Plant Geography of Chile

Transcription

Plant Geography of Chile
An Essay on Postmodern Biogeography
Den Naturwissenschaftlichen Fakultäten
der Friedrich-Alexander-Universität Erlangen-Nürnberg
zur
Erlangung des Doktorgrades
vorgelegt von
Andrés Moreira-Muñoz
aus Los Angeles, Chile
Gedruckt mit Unterstützung des Deutschen Akademischen Austauschdienstes (DAAD)
Als Dissertation genehmigt von den Naturwissenschaftlichen Fakultäten der
Universität Erlangen-Nürnberg
Tag der mündlichen Prüfung: Donnerstag 14. Juni 2007
Vorsitzender der
Promotioskommision:
Prof. Dr. Eberhard Bänsch
Erstberichterstatter:
Prof. Dr. Michael Richter
Zweitberichterstatter:
Prof. Dr. Tod Stuessy
A Paola, Silene, Coyán y Sayén
Contents
Zusammenfassung / Summary...............................................................................................8
1 Introduction........................................................................................................................... 11
1.1 Inventing plant geography............................................................................................. 13
1.2 Chorology and Arealkunde............................................................................................ 13
1.3 Naming the plant world................................................................................................. 19
1.4 From the map to the tree: cladistics and biogeography................................................. 24
1.5 The fragmented map of modern biogeography.............................................................. 27
1.6 Postmodern biogeography: de(re)constructing the map................................................ 29
2 Chile, a Remote Corner on Earth.......................................................................................... 37
2.1 Romancing the South: the discovery of a virgin world................................................. 37
2.2 Classification of the Chilean plants: a modern (but not definitive) synthesis................ 42
2.3 Geographical classification of the Chilean flora............................................................ 47
2.4 Excursus: vegetation maps and Vegetationsbilder......................................................... 60
2.5 Geographic ranges in the latitudinal profile................................................................... 63
3 Geographic Relationships of the Chilean Flora.................................................................... 71
3.1 Pantropical floristic element.......................................................................................... 72
3.2 Australasiatic floristic element....................................................................................... 73
3.3 Neotropical (American) floristic element...................................................................... 75
3.4 Antitropical floristic element.......................................................................................... 77
3.5 South-temperate floristic element.................................................................................. 80
3.6 Endemic floristic element.............................................................................................. 81
3.7 Cosmopolitan floristic element...................................................................................... 85
4 Biogeographic analysis......................................................................................................... 89
4.1. To be or not to be disjunct............................................................................................. 90
4.2 The austral v/s the neotropical floristic realm................................................................ 92
4.3 Analysis of Endemism................................................................................................... 95
4.4 The disintegration of the endemic and the south-temperate elements......................... 102
4.5 Plant geography of the Chilean Pacific islands............................................................ 106
5 Palaeogeography: insights into the Evolution of the Chilean Flora................................... 117
5.1 Continental movements: fragmenting the Earth’s surface........................................... 117
5.2 Oceanic transgressions over the continental surface................................................... 126
5.3 The Andean uplift........................................................................................................ 128
5.4 The last 30 000 years: surviving the Ice Age............................................................... 129
5.5 Alternative palaeogeographies: against scientific consensus....................................... 131
6 Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography................. 139
6.1 Molecular dating, the reigning paradigm..................................................................... 139
6.2. Phylogeny of Chilean plants....................................................................................... 141
6.3 Vicariance v/s dispersal in the Chilean Flora............................................................... 145
6.4 Sloppy biogeography v/s harsh geology?.................................................................... 149
6.5 Species and speciation................................................................................................. 151
6.6 Re-inventing an origin for the land plants................................................................... 157
7 Back to Postmodern Biogeography.................................................................................... 167
7.1 Biogeography as a social science................................................................................. 167
7.2 Biogeography toward a science of qualities................................................................ 178
8 Conclusions: toward a biogeographic synthesis of the Chilean Flora................................ 183
Footnotes................................................................................................................................ 186
References.............................................................................................................................. 188
Appendices............................................................................................................................. 221
Appendix A: List of Chilean genera, geographic distribution, floristic elements.............. 221
Appendix B: Genera shared by several Chilean regions................................................... 247
Appendix C: Matrix for PAE: Distribution of Endemic Genera........................................ 255
Appendix D: Native Genera in the Chilean Pacific Islands.............................................. 263
Agradecimientos / Danksagung / Acknowledgements.......................................................... 265
Lebenslauf.............................................................................................................................. 267
Zusammenfassung
Die historische Biogeographie hat sich in den letzten Jahrzehnten sowohl theoretisch als auch
methodisch stark weiterentwickelt. Konzepte wie Dispersion, Vikarianz oder Panbiogeographie
zeigen konstante Synergien im Zuge der Erneuerung von Theorie und Praxis. Die historische
Biogeographie muss sich dabei, wie alle Naturwissenschaften, der großen Herausforderung
stellen, sich der Moderne und der Postmoderne anzupassen. Die vorliegende Arbeit analysiert die
wichtigsten phytogeographischen Gegebenheiten in Chile unter besonderer Berücksichtigung der
geographischen Verbreitung von Gattungen und ihrer globalen Beziehungen. Die Auswirkungen
dieser Verhältnisse auf die Entwicklung der chilenischen Flora werden dabei besonders unter
verschiedenen paläogeographischen Szenarien diskutiert. Außerdem wird die Situation im
Kontext der modernen und postmodernen Wissenschaftstheorie betrachtet. Das erlaubt eine
Kontext-orientierte Synthese der chilenischen Pflanzengeographie und stellt den Ausgangspunkt
für weitere Forschungen in dieser Übergangsdisziplin zwischen Geographie und Biologie.
Schlagwörter: Pflanzengeographie, Panbiogeographie, Postmoderne, Floristische Elemente,
Chile.
Summary
Historical biogeography has seen a rapid theoretical and methodological developed in the last
decades. Concepts such as dispersal, vicariance, or panbiogeography show a constant synergy
in the renewal of the theory and the praxis. The present thesis examines the most important
phytogeographical issues in Chile with special consideration of the distribution and geographical
relationships of the genera. The consequences of these relations on the evolution of the Chilean
Flora are discussed thereby, particularly under different possible palaeogeographical scenarios. In
addition the conditions in the context of the modern and postmodern science theory are considered.
This permits a context-oriented synthesis of the Chilean plant geography, that set the starting
point for further research in this crucial field between geography and biology.
Keywords: Plant Geography, panbiogeography, postmodern science, floristic elements, Chile.
1 Introduction
From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden
Erlangen, March-December 2006
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1 Introduction
“Between geographers and historical biogeographers there has been relatively little
communication and, like Stoddart [1985], 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 1989).
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 100 years ago by Karl Reiche
(1907). The German botanist was appointed at the Museo Nacional de Historia Natural in 1896
by Federico Philippi, the son and scientific successor of the great naturalist Rudolf A. Philippi,
who arrived in Chile 1851 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 2), plus his incredible capacity, Reiche could publish his Flora de Chile in five volumes
(1896-1911) and his Plant Geography, this latter requested for the series Die Vegetation der Erde
by the German botanists Adolf Engler and Oscar Drude (Reiche 1907).
The Flora de Chile is under a new revision in a current effort lead by the Universidad de
Concepción, in a plan that comprises six volumes, from which the second has been almost
completed (Marticorena & Rodríguez 1995-2005). It seems to be also the time for a renewal of
Reiche’s Plant Geography. Not few things have changed in 100 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 1958, 1962). On the other side, the
biogeographic arena is getting more and more fragmented due to a plethora of methods (Ebach et
al. 2003), and the ultimate synthesis is getting more and more elusive (Lomolino & Heaney 2004).
Some speak about the crisis of biogeography (e.g. Riddle 2005), but this crisis is not restricted
to biogeography and seems to be more general, as the crisis of reductionistic modern science
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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 1st chapter sets the theoretical basis for the endeavour of doing plant geography in the 21th
century, in a constantly changing world (sensu Ebach & Tangney 2007). The 2nd 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 3rd 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 1st 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.
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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 [1989, 2001] 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 2003).
Plant geography, phytogeography, geobotany, chorology, Arealkunde. These concepts have been
used for two centuries in distinct and similar ways since Alexander von Humboldt’s Essai sur
la géographie des plantes (Humboldt 1805). The limits between these fields have never been
clear and it seems to be a waste of time delimiting them strictly. Augustin Pyramus de Candolle
(1820) called the subject just Géographie botanique and his son Alphonse de Candolle (1855)
added the adjective [Géographie botanique] raisonné. Later Schröter wrote about genetische
Pflanzengeographie oder Epiontologie (Schröter 1913), while Meusel preferred Vergleichende
Arealkunde or Geobotanik (Meusel 1943), and Walter added the adjective floristisch-historische
[Geobotanik] or reduced it to Arealkunde (Walter 1954, Walter & Straka 1970). Schmithüsen
proposed that geobotanists emphasize the plant and therefore should call their study plant
geography = Pflanzengeographie; geographers should have their emphasize in the physiognomy
of plant communities, i.e. the vegetation, and therefore should call it vegetation geography =
Vegetationsgeographie (Schmithüsen 1968). This proposal has not been followed by geographers
like Richter (1997) 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 1909), others about geographic botany (e.g. Raup 1942)1. For the present thesis I
took the original concept of A.P. de Candolle, simply traduced as plant geography. In this form I
can aside honour Stanley A. Cain’s great piece Foundations of plant geography (Cain 1944). 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)2 and spatiology; but perhaps the best term for the science
of area is areography, that portion of geography which deals especially with area” (Cain
1944, 147).
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1.2.1 Origin of chorology
The term chorology was introduced by Ernst Haeckel in the natural sciences (Friis 1998), from
the Greek word Chora (χώρα) = the residence, the distribution district.
„Unter Chorologie verstehen wir die gesammte Wissenschaft von der räumlichen
Verbreitung der Organismen, von ihrer geographischen und topographischen Ausdehnung
über die Erdoberfläche. […] Im weitesten Sinne gehört mithin die gesammte „Geographie
und Topographie der Thiere und Pflanzen“ hierher, sowie die Statistik der Organismen,
welche diese Verbreitungs-Verhältnisse mathematisch darstellt“ (Haeckel 1866, pp. 286287).
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 19th century (Drude 1890), later replaced by genetische
Pflanzengeographie (Irmscher 1922, 1929). Suessenguth (1938) and Vester (1940), 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 1938).
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
20th 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 (1927), although some accuse him, as having not yet
abandoned a descriptive science (see Holt-Jensen 1999). 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 60’s geography saw a real change, mainly due to the work of William Bunge (1962). 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 19th 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)3. 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 1.1 Distribution map of morphologic similar Gentiana species (section Endotricha), applying the
geographic-morphologic method (von Wettstein 1898)
theoretical systematics“ (Hannig & Winkler 1926-1940). 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 24 families, 258 genera and 1468 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 1943). Afterwards came
Walter’s Arealkunde (Walter 1954, 2. ed. by Walter & Straka 1970). 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
15
Figure 1.2 Donat‘s distribution maps in Hannig & Winkler (1926-1940)
arealkundliche school at Halle/Saale brought Arealkunde/Chorologie to its highest point, with
the work Vergleichende Chorologie der Zentraleuropäischen Flora, published in three volumes
(Meusel et al. 1965, 1978; Meusel & Jäger 1992). From the 50’s to 70’s we can mention also the
large mapping work of NW Europe from Swedish E. Hultén, and his Circumpolar plants (Hultén
1950, 1964, 1971). 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 1930, Milbradt 1976, Welß & Lindacher
1994).
The earlier maps were an auxiliary tool for solving more general problems about the relationships
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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 (1962), geobotanists are more concerned with dot/grid methods
„The influence of this first national atlas following the dot/grid method led in Europe to a true
renaissance of floristic mapping and consequently to an indeterminable number of atlases at the
national, regional and local scale” (Haeupler 2005). Examples are the floristic atlas of Germany
(Haeupler & Schönfelder 1988), of Bavaria (Schönfelder & Bresinsky 1990), or the Flora of the
Regnitz area (Gatterer & Nezadal 2003). 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. (1998) as successors of F. White and
its extensive work on Africa (White 1971, 1993); or Cope (2000) as the last representative of the
chorological work with grasses at Kew. Already in the preface of his work Meusel (1943) realized
“…During the boom that affected plant-geographical research in the last decades, chorology stayed
remarkably behind vegetation science (phytosociology). This may be partially connected with the
fact that the methods of modern ecology are better suited for vegetation analysis than for floristic
problems”. Afterwards the vegetation science developed much more strongly, by integration of
concepts of the ecology (e.g Clements), the Zürich-Montpellier school (Braun-Blanquet) and
phytosociology (Tüxen). This was also recognized by Rothmaler: “…Working directions like the
ecology detached themselves as independent branches of science, so that the original (floristic)
plant geography went adrift. This however stands in close connection with taxonomy; a complete
separation of both spheres of activity is not possible” (Rothmaler 1955). 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 (1944) 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. 2006). To areography belong the analysis of geographic ranges (e.g. Gaston
2003) and ecogeographic rules, like Bergmann’s rule, Allen’s rule, and perhaps the most famous
Rapoport’s rule, about latitudinal richness gradients. The Argentine ecologist E. Rapoport can
certainly be recognized as the re-finder of areography (Rapoport 1982), although he most surely
borrowed the term (not the meaning) from Cain (1944). 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 1995, Blackburn & Gaston 2003). Indeed Rapoport’s rule is still
17
constantly examined and discussed in ecological and biogeographical journals 20 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 2005, Lomolino et al. 2006). 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 1997, Schroeder 1998, Frey & Lösch 2004). 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 2007, p. 29). “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 1926-1940). In this sense
chorology is in the line of Parenti and Ebach’s comparative biogeography, and is closer attached
to systematics (Rothmaler 1955, Stuessy et al. 2003, Parenti & Ebach in prep.). Chorology and
Arealkunde have trespassed their essence to current more integrative approaches, like Croizat’s
panbiogeography (Croizat 1958). 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 (2006) and
Cullen & Walters (2006), as resumed from earlier studies, e.g. Croizat (1945), Lawrence (1951),
Davis & Heywood (1963), Griffiths (1974), Morton (1981), Walters (1986), and Stevens (1994).
