch-Korrektur der Dissertation bernahm Iris Burchardt. Nochmals vielen Dank! Marian Jngling gehrt zwar nicht zur Geographie, doch war seine sprachliche Hilfe entscheidend fr den guten Verlauf des Rigorosums. Speziellen Dank an Thomas Sokoliuk fr seine langjhrige Untersttzung und seine unerschpfliche Feststimmung (danke fr den Fisch, Thomas!) Bei Rolf Kastner mchte ich mich unter anderem fr die berlassung seines Computers bedanken und bei Stefan Adler fr die groe Hilfe beim Einleben in Deutschland im ersten Jahr. Irma Richter, Anette Wel, Arietta Eberstadt sowie Coni Augustin sorgten auch fr das gute Eingewhnen und stets fr das seelische und leibliche Wohl. Bei Frau Dr. Brigitte Perlick, Akademisches Auslandsamt der Universitt, mchte ich mich fr ihre unendliche Energie bedanken und bei Frau Merker vom Promotionsbro fr den besten Rat in einer schwierigen Situation. Schlielich danke ich dem DAAD fr die gewhrte finanzielle Untersttzung. Besonders der zustndigen Referentin Maria Hartmann gebhrt fr die schnelle Lsung unlsbarer Probleme mein herzlicher Dank. Der Zantner-Busch Stiftung gilt auch mein Dank fr die finanzielle Untersttzung 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 Im alone responsible for the interpretation of these papers. Am Schluss mchte ich den Erlangern danken, die Stadt bot uns fast vier Jahre lang ein Zuhause. Und einen Ratschlag mchte ich zu guter Letzt geben: Nehmt alles nicht so ernst! Das Leben ist zu kurz dafr! Ich wnsche mir, Deutschland htte die WM 2006 doch gewonnen Die gute Stimmung htte vielleicht ein paar Jahre angedauert.



LEBENSLAUF

Persnliche Daten Name: Eltern: Geburtsdatum: Familienstand: Kinder:

Andrs Moreira-Muoz Sergio Moreira / Mlica Muoz-Schick . Feb. 97 in Los Angeles, Chile verheiratet mit Paola Soto aus Via del Mar, Chile Sayn (4), Silene (), Coyn (/4)

Schule 97 - 988 Abschlu: Studium 989 - 994 Abschlu:

Deutsche Schule, Santiago de Chile Prueba de Aptitud Acadmica (entspricht Abitur)

Studium der Geographie an der Universidad Catlica de Chile Diplom-Geograph. Thema der Diplomarbeit: Naturschutzgebiete an der Kste Zentral Chiles Studium der Angewandte Geographie an der Universidad Catlica de Chile Gegrafo Profesional. Thema der Abschluarbeit:: Umwelt-Erziehung im Natur-Reservat Yerba Loca, Andes Zentral Chile

994 - 99

Abschlu:

Promotion 00 - 007

Institut fr Geographie, Friedrich Alexander Universitt Erlangen - Nrnberg Thema: Plant Geography of Chile

Andrs Moreira-Muoz Erlangen, 4. Juni 007

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