Integrating Stuessy’s and Cullen & Walters’ views we can identify seven phases in the history of
plant taxonomy (table 1.1.)
Table 1.1 Phases in the history of plant taxonomy From Stuessy (2006) and Cullen & Walters (2006).
Phase
Period
Issues
1
300 BC - 1460
Pioneer Greek Theophrastus wrote several manuscripts dealing with
plants, translated as Enquiry into Plants (Hort 1916, quoted by Stuessy
2006).
2
c. 1460-1660
Age of the Herbalists, practical knowledge against diseases
3
1660 - 1753
Classification per se, searching for revealing ‘God’s system of
classification used at the Creation’, i.e. as artificial systems, (e.g.
Caesalpino, Bauhin, Ray de Tourneffort, culminating in the system of
Linnaeus’ Species Plantarum of 1753.
4
1770-1880
The search for natural systems (e.g. Jussie’s Genera Plantarum, de
Candolles’ Prodromus).
5
1880-1950
Phylogenetic systematics, mainly enforced by Darwin’s Origin of
Species (1859). 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 2006) (e.g.
systems of Engler & Prantl, Hutchinson, Takhtajan, Cronquist).
6
1950-1980
Phenetic studies avoiding phylogenetic speculation and returning to a
‘more natural’ approach, sometimes using numerical methods
7
1980 onwards
Resurgence of phylogenetics mainly based on cladistics and molecular
information.
Cullen & Walters (2006) 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
19
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 2006, 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 2006, p. 34). This question lies on the basics of systematics and have been largely
debated, mostly by philosophers of science (e.g. Endersby 2001, Hull 2001). The early words of
the German philosopher E. Cassirer are still appropriate to discuss the subject. Cassirer speaks
about “the naïve view of the world”. According to his view “systematics consists of grouping
things according to their similarities. Our concepts of groups or classes are supposed to arise by
our distinguishing the similarities of things from their differences; as we ascend in abstracting
thought to ever more inclusive classes, the similarities which we recognize become ever fewer,
until I suppose they eventually disappear. From an ontological viewpoint this kind of thinking must
be judged naïve, because it involves presuming that the similarities (common attributes) of things
have a simple hierarchical distribution. But there is in nature no unique hierarchy of similarities,
as can be seen from the following example taken from Popper [1959, p. 421]” (Cassirer 1923, as
quoted by Griffiths 1974, p. 88). Popper’s 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 states5 under evaluation, or in providing a priori a phylogenetic framework to
organize the similarities. As shown in table 1.1, phylogenetic systematics arose only after the
general acceptance of Darwin’s Origin of Species, and the consequent changes in biological
theory.
Woodger (1952) explained this change in biological theory: “In the Linnean system of classification
20
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
(2006) recognizes three different schools of systematics that developed over the past 40 years:
phenetics (end 1950s into the 1960s), cladistics (1970s-1990s) and phyletics (2000 onwards).
In the following the emphasis will be put on some aspects of phenetics and cladistics (figure
1.4), being the latter the current dominating paradigm in systematics since the translation of Willi
Hennig’s Grundzüge einer Theorie der phylogenetischen Systematik into English (Hennig 1950,
1966).
Figure 1.4. David Maddison‘s graphical representation of the debate between cladistics and phenetics during
the 80‘s. [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 1963)”.
These latter authors already knew Hennig’s work, but they were not impressed by the various rules
Hennig set out for inferring phylogeny, nor by Hennig’s methods of reciprocal illumination. They
found them too liable to produce error, like reasoning in a circle, “the piling of hypotheses upon
hypotheses … Their goal was to develop clear, logical, straight-line methods of classification.
First a systematist must do A, then B, and only then C – no circles, no spirals, no back-tracking.”
(Hull 2001, p. 230).
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 2006).
21
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 1.1 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/gloss1phylo.html
Understanding evolution: http://evolution.berkeley.edu/evolibrary/article/phylogenetics_02
The methods proposed by Hennig (1950, 1966) 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 (2006), in three a priori axioms:
a) monophyly: principle that any classificatory group should be monophyletic, in contrast to
polyphyletic or paraphyletic (box 1.1);
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.
22
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 2001, p. 236).
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 (1998, 2003) 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” 6
The rise of cladistics came to a peak in 1998, when the new angiosperm classification (APG
1998) 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 2001).
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 1998). 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;
23
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 (2001) 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 1999, quoted by Endersby 2001).
“…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 1999, quoted by Endersby 2001).
“…we didn’t want it to be cited as ‘somebody et al’, because it did represent the work
of so many different people in different places… we wished to stress the fact that this
wasn’t a system of classification done by an expert - that is somebody like a Cronquist,
or a Takhtajan, or Thorne ... We wanted to explicitly say that botanists had gone beyond
the need for authorities in this way... (Chase 1999, quoted by Endersby 2001).
But APG’s effort just changed the authority of somebody toward the authority of the group, the
consensus, valid only for insiders and not for outsiders of alternative taxonomy. However, where
the supposedly objectivity stayed is a question that still remains (see further discussion in section
1.6.1).
1.4 From the map to the tree: cladistics and biogeography
„Ich setzte also im Erwachsenenalter die Spielereinen meiner Kindheit mit Karten
fort: ich verband Städte gleicher Größe durch gerade Linien, einmal, um festzustellen,
ob im Eisenbahn- oder Straßennetz gewisse Regeln erkennbar seien, ob es regelhafte
Verkehrsnetze gäbe, zum anderen, um die Abstände zwischen gleich großen Städten zu
messen. Dabei füllten sich meine Karten mit Dreiecken, oft gleichseitigen Dreiecken
– die Abstände gleich großer Städte untereinander waren also annährend gleich --, die
sich zu Sechsecken zusammenschlossen. Ich stellte weiter fest, daß in Süddeutschland
die kleinen Landstädte sehr oft sehr genau einen Abstand von 21 km voneinander haben.
Mein Ziel war abgesteckt: Gesetze zu finden, nach denen Anzahl, Größe und Verteilung
der Städte bestimmt sind...“ (Christaller 1968, p. 96) 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 20th 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 19th century by Lamarck & de Candolle (Ebach & Goujet 2006) and F. Schouw
24
(Mennema 1985) (see sections 1.2 and 2.3). 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 1986).
Box 1.2 Basics of cladistic biogeography
C
A
B
C
A
D
A
B
C
B
D
A
C
D
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-
25
1.4.1 Vicariance biogeography v/s dispersalism
Gareth Nelson and Norman Platnick developed this approach based on the work of León Croizat
(Nelson & Platnick 1981), but vicariance as an explanation of disjunct distributions is already
found in the early works of J.D. Hooker (Turrill 1953), and in Ratzel’s biogeography (Ratzel
1901). But more specifically, as Nelson and Platnick recognized, their views had grown out of the
work of Hennig, Croizat and Karl R. Popper. “Although they [Nelson & Platnik] admitted that
Croizat found Hennig’s principles of phylogenetic systematics antithetical to his own principles
of panbiogeography, and that neither Hennig nor Croizat had ever mentioned Popper (and vice
versa), nevertheless Nelson and Platnick considered their own work as synthesizing and extending
the views of these three authorities” (Hull 2001, p. 234). They extendedly discussed the science
of branching diagrams - cladistics in its most general sense (box 1.2). “It was primarily through
the championing of Hennig’s principles of biogeography, first by Lars Brundin of the Swedish
Museum of Natural History and then by a young American ichthyologist, Gareth Nelson, that
Hennig finally gained some recognition among English-speaking biologists” (Hull 2001, p.
231).
Nelson and Platnick’s approach was further extended by other authors that found in this approach
a real changing paradigm from the Darwinian dispersalist biogeography. Dispersalism long
dominated the biogeographic scene since the publication of The Origin of Species, and was further
developed by the New York School of biogeography, in the figures of W.D. Matthew (1871–1930),
K.P. Schmidt (1890–1957), G.G. Simpson (1902–1984), P.J. Darlington, Jr (1904–1983) and G.S.
Myers (1905–1985), (reviewed by Nelson & Ladiges 2001).
Dispersalist biogeography is based on the assumption that taxa originated in relatively small areas
(centres of origin) and therefore tries to reconstruct the routes that organisms covered to colonize
known past or present ranges. A good example of this view is Simpson’s Splendid Isolation: The
Curious History of South American Mammals (Simpson 1980). In fact, Croizat called dispersalism
“the science of the curious, the mysterious, the improbable” (Croizat 1958).
Many authors saw the problems in the search for the centre of origin and migration routes, like
botanist Stanley Cain (1902-1995), who analized Adams’ 13 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 1944, p. 185).
After almost 150 years of dispersalism, vicariance biogeography was taken by biogeographers as
the new paradigm, and most of the literature in the 80’s and 90’s was engaged with the analysis
of area cladograms.
But dispersalism was only taken a rest, and returned like the Fenix, with the new methods of
molecular dating, as well expressed in the apologia of de Queiroz (2005), and recent papers of
26
emphatic advocates (e.g. Renner 2005, McGlone 2005) (see the discussion in chapter 6). An
interesting point expressed by Grehan (2007) regarding current biogeographic studies is in direct
relationship to Christaller’s quotation at the beginning of this section: “To discern the geographic
context of evolution, Darwinian biogeographers look to historical theories of ecology, systematics,
molecular clocks, and geology – anything but distributions themselves – as the empirical data of
biogeography… One only has to see how often distribution maps are left out of the picture, even
for papers focusing on biogeographic theory and method” (Grehan 2007, pp. 83-84).
Table 1.2 Adams’ 13 criteria for centres of origin, as revised by Cain (1944, pp. 185-211).
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
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
13. 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 1966, p. 130-131) 8.
27
a
b
Figure 1.5 Hennig‘s fragmented map as test of the criterion of veracity (Hennig 1966)
In section 1.3 cladistics was briefly introduced, therefore Hennig’s metaphor serves here more
as an excuse to discuss the conflicting view in the current fragmented biogeographical scene. As
stated by Riddle (2005 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. 2003 and Morrone 2005) leave no
opportunity for this integration. Nelson & Ladiges (2001) characterized current biogeography as
a “mess of methods”.
Critics come and go between the exponents of different approaches: Ebach & Humphries (2002)
and Ebach et al. (2003) 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. [2003] is no more than a repeat of the pretectonic ideology that looks for centres of origin and direct lineages” (Ebach et al. 2003). 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. 2003). Furthermore, Donoghue &
Moore (2003) propose that cladistic biogeography has failed to become a truly productive research
programm because its scope has been fatally over-simplified by being restricted to assessing the
topological congruence of phylogenetic trees as depicted by general area cladograms. But Parenti
& Ebach (in prep.) reject Donoghue & Moore’s integrative biogeography program because in
their view it has been used to reinforce equivocal divisions, such as that between phylogenetically
and ecologically focused methods.
“In our view, biogeography is one field with one aim – to explain the distribution of taxa today
as elements of form, time and space regardless of whether the data is morphological or molecular
and whether taxa are represented as species, sub-species or populations…The dismissal of Earth
process from historical and ecological biogeography may have a sociological, political and
perhaps an emotional raison d’etre. Whatever the reason, the rationale is unclear and alienating
to non-biological disciplines. There is no reason to doubt the role of geological process, namely
tectonics as the primary cause for geographical and environmental composition (soils, water)
28
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 2003).
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 (2004) 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 1800s 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 2004).
Lomolino & Heaney (2004) continue with their optimism: “the revitalization will continue in
earnest, largely through the efforts of broad-thinking scientists who no longer shy away from but
embrace the complexity of nature, and who foster collaborations and conceptual reticulations in
modern biogeography”.
1.6 Postmodern biogeography: de(re)constructing the map
„Vivimos tiempos en que constantemente el postmodernismo confronta el discurso
hegemónico de la ciencia para denunciar sus excesos y exponer sus límites“ (Morrone
2003, p. 87) 9
Lomolino & Heaney (2004) 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 (2004) 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 2006).
29
Furthermore, in proposing to deconstruct current approaches and promoting creative applications,
Lomolino & Heaney are going beyond from modern reductionistic science and enter the complex
and more unpredictable region of postmodern science. In the search of the grand view, the ultimate
synthesis across space and time we come unavoidably to León Croizat’s analytical and synthetical
biogeography: Panbiogeography (Croizat 1960). In fact, Croizat’s catch-phrase the earth and the
biota evolve together has become one of the most popular slogans in the history of biogeography.
Croizat expressed this explicitly in dozens of articles and several books like Space, time, form,
the biological synthesis (Coizat 1962) (see Heads & Craw 1984 for a complete bibliography).
Croizat’s synthetical view is what biogeographers have been searching since the beginning of the
science as a modern task. Croizat, while opposing the dominant dispersalism of his time, is one of
the major responsible of the revitalization, quite possible a scientific revolution, that biogeography
is today experiencing (Lomolino & Heaney 2004). The deconstruction suggested by Lomolino &
Heaney (2004) is in tune with the revision that is today occurring in many sciences, specially the
social sciences. The critical view of postmodernism left nothing untouched and our world view is
changing more rapidly as many like to recognize it.
1.6.1 Objectivity and postmodern science
The philosopher Frederick Ferré wrote a challenging essay for the mainstream book Principles
of Conservation Biology (Meffe & Carroll 1997) about the limitations of reductionist modern
science in responding to the real world problems. Only postmodern science would “nurture the
human hunger for quality: for beauty, balance, creative advance” (Ferré 1997). The reaction of
Cotterill (1999) for such a naïve view was drastic: “postmodernism has little, if anything, to offer
these nor any other science – especially where conservation biology is tasked with informing
and changing policy… this uncritical sanction in a mainstream textbook on conservation biology
kindles suspicions that a deeper problem afflicts the discipline’s scientific integrity” (Cotterill
1999). Ferré’s essay got the sanction, and disappeared from the 3d edition of the book (Groom et al.
2006). 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 2000). 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 (2000) 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
30
described as fashionable salon philosophy by Dawkins [1995], who questions the entire credibility
of the postmodernist programme [Dawkins 1998]. In its extreme manifestations, postmodernism
is the antithesis of science [Wilson 1998]” (Attwell & Cotterill 2000, p. 560).
Remarkably, Attwell & Cotterill (2000) 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 2000, p. 560).
1.6.2 Postmodernism as the villain?
From the same tribune that holds Attwell & Cotterill’s (2000) 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 1995, p. 787).
Howarth (1995) 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 [1962] 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 1962, as quoted by Howarth 1995, 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 concerns10. Chorley expressed it with an excellent aphorism applied
to the theory of geomorphology:
31
“Whenever anyone mentions theory to a geomorphologist, he instinctively reaches for
his soil auger” (Chorley 1978)
But even in such hard disciplines like geomorphology, the philosophical analysis to strengthen
its intellectual foundation is on its course (e.g. Chorley 1978, Osterkamp & Hupp 1996, Rhoads
& Thorn 1996).
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 1988, quoted by Heads 1990). 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. 1999). More recently Heads
(2005a) 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 (1967) 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 2004). Brown & Lomolino
(2000) 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. (2004) 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.
32
“For history is like a nymph glimpsed bathing between leaves: the more you
shift perspective, the more is revealed. If you want to see her whole you have to
dodge and slip between many different viewpoints. This technique, which ought
to commend itself to historians, might also restore to subjectivists a prospect of
objectivity. Even the most dedicated subjectivist should be able to imagine what it
would like: objectivity would be the result of compiling or combining all possible
subjective points of view. Every time we take notice of each other, therefore, we get
a little closer to truth. Those who refuse to acknowledge anyone’s existence save
their own must approach truth by imagining a variety of perspectives. To see things
from no point of view is not even theoretically possible. If we try to see from every
point of view, we shall never attain our goal, but it is at least meaningful to speak
of seeing from every point of view, whereas it is literal nonsense to speak of seeing
from none.” (Fernández-Armesto 1997, Truth, p 228).
Following Fernández-Armesto’s idea, along the thesis the plant geography of Chile will be
analized from different point of views: from the traditional views to the alternative ones. The
effort has to be seen still as a work in progress, since:
“...Toda división fitogeográfica deberá, entre tanto, considerarse como provisional
o aproximada. Podrán expresarse opiniones, pero discutir divisiones, límites o
nomenclatura, o tratar de imponer ideas en forma dogmática, será solo gastar tiempo
y papel sin mayor provecho para la ciencia.” (Cabrera 1953, as quoted by Ribichich
2002)11
33
34
2 Chile, a Remote Corner on Earth
35
From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen,
March-December 2006
36
2 Chile, a Remote Corner on Earth
“Durante la Conquista, cuando los españoles recorrían América buscando esos metales
[plata y oro] y llevándose todo lo que encontraban al paso, Chile se consideraba el culo
del mundo, porque comparado con las riquezas del resto del continente tenía muy poco
que ofrecer” (Isabel Allende, Hija de la Fortuna, 1999)
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 Sebastián Elcano. A detailed relate of early collectors, botanists,
and naturalists that contributed to the knowledge of the Chilean flora has been done by Reiche
(1907) and Marticorena (1995) (there is also a complete bibliography in Marticorena 1992, and
an online summary by Muñoz-Schick & Moreira-Muñoz 2002). Some highlights in the discovery
of the Chilean plant world have been briefly resumed in box 2.1.
Box 2.1 The discovery of the Chilean plant world
The discovery of the Strait of Magellan by Hernando de Magallanes and his crew (21 october 1520) set the start point for the
European exploration of South America. Magallanes gave the
name ‘Cabo de Las Vírgenes’ to the eastern Cape that allowed
them to get into the Strait. During the first days of exploration,
the observations are about a native wood to make fire which
smoke smell well. They refer undoubtedly to the wood of ‘canelo’, Drimys winteri. The sailors early noted the properties of
canelo’s 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.
Hipólito Ruiz and José Pavón did the first specific botanical excursion to
Chile. With the purpose of extending the collections of the Real Botanical
Garden of Madrid, king Carlos III of Spain sent several naturalist expeditions
to different countries; Peru and Chile corresponded to Ruiz and Pavón. They
made collections in some points between Talcahuano and Santiago between
1782 and 1783. The work they published later in 3 volumes has more than
300 illustrations. Here as example‚ the ‚copihue‘, Lapageria rosea.
37
Box 2.1 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 (1768), Molina published in Italy his Compendio della storia geografica, naturale, e civile del Regno del Chile (Molina 1776). This
and coming works (Molina 1782, 1810) were for a long time the
main source of knowledge on natural sciences of Chile. The illustration represents araucaria, the Chilean palm (Jubaea) and ‚culén‘
(Otholobium glandulosum).
In 1827 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 3 volumes
and 300 illustrations (written with S. Endlicher), was accompanied for the first time by
coloured drawings. The night from 9. to 10.
January 1827 Poeppig crossed for the first
time the equator aboard the ship ‘Gulnare’.
“Leicht war die Stunde, in der wir aus der nördlichen Halbkugel schieden, einer der feierlichten und bedeutsamsten eines langen Reiselebens
… Fast will es dem Reisenden dünken als trete er mir dem Eintritte in
eine neue Welt auch in ein neues Leben…“. (after Morawetz & Rösser
1998). The illustration corresponds to Puya alpestris.
The English naturalist Joseph Dalton Hooker (1817-1911) accompanied the expedition to the South Pole of John Clark Ross
(1839 - 1843). He was the first who collected intensively in the
southern lands (Kerguelen, 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 1844-1860.
38
The great Charles Darwin (1809-1882), same as J.D. Hooker, was very young
when he traveled around the globe aboard the Beagle between 1831-1836. His
impressions abouth the southern biota and geology were fundamental for the
later development of the theory of evolution. He traveled in Chile between 18341835 and collected almost 1.500 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 1839).
After its Independence from the Spanish dominion in 1810, the young Republic of Chile had to better know its
natural resources. The French professor of natural sciences, Claude Gay
(1800-1873), already living in Chile,
was contracted by the government to
do the first intensive scientific exploration of the territory. He intensively
traveled between Copiapó and Chiloé
(1830 - 1842) (Muñoz Pizarro 1944).
Afterwards he published his masterpiece Historia Física y Política
de Chile, which consists of 28 volumes (8 of botany) and 2 illustration atlases (one cultural and one physical) (Gay 1845-1854). 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 300 species and in his
work 3767 new species are described. Illustration of Desfontainia
spinosa (source: www.memoriachilena.cl)
Rudolf Amandus Philippi (1808-1904) had to leave Germany due to political
problems and arrived in Chile in 1851. He is recognized as the main naturalist
in the history of Chilean science (Castro et al. 2005), having done studies in
botany, zoology, geology, paleotology, etnology, etc. He gave a big impulse
to the Museo Nacional de Historia Natural.
F. Philippi
R.A. Philippi got soon the collaboration of his son
Federico Philippi (1838-1910), 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 3767 species described to that
date rises to more that 7497 species. Of the 3730 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.
39
Karl Reiche (1860-1929), a German professor of natural sciences, was
appointed as encharged of botany in 1902 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 6 volumes
(Reiche 1896-1911). He also wrote the first plant geography of Chile
(Gründzuge der Pflanzenverbreitung in Chile, 1907, translated to Spanish by Gualterio Looser as Geografía Botánica de Chile).
Federico Johow (1859-1933) arrived in Chile 1889 to teach natural sciences at the Instituto Pedagógico, Instituto Nacional and Universidad de
Chile. He is the author of the milestone Estudios sobre la flora de las Islas
de Juan Fernández, still an obligate reference for the islands (Johow 1896).
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 (1879-1934) was incharged of the National Herbarium
in 1911, as Karl Reiche leaved the country and accepted an appointment in
Mexico. Fuentes made the first study on plants of the Easter Island (Fuentes
1913) and on Monocots that had not been treated in Reiche‘s Flora.
Ivan Murray Johnston (1898-1960), US American prominent botanist, studied the coastal desert flora and vegetation. His papers „The coastal flora
of the departments of Chañaral and Taltal“, and „The flora of the Nitrate
Coast“ (1929 a,b), are considered classical studies of the northern Chilean
coast, where vegetation is so highly related to sporadic precipitacion due
to El Niño events (Dillon et al. 2003) and the fog‘s water content (MuñozSchick et al. 2001).
Gualterio Looser (1898-1982) showed a wide interest in the natural sciences,
having been appointed as chief of anthropology at the National Museum. Since
a journey to Juan Fernández in 1925 he demonstrated more interest in botany,
publishing till 1971 abouth 140 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
Carl Skottsberg (1880-1963), the Swedish botanical eminence, president of the 7th International Botanical Congress in 1950, is one
of the most impressive botanists that explored the Chilean territory.
He came first with the Swedish expedition to the South Pole between 1901 and 1903 and afterwars explored several times mainly
Patagonia and the Pacific islands. He published about 128 articles
and several books about the Chilean botany and plant geography.
His Vegetationsverhältnisse längs der Cordillera de los Andes südlich von 41 Grad... (1916) and his Natural History of Juan Fernandez and Easter Island (1920-1956) are till today the most complete
works that treat these territories.
Carlos Muñoz Pizarro (1913-1976), 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 1942. He initiated a program of organisation of the collections.
Around 30,700 exemplars were revised, catalogued and organized. In 1961, with a grant from the
Rockefeller Foundation, Muñoz Pizarro explored, together with his wife Ruth Schick, the principal
European herbaria, carrying out the photographic registry of the type collections of Chilean plants.
The acquired knowledge allowed him to publish several important books: Sinopsis de la Flora Chilena (1959, 2d ed. 1966), the principal synthetical work of the Chilean flora still in use; Flores Silvestres de Chile (1966); Chile: Plantas en extinción (1973).
Mutisia decurrens, from Sinopsis de la Flora Chilena (1966)
41
2.2 Classification of the Chilean plants: a modern (but not definitive)
synthesis
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“
Tr
a
Monilophytes = ferns
Spermatophytes
Lycophytes
Tracheophytes
Euphyllophytes
Keeping in mind the discussion in section 1.2, there are several classification systems in which we
can organize the Chilean Flora. Taxonomy is mainly a modern European task, transferred to the
rest of the world since Linnaeus invented the binomial classification. There are also traces of an
aboriginal taxonomy, and Villagrán (2003) suggests to recognize this millenary task as a ciencia
indígena. But other languages dominate the sciences, and most of the aboriginal knowledge went
lost con la cruz y la espada.
The system used in the present thesis for organizing and analysing the Chilean families and genera
is the one proposed by Pryer et al. (2004) for the vascular plants (figure 2.1) and specifically the
APG system in its 2d version (APG II 2003) for the angiosperms (figure 2.2), with further updates,
as continuously tracked by Stevens (2001 onwards).
Leptosporangiate ferns
x
x
x
x
Horsetails
x
x
x
x
Marattioid ferns
x
x
x
x
x
Ophioglossoid ferns
x
x
x
x
x
Wisk ferns
x
x
x
x
x
Gnetophytes
x
x
x
Conifers
x
x
x
Ginkgo
x
x
x
Cycads
x
x
x
Angiosperms
x
x
x
Quillworts
x
x
x
x
Spikemosses
x
x
x
x
Clubmosses
x
x
x
x
Monophyletic
x
x
x
x
Paraphyletic
Figure 2.1 Cladogram showing relationships among the major lineages of vascular plants (from Pryer et al.
2004)
2.2.1 Floristic composition: major families and genera
Reiche’s (1907) statistics were composed of 141 families and 716 genera for the Chilean Flora
(table 2.1), as he critically revised previous Philippi’s account of 863 genera, as preparing the new
Flora de Chile (Reiche 1896-1911). Cullen & Walters (2006) recognize a trend of increase in the
number of families in different classifications in time, and in fact the number of genera and families
has grown since Reiche’s revision. Muñoz Pizarro (1966) in his Sinopsis de la Flora Chilena gave
a number of 182 families and 960 native genera13, and Marticorena’s (1990) checklist considered
175 families an 827 genera (table 2.1).
42
Amborellaceae
Nymphaeaceae
angiosperms
Austrobaileyales
Chloranthaceae
monocots
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
eudicots
Gunnerales
1/1
Berberidopsidales 2/2
Caryophyllales
12/69
3/9
4/10
+ Vitales 1/1
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
rosids
core eudicots
Santalales
Saxifragales
asterids
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
eurosids I
+ Zygophyllales 2/7
eurosids II
euasterids I
+ unplaced Boraginaceae /13
+ unplaced Icacinaceae /1
euasterids II
+ unplaced Desfontainiaceae /1
+ unplaced Escalloniaceae /2
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 2.1 Historical statistics for the Chilean native vascular plants
F. Philippi 1881
Reiche 1907
Muñoz Pizarro 1966
Marticorena 1990
Orders
91
-
Families
141
182
Genera
863
716
960*
175
827
* 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. 2005). Several genera should join the new proposed families Gratiolaceae or Linderniaceae
(Rahmanzadeh et al. 2005). On the other side, also former Scrophulariaceae have been erected to
43
family status, i.e. Calceolariaceae (Olmstead 2001). Chenopodiaceae are now being submerged
into Amaranthaceae; Empetraceae and Epacridaceae into Ericaceae (Kron 1996, Kron & Luteyn
2005).
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 1990). 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 2.3.
Table 2.2 Summary of Chilean plants (updated 31 December 2006)
ferns and fern allies
gymnosperms
monocots
dicots
TOTAL
Orders
Families
18
2
6
33
59
24
4
30
121
179
Genera
50
9
163
591
813
Native species
121
16
880
3,316
4,333
Table 2.3 Genera not found in continental Chile (* = family also absent from the continent)
Juan Fernández Islands
gymnosperms
Genus (family)
Dicksonia (Dicksoniaceae)*
Thyrsopteris (Dicksoniaceae)*
Arthropteris (Oleandraceae)*
-
monocots
Juania (Arecaceae)
Machaerina (Cyperaceae)
Megalachne (Poaceae)
Podophorus (Poaceae)
dicots
Centaurodendron (Asteraceae)
Dendroseris (Asteraceae)
Robinsonia (Asteraceae)
Yunquea (Asteraceae)
Selkirkia (Boraginaceae)
Haloragis (Haloragaceae)
Cuminia (Lamiaceae)
Lactoris (Lactoridaceae)*
Zanthoxylum = Fagara (Rutaceae)
Coprosma (Rubiaceae)
x Margyracaena (Rosaceae)
Santalum (Santalaceae)
Boehmeria (Urticaceae)
44
Desventuradas Islands
gymnosperms
monocots
dicots
Lycapsus (Asteraceae)
Thamnoseris (Asteraceae)
Nesocaryum (Boraginaceae)
Sanctambrosia (Caryophyllaceae)
Isla de Pascua
Davallia (Davalliaceae)*
Microlepia (Dennstaedtiaceae)
Diplazium (Dryopteridaceae)
Dryopteris (Dryopteridaceae)
Microsorum (Polypodiaceae)
Psilotum (Psilotaceae)*
Vittaria (Vittariaceae)*
Doodia (Blechnaceae)
gymnosperms
monocots
Kyllinga (Cyperaceae)
Pycreus (Cyperaceae)
Axonopus (Poaceae)
Stipa (Poaceae)
dicots
Triumfetta (Malvaceae)
Ipomoea (Convolvulaceae)
In the angiosperms, richest Chilean orders are the Lamiales (15 families, 58 genera), Caryophyllales
(12/69), Asparagales (10/38), Malpighiales (10/20), Poales (7/102), Alismatales (7/15), Ericales
(6/17) and the Asterales (5/137) (figure 2.2).
The Asteraceae, usually the largest family in floras of arid or semi-arid regions (Goldblatt &
Manning 2000), is also the most species- and generic-rich family in the Chilean flora (table 2.4).
The second largest family is the Poaceae, what is also expected due to the global high richness
of the family. Apiaceae, Brassicaceae, Fabaceae and Solanaceae follow in size, with Fabaceae
behind Poaceae in species numbers.
Table 2.4 Ranking of the 20 largest families in the Chilean flora by size
FAM
Asteraceae
Poaceae
Apiaceae
Brassicaceae
Fabaceae
Solanaceae
Caryophyllaceae
Cactaceae
Cyperaceae
Boraginaceae
Rosaceae
Verbenaceae
Malvaceae
Alliaceae
Myrtaceae
Amaranthaceae
Iridaceae
Campanulaceae
Lamiaceae
Plantaginaceae
N° genera N° Chilean species
121
67
24
23
22
22
18
16
16
13
12
11
11
9
9
9
9
8
8
8
45
863
392
93
156
275
164
66
96
146
107
42
83
73
39
23
47
38
17
29
43
The most species-rich genera are Senecio, Adesmia, Oxalis, Viola, Haplopappus, Poa, Carex,
Solanum, Calceolaria, Berberis (table 2.5). Some of these genera are cosmopolitan species-rich
genera (e.g. Senecio, Oxalis, Viola), while others are strict neotropical genera (e.g Haploppapus,
Calceolaria). Adesmia is a very interesting case, being the most species-rich genus restricted to
the southern Andes of Chile/Argentina (with some representation in Perú and southern Amazonas)
(a recent biogeographic analysis of Adesmia has been done by Mihoc et al. 2006).
Table 2.5 Ranking of the 20 largest genera in the Chilean flora (including Hypochaeris, Alstroemeria,
Verbena each with 33 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
218
132
117
70
64
61
55
51
50
47
46
43
43
43
41
39
37
36
36
33
2.2.2 Endemic families
The Chilean flora comprises three monotypic or dytipic endemic families, Gomortegaceae,
Francoaceae, and Lactoridaceae, this latter endemic to Juan Fernández (table 2.6). Also there are
four subendemic families restricted to Chile and adjacent territories in Argentina and Peru (table
2.7). The Chilean flora harbours 83 endemic genera, that will be further analysed in chapter 3.6.
Table 2.6. Endemic families of the Chilean flora
Family
Genera
N° species
Gomortegaceae
Francoaceae
Gomortega
Francoa
Tetilla
Lactoris
1
1
1
1 (Juan Fernández)
Lactoridaceae
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)
1 (1)
8 (8)
1 (1)
1 (1)
24 (18)
46
Just as a brief comparison, Argentina harbours one endemic family (Halophytaceae) and 45
endemic genera (Zuloaga et al. 1999). The flora of Perú, composed of more than 17.000 species,
and around 2.400 native genera14, has 51 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 23 endemic genera and also none endemic family (Jørgensen & León-Yáñez 1999).
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 5, 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. 5: 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 2001),
others have shown that the biogeographical hierarchies are real (McLoughlin 1992, 1994).
Hereafter I present a summary of the different ways in which the Chilean flora has been classified
at the global level (table 2.8).
Table 2.8 The geographical classification of the Chilean flora
N°
kingdoms
N°
regions
G.R. Treviranus 1803
8 ‘Flor’
-
Antarktische Flor
August Pyramus de
Candolle 1820
Schouw 1823
-
20
le Chili and les terres Magellaniques
25
-
Alphonse de Candolle
1835
Grisebach 1872
-
45
-
24
Engler 1879-1882
4
32
Drude 1884
14
55
Drude 1890
Diels 1908
Good 1947 (1974)
14
6
6
55
7
37
Mattick 1964
6
43
South of 42° = Antarktisches Florenreich; 42° - 23° = Reich der
Holzartigen Synanthereen (Compositae); North of 23° = Reich
der Cactus und Piper
Region 35, Le Chili, and region 36, la Patagonie, la terre de
Feu et les iles Malouines (Falkland).
Tropische Andean Flora = North to 23°; Chilenisches
Übergangsgebiet = 23°-34°, Antarktisches Waldgebiet = 34°56°
Südamerikanisches 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 41°;
Neotropical kingdom, Andean region, North of 41°.
South = Antarktisches Florenreich; North = Neotropisches
Florenreich
47
Takhtajan 1961
6
37
Takhtajan 1978
6
35
Cox 2001
Morrone 2002
5
3
12
Antarctic kingdom, Patagonian region, South of 40°;
Neotropical kingdom, Andean region, North of 40°.
Holantarctic kingdom, Chile-Patagonian region, South of 25° to
Antarctic peninsula and Malvinas Islands (Falkland Is.)
Southamerican kingdom
Austral kingdom, Andean region
2.3.1. The global classification
Gottfried R. Treviranus (1776-1837), 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 1803). 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 (1740-1830),
J. Banks (1743-1820), and G. Forster (1754-1794)15. 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. 132).
At this early stage biogeographic representation was inexistent, since the discipline was at its early
stages of development. Only 2 years later Jean B. Lamarck and Augustin-Pyramus de Candolle
published the “first biogeographical map” for the third edition of the Flore française (Ebach &
Goujet 2006). A-P. de Candolle, in his Géographie botanique further classified the world in 20
floristic regions: Chile fitted into two regions, le Chili and les terres Magellaniques (de Candolle
1820). But de Candolle’s world classification still lacked a map. Three years later, Danish
botanist Joakim Frederik Schouw (1789-1852) published the first ptytogeographical world map
(Schouw 1823, Mennema 1985). Schouw proposed 25 floristic realms =Florenreiche. Southern
South America was classified into two realms: the Reich der Holzartigen Synanthereen, and the
Antarktisches Reich, from 40°S to the South. Applying the concept of endemism recently proposed
by A-P. de Candolle, Schouw exposed explicitly the criteria for classifying and delimiting the
floristic realms (Drude 1884, p. 13):
1. half of the known plants have to be native to the territory in question;
2. ¼ of the genera had to be endemic or have their maximal distribution there;
3. A plant realm had to have some endemic families.
De Candolle’s son Alphonse de Candolle (1835) disputed the criteria Schouw used and proposed
45 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 régions, proposées jusqu’à
present, pour des systèmes artificiels.... Elles ont nui a la science”16 (A. de Candolle 1855,
p. 1304-1305, as quoted by Nelson 1978). Later he states, with reference to his Geographie:
“ouvrage du reste complètement différent de celui auquel mon père pensait, car les documents
etaient devenus plus nombreus, et mes idées s’étaient singulièrement éloignees de celles qui
régnaient dans la science depuis vingt ans “ 17(A. de Candolle Mémoires, p. 395, as quoted by
Nelson 1978).
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 (1817-1911). While sailing on board James
Cook’s Endeavour, he notably improved the floristic knowledge of the southern hemisphere.
Hooker’s publications compiled as The Botany of the Antarctic Voyage (Hooker 1844-1860), were
almost as epoch-making as Darwin’s Origin of species (Thiselton-Dyer 1909).
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 (1814-1879) to publish his Vegetation der Erde, which related explicitly the
plant world with the regional climates (Grisebach 1872). In Grisebach’s view, Chile had to be
classified into 3 regions =Gebiete: 1. a Chilean floristic core, Chilenisches Übergangsgebiet
(transition zone) from 23° to 34°, that holds a “unique flora”; 2. an Antarctic region (Antarktisches
Waldgebiet), ranging from 34° to 56°, characterized especially by the genus Nothofagus; and 3. a
tropical Andean flora that ranges from Ecuador to northern Chile (figure 2.4).
Adolf Engler (1844-1930), one of the most prominent scientists in botanical
history, working at the Botanical Garden in Berlin, was the first one to
try a synthesis of the evolution of the plant world on the earth surface
(Engler 1879, 1882). He classified the world flora into four realms and 32
regions, dividing also each region into diverse provinces and districts, thus
constructing the hierarchical system that forms the base of all the following
classification systems. Chile was classified into the Südamerikanische
Florenreich (recognizing the Nordchilenische Provinz as the Chilenische
Übergangsgebiet from Grisebach) and in the Altoceanisches Florenreich, South of 36°S. This
Altoceanisches Florenreich grouped southern Chile with New Zealand’s 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 2005, p. 26).
49
50
Figure 2.4 Vegetation of the world, after Grisebach (1872, 2d edition 1884)
51
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 1899, p. 149) as well as adding a fifth kingdom, the
Ozeanisches Florenreich, which was composed of the aquatic plants from the vast oceans.
Figure 2.6 Floristic realms from Drude (1884)
Oscar Drude (1852-1933) 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
52
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 41°S corresponds to the Andines
Florenreich and southern Chile to the Antarktisches Florenreich (figure 2.6).
Drude’s concern over about the floristic and ecological differences led him to publish separate
maps for a floristic classification and for a vegetation classification (Drude 1887). Three years
later he abandoned the floristic classification altogether: “The maps published in the geographical
reports of 1884 about my floristic classification show the uncertainty of the boundary lines due
to numerous migration routes and directions of dispersal, which overlap from one to the other
realm; it is long a known fact that each attempt to draw strict floristic boundaries, is itself ruinous”
(Drude 1890, p. 329). Drude decided to join the more physiognomic approach from Grisebach,
and he modified the classification on the basis of the new climatological basis provided by
Wladimir Köppen (1884). The floristic kingdoms still numbered 14, and South America remained
unmodified.
At the beginning of the 20th century, Ludwig Diels (1874-1945), successor of Engler in Berlin,
synthesized Drude’s classification into six floristic realms, following the early proposal from
Engler (1882) but obviating the oceanic realm (Ozeanisches Florenreich), and dividing Engler’s
Altoceanisches Florenreich into an Antarktis, an Australis and a Capensis (figure 2.7). Diels
(1908) was the first to raise the African Cape region to the category of a realm. Interestingly he
considered the Australisches Florenreich (Australis) as comprising only Australia and Tasmania -
Figure 2.7 Floristic realms from Diels (1908)
53
- and considered Malesia and New Zealand as part of the Paläotropis. South America and Central
America including Mexico and Baja California was part of the Neotropis, but southernmost
South America retained its designation as a realm, the Antarktis. Diels (1908) little book was
reprinted five times until 1958, and his realm classification was retained adding only more details
at the regional scale by Mattick (1964) and later popular authors like Good (1974) and Takhtajan
(1978). Diels’ proposal modified by the mentioned authors is still preferred in all modern German
phytogeography textbooks (e.g. Richter 1997, Schroeder 1998, Frey & Lösch 2004).
Figure 2.8 Floristic regions from Good (1947)
English botanist Ronald Good’s (1896-1992) 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 1947 till 1974. He followed Diels with the 6 realms, dividing them in 37 regions
(figure 2.8). Chile south of 41°, was classified into the Antarctic kingdom, and as the Neotropical
kingdom to the north. The scheme is very similar to that of Russian botanist Armen Takhtajan,
which became very popular after its translation into English. In a first version he maintained
the 6 realms from Diels and the 37 regions from Good (Takhtajan 1961) (figure 2.9), but in his
definitive proposal he reduced the regions to 35. Diel’s scheme of 6 realms stayed unchanged
(Takhtajan 1978) (figure 2.10). In his first classification he considered Central Chile as part of
the Neotropical region (Takhtajan 1961) (figure 2.9), but in his later work he classified all the
southern cone S of 25° into the Holantarctic kingdom (Takhtajan 1978) (figure 2.10).
The basic scheme of 6 floristic realms proposed by Diels (1908) stayed unchanged during the 20th
century; only at the beginning of this century, Cox (2001) 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.
54
Figure 2.9 Floristic regions from Takhtajan (1961)
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 (2001) discuss the Takhtajan’s Holantarctic kingdom listing the 11 endemic families and
34 endemic genera proposed by Takhtajan (1978). Takhtajan’s Holantarctics is based at the
family level mainly on American endemic families (e.g. Thyrsopteridaceae, Lactoridaceae,
Gomortegaceae, Aextoxicaceae). The dominance of American families seems to give reason to
Cox while transferring them to the South American kingdom.
The last word stays in Morrone (2002), who challenged Cox’s proposal and compiles the floristic
and faunistic knowledge in one synthetic classification. The result is a scheme of only 3 biotic
realms: the Holarctic kingdom, the Tropical kingdom (=East Gondwana), and the Austral
kingdom (=West Gondwana). Morrone (2002) related the classification to the history of these
biotas, as was Engler’s early intention (1879, 1882). In fact the result is remarkably similar to
Engler’s, but grouping the paleotropis and neotropis in one tropical realm.
In Morrone’s proposal the austral kingdom is composed by S Australia, NZ, S Africa and S South
America, extending through the Andes till 5°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 (2005, p. 236) for intercontinental relationships.
55
56
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 Reiche’s
analysis of the distribution of the Compositae family
in the country (Reiche 1905) (figure 2.11). On a
second map he also proposed possible migration
routes for these taxa (Reiche 1905) (figure
2.12). The first (and only) intend of a synthetical
cartography for the Chilean flora is expressed in
the two maps that accompanied Reiche’s Plant
geography (Reiche 1907) (figure 2.13, 2.14). Here
Reiche proposed distribution ranges and limits
for some key taxa in his view. In a second map
he divided the country in several floristic units,
integrating floristic and physiognomic knowledge.
Reiche proposed the limit between the antarctic
and neotropical reamls at around 41°S. In spite of
appearing so simple, it is notably the first intend
of a floristic cartography for Chile. 50 years later,
Schmithüsen drew a schematic representation
of the different floristic elements composing the
woody flora between 30° and 42°S. (Schmithüsen
1956) (figure 2.15).
The floristic cartographic task at the national level
seems since then to be frozen in time, having
been replaced by more general biogeographical
classifications (e.g. Cabrera & Willink 1973;
Rivas-Martínez & Navarro 2000; Morrone 2001,
2006; see a discussion in Ribichich 2002 for the
task in adjacent Argentina).
Figure 2.11 The first plant geographical map for
Chile (Reiche 1905)
57
Figure 2.13 Geographic ranges and distribution
limits of selected taxa (Reiche 1907)
Figure 2.12 Possible migration routes for the
Chilean Compositae (Reiche 1905)
58
1. General floristic area of
neotropical origin
2. Forest flora of south-hemispheric-subtropical origin
3. Laurophyllous neotropical
and suth-hemispheric origin
4. Neotropical sclerophyllous
element
5. 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 2.14 Plant geographical divisions of Chile
(Reiche 1907)
59
Figure 2.15 Diagram of floristic elements
in Central Chile (Schmithüsen 1956)
2.4 Excursus: vegetation maps and
Vegetationsbilder
Contrary to the floristic cartography, the vegetation
cartography has seen a good development in Chile.
Mainly on the base of the early climatic classification
(Köppen 1930), followed by bioclimatic proposals
for the country (di Castri 1968, di Castri & Hajek
1976, Quintanilla 1974, Amigo & Ramírez 1998),
the vegetation cartography has developed in
hand of Schmithüsen (1956) (figure 2.16), Hueck
(1978), Pisano (1977, 1981, 1983), Quintanilla
(1983, 1985, 1988), Gajardo (1994), and Luebert
& Gajardo (2005). These efforts and the use of
global climatic surfaces on a GIS-based platform
allowed Luebert & Pliscoff (2006) to publish the
most accurate bioclimatic and vegetation synthesis
to date. The early works of C. Skottsberg deserve
special attention (Skottsberg 1910a, 1916). Due to
the impressive floristic knowledge of the eminent
botanist, his maps can be considered a good
synthesis of floristic and physiognomic information
(figure 2.17). Remarkably are also Skottsberg’s three
volumes on the Natural History of Juan Fernández
and Easter Island (Skottsberg 1920-1956), as well
as his Vegetationsbilder, that spread the images
of Chilean landscapes at the beginning of the 20th
century (Skottsberg 1906, 1910b) (figure 2.18).
Figure 2.16 Schmithüsen‘s vegetation map (1956):
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.
60
Figure 2.17 Phytogeographic map from Skottsberg (1910a)
61
Figure 2.18 Vegetationsbild from Skottsberg (1910 b)
62
2.5 Geographic ranges in the latitudinal profile
The distribution of the genera along the latitudinal profile in Chile is based on the collections of
the National Herbarium (SGO), that have been partially transferred to a data base with geographic
coordinates (see Muñoz-Schick & Moreira-Muñoz 2002). The data base is a work in progress,
and ca. 60 000 records have been checked so far, from ca. 100 000 specimens of native vascular
plants registered at SGO. Dot maps have been done as an event theme in ArcView 2.3 GIS
program. Every genus has been checked for false or lacking coordinates, and for evident wrong
determinations. Also very helpful for plotting the distribution has been the revision of several
regional floras and checklists (Skottsberg 1916, Moore 1983, Henríquez et al. 1995, Marticorena
et al. 1998a, 2001) and local floras and checklists (e.g. Muñoz-Schick 1980, Teillier et al. 1994,
2005; Richter 1995, Rundel et al. 1996, Arroyo et al. 1998, 2000, 2002, 2003; Luebert & Gajardo
2000, 2005; Luebert et al. 2002, Villagrán 2002).
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) 102 genera occur in Chile in a very small range (<1 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) 115 genera show a small distribution range between 1 to 5 degrees of latitude (e.g. Aphanes,
Traubia, Guindilia, Orites);
c) 154 genera have a medium- small range of distribution from 5 to 10 latitudinal degrees (e.g.
Luciliocline, Hebe, Quillaja, Aa, Lepidoceras);
d) 231 genera show medium-large ranges of distribution, from 10 to 20 latitudinal degrees (e.g.
Acacia, Podocarpus, Cissus, Oziröe, Azara);
e) 211 genera have ranges larger than 20 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 (17°35’) to Cabo de Hornos (56°).
63
-57
Also the distribution of the taxa was
summarized in 10 degree latitudinal bands,
from 17,6° to 27° (arid tropical zone) (figure
2.20); 27° to 37°S (Mediterranean-type zone)
(figure 2.21); and 37° to 56S° (temperate
zone) (figure 2.22).
Figure 2.19 Latitudinal ranges for Chilean
families
64
-47
-37
-27
-17
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
-57
-47
-37
-27
-17
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
Oxychloë
Pitraea
Polyachyrus
Nototriche
Malacothrix
Amblyopappus
Tetragonia
Copiapoa
Lenzia
Prosopis
Lycium
Chuquiraga
Bakerolimon
Echinopsis
Malesherbia
Pennisetum
Sicyos
Figure 2.20 Latitudinal ranges for Chilean genera with average distribution north of 27°S
Cristaria
Argemone
65
-47
-37
-27
-17
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
-57
-47
-37
-27
-17
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
66
Figure 2.21 Latitudinal ranges for Chilean genera with average distribution in Central Chile between 27°S and 37°S
-57
-47
-37
-27
-17
-57
-47
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
-37
-27
-17
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
67
Figure 2.22 Latitudinal ranges for Chilean genera with average distribution in the temperate zone south of 37°S
-57
68
3 Geographic Relationships of the Chilean Flora
69
March-December 2006
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 1950, p. 204, see also
Wangerin 1932). In any case Wulff (1950) was the opinion that geographical [floristic] elements
are fundamental for an understanding of a flora, and that an analysis of a flora should begin with
these elements. More recently, floristic elements have been analysed for the North American flora
north of México (Qian 1999), the East Asian flora (Qian et al. 2003), the flora of the Bolivian
Andean valleys (López 2003), and the Ecuadorian superpáramo (Sklenar & Balslev 2007). In the
case of the Chilean flora, the first attempt is the one of Reiche (1907). He was able to identify
seven elements or Kontingente (table 3.1). Later, Villagrán & Hinojosa (1997) 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 3.1 Floristic elements in the Chilean Flora
Reiche 1907 (Kontingente)
Villagrán & Hinojosa 1997
1
Tropical-American
Neotropical (amplio / disyunto)
2
Andes Chile/Argentina
3
California/Mexican
Endémico bosques subantárticos /
Endémico de Chile
-
4
Antarctic
Australasiano (cálido/antártico)
5
Boreal (Europa, S Chile)
Amplio
6
Pantropic
Pantropical
7
Introduced flora
-
The present work is the second attempt, after Reiche (1907), 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 (1952, 1958)18, and have been successfully used for the analysis and interpretation of
disjunct distribution patterns (e.g. Katinas et al. 1999 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. (2006) 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 1947), and this
approach will be followed here. The global distribution for each genus has been obtained from
71
Wielgorskaya (1995) and Mabberley (1997). 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 2.2, 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 3.2 Floristic elements for the vascular plant Chilean genera (update 07 February 2007)
Floristic elements
1. Pantropical
2. Australasiatic
3. Neotropical
4. Antitropical
5. South-temperate
6. Endemic
7. Cosmopolitan
N° Genera
88
59
216
152
81
83
134
Total
813
%
11
7
27
19
10
10
16
100 %
Table 3.3 Generalized tracks for the vascular plant Chilean genera (update 07 February 2007)
Tracks (follow subsections in the text)
N° of genera
3.2.1 Austral-antarctic track
31
3.2.2 South Pacific tropical track
25
3.2.3 Circum-austral track
3
3.3.1 Wide Neotropical track
64
3.3.2 Andean track
113
3.3.3 South Amazonian track
39
3.4.1 Wide antitropical track
84
3.4.2 Antitropical disjunct Pacific track
56
3.4.3 Circum-Pacific track
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 (6 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,
72
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 1960
Acacia (Fabaceae)
Bacopa (Gratiolaceae), sensu Dawson 1968
Cotula (Asteraceae), Meusel & Jäger 1992, Bremer 1994
Nicotiana (Solanaceae), sensu van Steenis & van Balgooy 1966
Lobelia (Campanulaceae), sensu Meusel & Jäger 1992
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 (1995) 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 3
genera. Within these genera, three generalized tracks based on superimposing distributions have
been identified.
73
Azorella distribution, sensu Baumann 1988
Berberidopsis distribution, sensu Ronse De Craene 2004
Coprosma distribution, sensu van Steenis & van Balgooy 1966
Hebe distribution, sensu van Steenis & van Balgooy1966
Jovellana distribution, sensu Dawson 1968, Heads 1994
Lagenophora distribution, sensu Cabrera 1966
Leptinella distribution, sensu Bremer 1994
Oreomyrrhis distribution, sensu Mathias & Constance 1955
Rytidosperma distribution, sensu Linder & Barker 2000
Ourisia distribution, sensu Meudt & Simpson 2006
Figure 3.2 Examples of the australasiatic floristic element
74
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.
(1999).
It numbers 31 genera, e.g. Luzuriaga, Eucryphia, Prumnopitys, Jovellana, Pseudopanax.
Most of these genera are restricted to the southern temperate forests, but some of them
reach northernmost Chile (e.g. Hypsela, Azorella, Cortaderia) and some of them even
reaching Ecuador. Some genera reach Central Chile, disappear in the Atacama and
reappear in the northern Andes (e.g. Fuchsia, Lomatia, Prumnopitys). Haloragis is an
austral genus that reaches the Juan Fernández islands, but not the continent.
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 25 genera, e.g. Araucaria, Nothofagus, Lagenophora, Abrotanella, Coprosma,
and Hebe. Many genera are restricted to the southern temperate forests, but a few reach
northernmost Chile (e.g. Colobanthus, Muehlenbeckia, Pratia). Some reach central Chile,
disappear in the Atacama and reappear in the northern Andes (e.g. Citronella, Uncinia,
Oreobolus). Doodia is widespread in the Pacific till Isla de Pascua, not in continental
Chile. Dicksonia, Arthropteris, Coprosma, and Santalum are austral genera that are
represented in the Juan Fernández Islands, also absent from the continent19.
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 3 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 216 genera, mostly from the Asteraceae (53), followed by Solanaceae (12)
and Poaceae (12), Cactaceae (11), Brassicaceae (7) and Apiaceae (7). Three generalized tracks
have been identified within this element:
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.
75
Azara
Alstroemeria
Eremodraba
Cremolobus
Calceolaria
Escallonia
?
Eudema
Malesherbia
Mecardonia
Nototriche
Mathewsia
Vasconcellea
Figures 3.3 Examples of the neotropical floristic element (distribution from monographs and checklists)
76
The wide Neotropical track comprises 64 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 113 genera, e.g. Chuquiraga, Nototriche, Polylepis, Geoffroea. Some are
found in all of Chile (e.g. Perezia, Escallonia); others only in northernmost (Dunalia,
Stenomesson) or north-Central Chile (Exodeconus, Equinospsis, Lycopersicon); many
distributions are disjunct between southern Chile/Argentina and northern South America
(e.g. Llagunoa, Myrcianthes, Dysopsis, Blepharocalyx, Desfontainia, Myrteola). More
than 80 genera have a main massing in the central Andes of Perú, Bolivia, northern
Argentina and Chile. Most of them occur in a continuous range from northern Chile to
Peru/Bolivia (e.g. Neowerdermannia, Oreocereus, Tunilla, Pycnophyllum, Philibertia,
Balbisia, Acantholippia), others occur disjunctly between central Chile/Perú (Tetraglochin,
Weberbauera, Kageneckia, Eccremocarpus, Lucilia). 19 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 39 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. Welß (pers. comm.) correctly noted, the most
appropriate term would be antitropic, since amphitropical means both tropics. In Cox’s (1990)
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 2005, and
Parenti 2007).
(26), Poaceae (20), Fabaceae (8), Polemoniaceae (7), and Boraginaceae (7).
77
Hydrangea distribution, sensu Mabberley 1997
Antennaria distribution, sensu Meusel & Jäger 1992
Artemisia distribution, sensu Meusel & Jäger 1992
Euphrasia distribution, sensu Van Steenis 1966, Dawson 1968
E Asia
Osmorhiza, sensu Wood 1972
Tropidocarpum, sensu Al-Shehbaz 2003
Frankenia, sensu Wahlen 1987
Agoseris, sensu Wood 1972
Figures 3.4 Examples of the antitropical floristic element
78
Mancoa, sensu Al-Shehbaz comm. pers.
Larrea, sensu Hunziker et al. 1972
One of the naturalists to refer to this element was G. Treviranus (1803), even though he did
not recognise it formally. While discussing the components of his Antarktische Flor, he noticed
that there was a floristic relationship between southern South America and New Zealand, and
that some of this genera have their main distribution in the northern and southern temperate
zones (e.g. Pinguicula, Salix, Ribes) (Treviranus (1803, pp. 131-132). Du Rietz (1940) 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 1960s (Constance 1963). 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).
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 56 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).
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
Grammosperma
Melosperma
Onuris
Embothrium
Hamadryas
Menonvillea
Pilgerodendron
Fonkia
Laureliopsis
Monttea
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.
(9), Apiaceae (5), 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 21°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 83 genera, of which 67 genera are endemic to continental Chile (e.g. Oxyphyllum,
Bridgesia, Leontochir, Balsamocarpon, Trevoa, Jubaea), while 16 genera are endemic to the
Chilean Pacific islands, especially Juan Fernández (e.g. Dendroseris, Lactoris, Cuminia, Juania).
The endemic element will be further discussed in sections 4.3, 4.4, and 4.5.
Acrisione
Adenopeltis
Alona
Anisomeria
Figure 3.6 Endemic floristic element (collections from SGO)
81
Araeoandra
Avellanita
Bakerolimon
Balsamocarpon
Bridgesia
Desmaria
Dinemagonum
Gethyum
Gomortega
Calopappus
Dinemandra
Guynesomia
Cissarobryon
Conanthera
Epipetrum
Ercilla
Gymnachne
82
Gypothamnium
Copiapoa
Fascicularia
Cyphocarpus
Francoa
Hollermayera Homalocarpus
JF
Huidobria
Legrandia
Ivania
Hymenoglossum
Leontochir
Leptocarpha
Leucocoryne
Jubaea
Leunisia
Lapageria
Latua
Marticorenia
Metharme
JF
Microphyes
Miersia
Miqueliopuntia
Moscharia
83
Neuontobothrys
Notanthera
JF
Ochagavia
Placea
Tecophilaea
Oxyphyllum
Pleocarphus
Tetilla
Podanthus
Traubia
Peumus
Reicheella
Trevoa
Figure 3.6 Endemic floristic element (continues)
84
Phrodus
Sarmienta
Valdivia
Pintoa
Pitavia
Scyphanthus
Speea
Vestia
Zephyra
3.7 Cosmopolitan floristic element
There are not many genera that can be considered as really cosmopolitan (Good 1974), 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 134 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).
Bidens (Asteraceae), sensu Meusel & Jäger 1992
Apium (Apiaceae), sensu Meusel et al. 1978
Eryngium (Apiaceae), sensu Meusel et al. 1978
Hydrocotyle (Apiaceae), sensu Meusel et al. 1978
Lycium (Solanaceae), sensu Meusel et al. 1978
Mimulus (Phrymaceae), modified from Meusel et al. 1978
Figure 3.7 Cosmopolitan floristic element
85
86
4 Biogeographic analysis
87
March-December 2006
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 / 2] (see appendix A for the latitudinal limit of each genus) Results are
shown in table 4.1 and figure 4.1.
Table 4.1 Average of geographic range of floristic elements
floristic element
Average (°S)
-32,2
-41,5
-29,6
-35,1
-42,3
-32,2
-36,0
1. Pantropical
2. Australasiatic
3. Neotropical
4. Antitropical
5. South-temperate
6. Endemic
7. Cosmopolitan
The neotropical element has the northernmost
average (29,6°S), while the south-temperate
element has the southernmost average (42,3°S). The
australasiatic element has an average at 41,5°S. The
pantropical, antitropical, cosmopolitan and endemic
elements show an average latitude between 32°S and
36°S. The fact that most elements have their average
in Central Chile tends to reinforce the early view of
Grisebach (1872), Engler (1882), and Schmithüsen
(1956) to consider this region as a transition zone
with different converging elements.
18°
24°
neotropical
pantropical/
endemic
antitropical
cosmopolitan
australasiatic
south-temperate
30°
36°
42°
48°
After grouping the Chilean plants in seven floristic
54°
elements (Kontingente), Reiche (1907, p. 319)
proposed that the neotropical and antitropical
(Reiche’s 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 (~ 35°). Reiche illustrated these migration routes in his map
for the Compositae (Reiche 1905) (figure 2.12). Now, Reiche’s view can be complemented by
analysing the disjunct distribution patterns in all the floristic elements detected.
4.1. To be or not to be disjunct
Villagrán & Hinojosa (1997) classified some of the temperate woody genera in an elemento
neotropical disyunto. These authors suggest that the disjunct distribution is the result of the
former existence of a continuous subtropical forest from SE Brazil to central/South Chile during
Oligocene/Miocene times (see chapter 5).
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.
Wide Neotropical track (64)
Andean (113) and South-Amazonian (39) tracks
Austral-antarctic track (31)
South Pacific tropical track (25)
Circum-austral track (3)
Wide Antitropical track (84)
Pacific antitropical track (56)
Circum-Pacific track (9)
Panbiogeographic nodes
Figure 4.2 Generalized tracks for the Chilean plant disjunct ditributions (in parenthesis the number of
genera composing the tracks)
90
As noted by Moore et al. (2006), researchers have debated the causes of the antitropical disjunctions
for over a century (e.g. Bray 1900; Johnston 1940; Constance 1963; Raven & Axelrod 1974;
Hunziker et al. 1972; Solbrig 1972a,b; Carlquist 1983). Moore et al. (2006) 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 1940, Barbour 1969).
c) long-distance dispersal, probably by migrating birds; from the 1960s on, many researchers
have favoured this explanation as the main cause for the disjunctions (Raven 1963, 1972;
Cruden 1966; Carlquist 1983; Simpson & Neff 1985).
The commonly preferred scenario of long-distance dispersal (LDD) which was developed since
the 1960s is well described by Raven (1963, p. 153): “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 6.3).
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 (2006) opinion, these disjunct groups could provide evidence
both of earlier dispersal events, and of multiple dispersal events.
Moore et al. (2006) 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 1963, p. 153). 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).
91
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 (1882), Diels (1908) and Skottsberg (1916), which draw a boundary
between the neotropical and the austral floristic realms at around 47°S (figure 2.17);
b) Takhtajan’s (1978) newest view setting this line at around 23°S (figure 2.10);
c) the modern proposal of Cox (2001) that aligns all of the South American flora under an American
kingdom;
d) and recent Morrone’s (2002) classification that situates the Chilean biota plus the whole Andes
in an Austral kingdom.
The latitudinal range of several genera supports each of these views:
a) the boundary at ~47°S is shown by several genera that do not overpass this limit to the north,
like Tetrachondra, Grammosperma, Hebe, Oreomyrrhis, or Rostkovia.
b) the boundary at 23° poposed by Takhtajan (1978) is shown by most (41) of the genera that
compose the australasiatic element. The Fray Jorge fog forest, located at latitude 30°, has long
been recognized as the northern outpost of some of the subantarctic elements (Muñoz Pizarro &
Pisano 1947, Trocoso et al. 1980, Squeo et al. 2004). Some kilometres north of Fray Jorge, in
Pichasca, the northernmost fossil occurrence of the emblematic subantarctic genus Nothofagus
has been documented (Torres & Rallo 1981).
c) Cox’s (2001) vision neglects or minimizes the australasiatic element in the American flora and
therefore do not leave opportunity for comparisons.
d) In defense of Morrone’s (2002) 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
Desfontainia/
10°
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 4.3 Northward extent of spread of Gondwana
genera in tropical America (redrawn from Webster 1995)
answer the question. Since the latitudinal
92
distribution of the genera, as represented in figures
2.19 to 2.22, does not account for possible distribution
or collection gaps, four regional floras were taken for
a similarity analysis: Antofagasta (ANT), Coquimbo
(COQ), Biobío (BIO), and Magallanes (MAG) (figure
4.4). Three of the regions included in the anaylsis have
a published checklist (table 4.2). The checklist for
BIO was gently provided by Dr. R. Rodríguez from
CONC herbarium. Synonym taxa were homologized
(e.g. Lagenophora=Lagenifera, sensu Cabrera 1966)
. 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
ANT
Isla de Pascua
COQ
Juan Fernández
BIO
MAG
Jaccard similarity index was applied to the data Figure 4.4 Chilean continental regions included
set (Zunino & Zullini 2003, Cox & Moore 2005, in the similarity analysis and location of Chilean
Pacific islands.
Lomolino et al. 2006) (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
Antofagasta
126.049
N° native vascular
plant genera
316
COQ
Coquimbo
40.580
457
Marticorena et al. 2001
Biobío
37.063
465
CONC, R. Rodríguez pers. comm.
Magallanes
132.033
252
Henríquez et al. 1995
BIO
MAG
Regions
Area (km2)
Source
Marticorena et al. 1998a
Table 4.3 Floristic similarity and geographic distance different regions in Chile: Antofagasta (ANT),
Biobío (BIO), Coquimbo (COQ) and Magallanes (MAG). Geographic distance have been calculated as
the latitudinal difference between the geographic centroid of each region.
Compared regions
ANT/COQ
ANT/BIO
ANT/MAG
COQ/BIO
COQ/MAG
BIO/MAG
Shared genera
244
174
97
323
150
178
Similarity
Jaccard
0,46
0,29
0,21
0,54
0,27
0,33
93
Distance (km)
777
1527
3227
750
2450
1700
1.0
0.8
0.6
0.4
COQ/BIO
L
ANT/COQ
ANT/BIO
L
L
BIO/MAG
L COQ/MAG
0.2
Jaccard similarity
L
L
ANT/MAG
L
L
L
0.0
L
0
2000
4000
Figure 4.5 Similarity
8000
10000 versus geographic
6000
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
4.6).
Table 4.4 Floristic elements in each region
REG
ANT
COQ
BIO
MAG
pantropical
n
28
39
42
11
%
8,89
8,59
9,09
4,38
australasiatic
n
8
22
41
38
%
2,54
4,85
8,87
15,14
neotropical
n
115
114
100
29
%
36,51
25,11
21,65
11,55
antitropical
n
45
73
72
44
%
14,29
16,08
15,58
17,53
southtemperate
n
17
42
51
42
%
5,40
9,25
11,04
16,73
endemic
n
15
38
27
1
40,00
%
30,00
1.
2.
3.
4.
5.
6.
7.
20,00
10,00
Pantropical
Australasiatic
Neotropical
Antitropical
South-temperate
Endemic
Cosmopolitan
0,00
ANT
COQ
BIO
MAG
Regions
Figure 4.6 Floristic elements present in four regional floras (percentage)
94
%
4,76
8,37
5,84
0,40
cosmopolitan
n
82
117
123
84
%
26,03
25,77
26,62
33,47
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 (1882), Drude (1884), Reiche (1907, p. 282), Diels (1908),
and Skottsberg (1916).
4.3 Analysis of Endemism
Endemism, since Augustin P. de Candolle (1820) 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. 2006). Taxa can be endemic to a location due
to three different reasons: (1) they originated in that place and never dispersed; (2) their entire
range has shifted in locality after the origin; or (3) 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 1994, Linder 2001). In the words of Nelson & Platnick (1981) “the most elementary
questions of historical biogeography concern areas of endemism and their relationships”.
Hereafter, the patterns of endemism and its hierarchical arrangement in the Chilean continental
vascular flora will be analysed. Since the country has mainly natural borders, i.e. the Andes and the
Pacific, it can be interpreted as a natural unit for the analysis of endemism. Reports on endemism
at the genus level have been partially published by Muñoz-Schick & Moreira-Muñoz (2000).
50
N° endemic genera
40
30
20
10
0
1
2
3
4
5
6
7
8
9
N° species per genus
95
10+
Fig. 4.7 N° of endemic genera v/s N° of
species per genus
-57
-47
-37
-27
-17
Neuontobothrys
Metharme
Ivania
Leontochir
Marticorenia
Avellanita
Valdivia
Reicheella
Oxyphyllum
Gypothamnium
Pintoa
Balsamocarpon
Miqueliopuntia
Dinemagonum
Araeoandra
Guynesomia
Bridgesia
Pleocarphus
Tecophilaea
Leunisia
Gethyum
Placea
Traubia
Speea
Jubaea
Calopappus
Tetilla
Cissarobryon
Legrandia
Pitavia
Gomortega
Desmaria
Hollermayera
Latua
Dinemandra
Zephyra
Copiapoa
Bakerolimon
Phrodus
Cyphocarpus
Homalocarpus
Moscharia
Trevoa
Adenopeltis
Podanthus
Scyphanthus
Ochagavia
Miersia
Acrisione
Gymnachne
Vestia
Fascicularia
Leptocarpha
Huidobria
Alona
Conanthera
Anisomeria
Leucocoryne
Microphyes
Epipetrum
Peumus
Lapageria
Sarmienta
Notanthera
Fig. 4.8 Latitudinal ranges for the 67 Chilean endemic genera. Small distribution
ranges on top, large towards the buttom.
Francoa
Ercilla
Hymenoglossum
96
4.3.1 Geography of Endemism
The 67 endemic continental Chilean genera (section 3.6) belong to 36 families (appendix A).
In continental Chile, the family with the highest number of endemic genera is Asteraceae (11).
The Alliaceae are represented by 4 endemic genera, and the Brassicaceae, Solanaceae and
Tecophilaeaceae by 3 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 2.6).
The most species-rich genus is Copiapoa (Cactaceae), which contains 25 endemic species,
followed by Leucocoryne (11, 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 (46 genera).
The distribution ranges of the endemic genera were grouped in four categories: small (within one
latitudinal degree), medium-small (1 to 5 latitudinal degrees), medium-large (5 to 10 latitudinal
degrees) and large (more than 10 latitudinal degrees). Of the 67 endemic genera analysed in
this study, 6 (8,9% of the total) belong to the smallest category, i.e. are found only within 1
degree of latitude (figure 4.8). These are the genera Avellanita, Ivania, Leontochir, Metharme,
Neuontobothrys, and Valdivia. 28 genera (41,8%) occur in medium-small ranges of distribution
between 1 and 5 latitudinal degrees, e.g. Leunisia, Tetilla, Jubaea. 20 genera (29,9%) feature
medium-large ranges of distribution between 5 and 10 latitudinal degrees, e.g. Copiapoa, Miersia,
Vestia. 13 genera (19,4%) feature large ranges of distribution wider than 10 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 19 latitudinal degrees from
25-26
27-28
21° to 40° S.
29-30
Latitude (°S)
31-32
The analysis of the generic richness indicates
a latitudinal trend, with the greatest number of
endemic genera (35) ranging between 33°-34°
S (figure 4.9) The number of endemic genera
decreases constantly to the north and to the south
of this central area.
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
N° genera
Fig. 4.9 Endemic genera at each latitude degree.
97
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
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 4.10 Geographic
ranges of endemic Chilean genera
Fig. 4.11 Grid (111 x 111 km) for the
PAE application in continental Chile
Figure 4.12 Generic richness for each
111 x 111 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 1988, Morrone
1994). 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 1992, as quoted by Nihei
2006). 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 1992), i.e. endemic taxa that are found
98
in more than one area (Nihei 2006). 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 (1/0) 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 3.6). 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 100 replicates resulted in 410 most parsimonious trees. A TNT search resulted
in only five most parsimonious trees. Consensus trees were obtained using the majority-rule,
with confidence values calculated at each node (figure 4.13 shows the consensus tree of the TNT
search).
The consensus tree discriminate two major areas of accumulated endemism within a group of
central Chilean coastal cells (31-38°S) = a central coastal floristic block; and a group of central
Chilean Andean cells (33°-39°S) = a central Andean floristic block. A third group of southern
cells including Chiloé Island (44°S) can be interpreted as a transition toward the temperate zone,
that features fewer endemic genera. The cells from 31°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 39° to Chiloé is characterized
by Latua, Hollermayera, and Desmaria. Northern endemic taxa outside of the core areas of
endemism are Zephyra, Reicheella, Oxyphyllum, and Metharme (not shown in figure 4.14, but
see figures 3.6 and 4.8).
Pliscoff (2003) and Rovito et al. (2004) 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
Xco
100
Vco
Wco
80
60
100
Van
80
Tan
80
Uco
100
Uan
100
100
San
100
Tco
100
Sco
100
Qan
Ran
Rco
100
Qco
Oco
Pco
Figure 4.13 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. 5 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.
100
Central coastal floristic block
Central Andean floristic block
Southern floristic block
Fig 4.14 Floristic blocks based on
terminal cells from figure 4.13.
(geodiversity) (Cowling et al. 1996, Mutke & Barthlott 2005). The central area of endemism lies
within the Mediterranean-type climate zone of Central Chile, which is located between 27°37°S (Luebert & Pliscoff 2006). 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 34°S) seems to be very recent (Villa-Martínez et al. 2003). This suggests, for Chile as well
as for the European Mediterranean region (Verdú et al. 2003), that the endemic genera are not
directly related to the current climate, but rather to the Paleogene history of the biota (Hinojosa
et al. 2006, and chapter 5). Landrum (1981) proposed that the high proportion of endemic genera
present in central Chile is due to the long isolation of these forests, at least 10 to 30 mya.
Central Chile as a core area of endemism can be interpreted as a panbiogeographic node (sensu
Heads 2004) 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
101
historical geography are worth mentioning. In the opinion of some authors, e. g. Santos (2005),
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 (2005) is valid in the sense that one has to be aware of the
limitations and assumptions inherent to the methods20. But in a recent revision, Nihei (2006)
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 2005).
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. 2002, Szumik & Goloboff 2004, Domínguez et al.
2006, see a comparison of methods in Moline & Linder 2006). 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 (2002) paper.
Anyway, as demonstrated by the bibliographical survey done by Nihei (2006 + online Appendix
S1), 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 2006). 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 5).
The presence of a high number of monotypic genera suggests that most of the endemic genera are
representatives of old lineages that have lost must of their diversity due to extinctions.
Palaeogeographic scenarios show a ongoing exchange between neotropical and subantarctic
palaeofloras from the late Cretaceous to the Pliocene (Hinojosa & Villagrán 1997), or even
starting in the early Cretaceous (Barremian/Aptian). This affected the lineages of which extant
angiosperms originated (Troncoso & Romero 1998). 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.
102
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 (Villagrán & Hinojosa 1997). Peumus, joining other extant Lauraceae taxa
(Cryptocarya, Beilschmiedia, and Persea), was already a component of the middle Miocene flora
in central Chile (Hinojosa & Villagrán 1997). Renner (2004) proposed that Peumus diverged from
the Monimia/Palmeria line about 76 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 (24-21 mya; Nishida
et al. 1989, quoted by Renner 2004), but molecular analysis suggests an age of 100 mya for
Gomortega (Renner 2004). 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 4.15 Phylogenetic and geographic relationships of Gomortega and Atherospermataceae
Ripogonum (Australia/N. Guinea/NZ)
Lapageria (endemic Chile)
Philesia (Chile/Argentina)
Source: Vinnersten & Bremer 2001
Figure 4.16 Phylogenetic and geographic relationships of Philesiaceae
103
Neotropical relationship
Chilean endemic genera Dinemandra, Dinemagonum and Leontochir have an evolutionary
relationship with neotropical lineages (Davis et al. 2001, Aagesen & Sanso 2003), 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 43-53 mya (DeVore
& Stuessy 1995). However, endemic genera could be much more recent, e.g. the ancestor of
Moscharia (endemic) and Polyachyrus (Perú/Chile) possibly occupied a part of north-central
Chile during the last uplift of the Andes in Pliocene and Pleistocene times (Katinas & Crisci
2000). 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 22 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. 2003). For example, Calopappus
is considered by some authors as a synonym of Nassauvia (Bremer 1994), and Leontochir as a
synonym of Bomarea (Hofreiter 2006, but see Aagesen & Sanso 2003). Other endemic genera
like Guynesomia have only recently been recognized as being different from their closest relatives
(Bonifacino & Sancho 2004). 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 4.5), e. g. Pilgerodendron, Lebetanthus, Huanaca, Philesia, Asteranthera, Mitraria, and
Sarmienta.
104
Table 4.5 Australasiatic relationships of South-temperate genera
Taxon
Phylogenetic relationships
Source
Pilgerodendron
(Cupressaceae)
Genus from southern Chile/Argentina, closely related to
Libocedrus (New Zealand); some authors propose to unite them in
one genus.
Epacridaceous genus from (MAG/Argentina) is closely related to
Prionotes (Tasmania)
Gadek et al. 2000,
but see Farjon et al.
2004
Crayn et al. 1998
Huanaca
(Apiaceae)
Philesia (+ endemic
Lapageria)
Patagonian genus close related to New Zealand Stilbocarpa
Andersson et al.
2006
Vinnersten &
Bremer 2001
subfamily
Coronantheroideae
Members of the Gesneriaceae restricted to the Southern
Hemisphere that are closely related. A Gondwanan origin
for the whole family has been proposed. The small group of
Coronantheroid Gesneriaceae supposedly invaded the Americas
via Antarctica and southern South America and gave rise to the
Gesnerioid Gesneriaceae. While the Coronantheroid Gesneriaceae
became nearly extinct (the three Chilean/Argentinian genera
Asteranthera, Mitraria, Sarmienta being the last survivors), the
Gesnerioid Gesneriaceae evolved explosively in the American
tropics.
Lebetanthus
the only members of the South American Philesiaceae, are
supposedly splitted from the Australian–New Zealand Ripogonum
at 47 +- 8,4 mya.
Burtt 1998, Weber
2004
As example, Vinnersten & Bremer (2001) suggest that the ancestor of Ripogonum, Lapageria,
and Philesia was distributed in South America and New Zealand and possibly also in Australia.
The isolation of South America from Australia and New Zealand corresponds to the split of the
South American Lapageria and Philesia from the Australian–New Zealand Ripogonum estimated
to 47 +- 8.4 mya. The same interpretation is possible for the split between Alstroemeriaceae and
Luzuriagaceae, so it may be that termination of the Antarctic link during Eocene resulted in two
vicariance events within Liliales (Vinnersten & Bremer 2001).
105
4.5 Plant geography of the Chilean Pacific islands
The Pacific islands offshore Chile are mostly treated as oceanic islands, following the early
classification of Wallace (1880). According to the theory of equilibrium in island biogeography,
the biota of these islands is the result of immigration and extinction rates (MacArthur & Wilson
1967). The Chilean Pacific islands show different situations: Juan Fernández, Desventuradas,
and Easter Island (Isla de Pascua), are located at different distance and latitude in relation to the
Chilean coast, and show different floristic relationships.
Figure 4.17 Isla de Pascua
(from Skottsberg 19201956).
4.5.1 Isla de Pascua (27° 05’ S, 109° 20’ W)
Isla de Pascua (Easter Island), located at 3,765 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 1990). 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 2.2). 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 1991). Based in the works of Skottsberg (1920-1956) more recently revised by Marticorena
(1990) and Zizka (1991), a checklist with a total of 20 families and 29 genera was compiled
(appendix D). The best represented family is the Poaceae, with 5 genera. Several taxa that
appeared with a question mark in Zizka’s revision and considered as introduced by Marticorena
(1990) have been left out from the analysis (e.g. Caesalpinia, Calystegia), while some listed by
Marticorena as aliens have been retained due to the reasons exposed by Zizka, like Triumfetta
(Malvaceae), or Kyllinga (Cyperaceae). The result of phytogeographic elements analysis shows
a clear predominance of pantropical (41%) and cosmopolitan genera (49%), with little presence
106
1. Pantropical
2. Australasiatic
3. Neotropical
4. Antitropical
5. South-temperate
6. Endemic
7. Cosmopolitan
41%
49%
0%
3%0%
7%
Figure 4.18 Floristic elements
Isla de Pascua
0%
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 (1920-1956). 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 4.19 Isla San Félix. [www.cordell.org/
SFX/SFX_pages/SFX_Main.html]
Figure 4.20. Isla San Ambrosio (after Kuschel 1962)
4.5.2 Islas Desventuradas, San Félix (26°17’ S / 80°05’) and San Ambrosio (26°21’S /
79°53’W)
This relatively small volcanic archipelago is located approximately 850 km off the Chilean coast
(figure 4.4.). Desventuradas Islands consist of the two main islands San Félix (figure 4.19) and Isla
(de) San Ambrosio (figure 4.20) plus several rocks and stacks: Islote Gonzalez and Roca Catedral.
Together, the Desventuradas Islands have a surface area of only 10.3 km². The topography is very
rugged, with peak elevations of 193 m asl on Isla San Félix, 479 m a.s.l. on Isla San Ambrosio.
107
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
(1870), F. Philippi (1875), Skottsberg (1937, 1951, 1963), Johnston (1935), Gunckel (1951), and
Kuschel (1962). More recent treatments have been done by Marticorena (1990) and Hoffmann
& Teillier (1991)21. According to these authors, the vascular flora of the islands consists of 13
families, 18 genera and 25 native species. More recently Teillier & Taylor (1997) 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 13 families and 19 genera was
compiled (appendix D). The best represented family is the Amaranthaceae (4 genera formerly
classified under the Chenopodiaceae).
The floristic element that dominate in the islands is the cosmopolitan element (52%), but the
endemic element is also noteworthy, reaching a 21% (figure 4.21). This percentage is higher
than the one showed by the continental flora or Juan Fernández at the genus level. Furthermore,
Marticorena (1990) reports a level of endemism of 60,6% 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 4.21 Floristic
elements Islas
Desventuradas
The natural history of the archipele has been analysed by Skottsberg (1937), coming to a similar
conclusion that for the Juan Fernández Islands, in the sense that the Desventuradas flora do not
show a oceanic character but a continental one. This view can be challenged by the recent report
of the genus Maireana, formerly known only from Australia and hypothetized as the result of a
recent long-distance dispersal event (Teillier & Taylor 1997). The contemporary discovery of the
genus in the coast of Atacama region (Marticorena 1997) 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
6.3). On the other hand, the existence of four endemic angiosperm genera and 20 endemic species
reinforces the view of an old floristic history not just explicable by recent migration events.
108
Noteworthy, in his thesis about the biogeography of Brassicaceae, Brüggemann (2000) suggest
that the American Lepidium can have originated via long-distance dispersal from Australia or
North America, but on the other hand he suggests that the Pacific taxa have been translated by
Polynesian folks. Brüggemann (2000) mentions as example the absence of Lepidium from Juan
Fernández, which apparently was not reached by the Polynesian. The presence of the endemic
species L. horstii in the Desventuradas Islands is contrary to this suggestion.
Figure 4.22 Isla Alejandro Selkirk (Masafuera)
Figure 4.23. Isla Robinson Crusoe (Masatierra) (from Skottsberg
1920-1956)
4.5.3 Archipiélago de Juan Fernández
The archipelago, located between 667 and 850 km from the continent (figure 4.4), is formed by
two main islands and one islet:
- Alejandro Selkirk Island (33°46’ S / 80°47’W) (also known as Isla Más Afuera), 850 km from
the American continent. Its highest elevation is Cerro Los Inocentes (1.380 m a.s.l.) (figure 4.22).
- Robinson Crusoe Island, (33°38’ S / 78°51’ W) (also known as Isla Más a Tierra), located at
around 667 km from the continent. Its highest peak is Cerro El Yunque (915 m a.s.l.). Close to
Robinson Crusoe there are two islets: Islote Juanango, and Santa Clara, this latter located 1 km
southwest of Robinson Crusoe (figure 4.23).
The archipelago is world wide known as having been the scenary for the real history that inspired
the novel of Robinson Crusoe. Therefore the island Masafuera was renamed in honour of the
Schottich seaman Alejandro Selkirk, who survived 4 years and 4 months on Masatierra. Since its
official discovery in 1574 by the pilot Juan Fernández (maybe even before), Masatierra was an
obligate anchor place for seamen and bucaneers after trespassing the Cape Horn, and therefore
dozens of treasure histories characterize the island. But several botanists have long advice that the
real treasure of this archipele is in its unique plant world.
The flora of the archipelago has been long a subject of interest to botanists (e.g. Gay 1832, Johow
1896, Skottsberg 1920-1956, Muñoz Pizarro 1969, Stuessy et al. 1984b, Marticorena et al. 1998b,
109
Danton et al. 2000, Danton & Perrier 2003). The native vascular flora comprises 56 families
and 110 genera. The best represented family are the Poaceae (11 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 3.6), and 104 endemic
species (Stuessy et al. 1998a). Unfortunatelly the combining invasion of browsing animals and
continental plants place the island’s native flora at a competitive disadvantage (Dirnböck et al.
2003), so that at least 75% of the endemic flora is high endangered (Cuevas & van Leersum 2001,
Stuessy et al. 1998b). Swenson et al. (1997) reported a number of 227 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 1974). Oceanic islands are often explained as geologically new territories
and Juan Fernández is not the exception: Isla Robinson Crusoe has been dated at ca. 4 mya, Isla
Alejandro Selkirk at 1-2 mya, and Isla Santa Clara at 5.8 mya (Stuessy et al. 1984a). 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. (2006) 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 (1925, 1956) and he therefore suggested that the origin
of the island flora should be find in alternative palaeoscenarios. Skottesberg revised and discussed
all available evidence in floristic and faunistic elements, in geotectonics of the Pacific, and in the
continental Tertiary flora and came to the conclusion that the Fernandezian flora is not a one of
oceanic but of continental nature. Skottsberg therefore proposed a tentative sketch on the history
of the Fernandezian flora that is in concordance with Brüggen’s Tierra de Juan Fernández, an old
submerged landmass west of today South America (Brüggen 1950, Skottsberg 1956, p. 394; see
figure 5.12).
Skottsberg’s critical vision has been systematically oversimplified by modern authors, while
emphazising that he noted the close relationship with the American flora (e.g. Crawford et al.
1990, Bernardello et al. 2006). Skottsberg certainly recognized the floristic relationship between
the islands and the American continent, but he also noted the closest relatioship with the western
110
Pacific and especially with Australasia. Indeed, several genera not found in continental Chile
show a wider distribution in Australasia, like Coprosma (Rubiaceae), Arthropteris (Oleandraceae),
Haloragis (Haloragaceae) or Santalum (Santalaceae). The analysis of floristic elements of the
Juan Fernández flora shows a 14% of neotropical genera and 11% are australasiatic genera. Most
important is the cosmopolitan element (28%), and also the pantropical (17%) the endemic (14%)
and the antitropical elements (12%) (figure 4.24).
17%
28%
11%
14%
14%
4%
1. Pantropical
2. Australasiatic
3. Neotropical
4. Antitropical
5. South-temperate
6. Endemic
7. Cosmopolitan
12%
Figure 4.24 Floristic elements Juan Fernández
Furthermore, including Juan Fernández (JF) into the similarity analysis done between Chilean
regions (section 4.2) results in a closer floristic relationship between JF/MAG than between JF
and the other Chilean regions (table 4.6, figure 4.25).
Table 4.6. Floristic similarity between regions including Juan Fernández
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
0,29
1527
0,21
3227
0,54
750
0,27
2450
0,34
1700
0,18
2261
0,16
792
0,12
895
0,08
1439
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:
111
a) The Magallanes biota had once a more northward distribution, till central Chile, and from
there it reached the islands via long-distance dispersal. The presence of south-temperate
elements in the Fray Jorge fog forest at 30°40’ has been long suggested as an evidence
of the more widespread temperate forests during the Cenozoic. But the remnants of these
forests are still represented in the coast of BIO and there is no apparent reason why the
relation BIO/JF stays in the trend of similarity/distance, contrary to MAG/JF.
b) An alternative explanation for the floristic similarity between JF and MAG is a direct
connection of the two land masses. This explanation is contrary to current geological
consensus, since the islands seem to be geologically too young for this type of explanation.
The islands are located on top of the Juan Fernández Ridge, that controls the tectonic and
geological evolution of the southern Andes at 33°-34°S since the Tertiary (Yáñez et al.
2001). The seafloor age assigned by Müller et al. (1997) to the seafloor offshore Chile
between 18°S and 40°S, based on magnetic anomalies and relative plate reconstructions,
ranges from 20 to 48 mya. Juan Fernández rests on seafloor dated at around 20 to 33
mya.
0.6
COQ/BIO
0.4
ANT/COQ
ANT/BIO
0.2
Jaccard similarity
0.8
1.0
Some early geologists believed in a former land west of South America occupying the whole
Pacific (Burckhardt 1902, Beloussov 1968), or at least a portion of it as a Transandiner Kontinent
close to the South American coast (Muñoz Cristi 1942, Cañas 1966, Illies 1967). Brüggen (1950,
JF/BIO
0.0
JF/COQ
0
BIO/MAG
COQ/MAG
L
JF/MAG
ANT/MAG
JF/ANT
2000
4000
6000
8000
10000
Distance [km]
Figure 4.25 Floristic similarity between different regions in Chile: Antofagasta (ANT), Biobío (BIO),
Coquimbo (COQ) and Magallanes (MAG). The Jaccard similarity index as related to geographical
distance within Chile (squares), and between Juan Fernández and Chilean continental regions (circles).
The lines represent trend curves as an exponential curve. The relation JF/MAG (filled circle) is considered
as an outlier and is not included in the trend curve. The arrow show the possible position of the relation
JF/MAG following the trend (see discussion in the text).
112
p. 59) proposed the name Tierra de Juan Fernández for this larger continental land mass west of
current South America. Miller (1970) analysed different possibilities for the disappearing for this
land and concluded that the Ozeanisierung of Juan Fernández Land occurred in Late Tertiary, and
that this land was not at all at the same location of today’s Juan Fernández archipelago (Miller
1970, p. 934). From floristic similarity as analysed in the present thesis, this land should have
been existed much more to the South. To fit in the trendline of the relation similarity/distance
shown by other Chilean regions, the Juan Fernández islands (Juan Fernández Land) could be
located at the same longitude (80°W), but at 48°S instead of 33°, i.e. 15 degrees latitude or around
1.650 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 (69 mya). This has been corroborated
recently by the analysis of Wikström et al. (2001): 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 85 mya.
In Lammers et al.’s (1986) 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. 1999), thus suggesting that the Lactoridaceae were
widespread across the Southern Hemisphere
during the Late Cretaceous (Lammers et al.
1986). “Differences between Lactoripollenites
and Lactoris pollen imply that these represent
different clades within the Lactoridaceae or
that the former evolved into the latter genus
elsewhere in the Southwest Pacific region prior
to its migration onto Masatierra in the PlioPleistocene (Macphail et al. 1999). Crawford
et al. (2001) 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 4.26 Possible position of the hypotheaized
of long-distance dispersal” Crawford et al. Tierra de Juan Fernández, according to floristic
2001, p. 189).
similarity at the genus level (see figure 4.25).
113
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. 2003).
Stuessy et al. (1984a) were aware of the earlier view from Brüggen (1950) and Skottsberg (1956),
but they gave much value to the Potassium-argon dating. Thus most of the papers dealing with
the evolution of the Fernandezian flora in the last two decades start from the 4 mya date. The
dating of Santa Clara puts another question on the problem, since this little islet seem to be almost
two million years (5,8 +-2,1 mya) older than Robinson Crusoe. Stuessy et al. (1984a), 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 1981b). Also Stuessy et al. (1984b) recognize that the islands
could have been much more extense and have been rapidly eroded during the last 4 million years
(figure 4.27).
A
B
C
Figure 4.27 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. 1984b).
114
5 Palaeogeography: insights into the Evolution of the Chilean Flora
115
March-December 2006
116
5 Palaeogeography: insights into the Evolution of the Chilean Flora
Stuessy & Taylor (1995) suggest (based on van der Hammen & Cleef 1983) that there are three
principal sources of information for interpretating the origin of a flora:
a) evidence of geomorphological changes in the landscape;
b) the macro- and microfossil record;
c) biogeographical relationships between extant floras.
Stuessy & Taylor (1995) further mention the three principal geological influences that shaped the
present Chilean flora:
a) continental movements (due to plate tectonics);
b) oceanic transgressions over the continental surface;
c) the Andean uplift.
5.1 Continental movements: fragmenting the Earth’s surface
Plate tectonics is the current paradigm in geology (and related sciences like biogeography), after
the general acceptance of Alfred Wegener’s Kontinentalverschiebungs theory. The Earth’s surface
seems to rest on twelve major tectonic plates that interact at their borders. Under the paradigm of
plate tectonics, the floral history of southern South America is intrinsically related to the geologic,
tectonic and climatic history of the Gondwana continent.
The term Gondwana derive from the non-marine sedimentary rocks exposed in a series of grabentype basins of the Indian peninsula, that relate this land to the rest of Gondwanic landmasses
concentrated today in the southern hemisphere (McLoughlin 2001). 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 1944). Gondwanan has become synonymous with
the southern hemisphere biota, however, McLoughlin (2001) notes that the distinctive Indian
Gondwana sedimentary sequences and fossils were deposited while the southern continents were
united to Laurasia (Permian to early Cretaceous).
Several attempts have been made to relate the biogeographical information with the break-up of the
Gondwana continent (e.g. Villagrán & Hinojosa 1997, McLoughlin 2001, Sanmartín & Ronquist
2004). 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 119-105 mya, and the split between Antarctica and southern
South America (32-28 mya) (table 5.1).
Paleoreconstructions for Gondwana landmasses’s lateral movements since the Late Devonian
(360 mya) is presented in figure 5.1. The reconstruction has been done with the program Timetrek
v. 3.1 (Cambridge Paleomap Services Ltd.).
117
Table 5.1 Dating major palaeogeographic events = 3 stage break-up of Gondwana
Major
separation
events
Age
(mya)
Period
Possible cause
Source
(W Gondwana /
E Gondwana)
180150
Early
to Late
Jurassic
Breakup associated with development of a
series of deepseated mantle plumes beneath
the extensive Gondwanan continental crust
(e.g in S Africa (ca. 182 mya) and the
Transantarctic mountains (c. 176 mya)
Scotese &
McKerrow 1990,
Storey 1995,
Reeves & de Wit
2000
Africa – South
Am separation
119105
Early
Cretaceous
Opening South Atlantic Ocean, due to the
emplacement of Plume-related ParanaEtendecka continental flood basalts in Brazil
and Namibia (137-127 Mya). Final break-up
of Africa and S Am was completed only at 80
mya)
Turner et al.
1994, Jones 1987,
Barker et al. 1991,
Grunow et al 1991,
MacLoughlin 2001
West AntarcticaS Am
32-28
Palaeogene/
Oligocene
Beginning at ~35-30,5 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 120 mya,
and closer again at 80 mya).
Lawver &
Gahagan 1998
360 mya
250 mya
200 mya
145 mya
Figure 5.1 Palaeoreconstructions of Gondwana since Late Devonian (360 mya). Program TimeTrek v.
3.1., Cambridge Paleomap Services Ltd.
(trial version for download disponible at http://www.the-conference.com/CPSL/timetrek.htm).
118
100 mya
80 mya
60 mya
34 mya
10 mya
0 mya
Figure 5.1 (continued)
119
McLoughlin (2001) 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 5.2). 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 5.2 Distribution of the Permian Glossopteris flora in Gondwana (after McLoughlin
2001). Palaeo-biomes after Willis & McElwain 2002
Late Triassic (c. 200 mya)
Tropical summerwet
Subtropical desert
Winterwet
Warm temperate
Cool temperate
Dicroidium palaeoflora
Figure 5.3 Distribution of the Triassic Dicroidium
flora in Gondwana (after McLoughlin 2001). Palaeobiomes after Willis & McElwain 2002
Dicroidium leafs (www.rosssea.info/geology.
html)
120
121
200145
5. Jurassic
pteridospermconifer floras
Jurassic
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.
Seed-fern floras progressively replaced lycophyte floras.
Emergence of first arborescent plants: Lepidodendron and progymnosperms.
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).
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.
More diverse floras dominated by Dicroidium (see also Herbst et al. 2005)
(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.
250205
4. The
Dicroidium
flora
Triassic
Permian
300255
3. The
glossopterid
flora
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 world’s biota
during the Permian-Triassic extinction..
Carboniferous
330
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.
Late Devonian
370
Herbaceous to shrub-sized lycophytes attributed to Haplostigma, Leclercqia,
Archaeosigillaria and Protolepidodendron.
Fossils of Zosterophyllum and the lycopod Baragwanathia
present in both hemispheres suggest little floristic
provincialism (McLoughlin 2001), but several authors
recognize at least five biogeographical units (see Willis &
McElwain 2002)
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.
Free-sporing, herbaceous forms occupying moist habitat (Coastal areas, river flats,
delta swamps, lake margins) = Rhynopsida, Zosterophyllopsida, Trimerophytes and
lycopod fossils.
Silurian
– Early
Devonian
Palaeoenvironmental conditions
Composition
Period
Middle
Devonian
Age
(mya)
~430390
390
2. Arborescent
lycopod and
seed-fern floras
in a cooling
climate
1.
Cosmopolitan
early land plant
floras
Floras
Table 5.2 Gondwanan floras: five major phases of development before the appearance of angiosperms (sensu McLoughlin 2001)
For the Late Mesozoic (i.e. Cretaceous) and the Cenozoic, several models for the history of
the vegetation and climate have been proposed. Romero (1986, 1993) proposed three main
palaeofloristic types: a Neotropical palaeoflora, a Mixed, and an Antarctic palaeoflora. Troncoso
& Romero (1998) refined this classification and recognized twelve palaeofloristic types. Hinojosa
& Villagrán (1997) recognized five paleofloras (Tropical, Antarctic, Mixed, Sutropical mesic, and
Sutropical xeric), later renamed as Gondwanic, Subtropical Gondwanic, Mixed, and Subtropical
Neogene palaeofloras (Hinojosa 2005). The palaeoscenarios proposed by Romero (1993), Hinojosa
& Villagrán (1997), Troncoso & Romero (1998), and Hinojosa (2005) are summarized in table 5.3
and illustrated in figure 5.4, on the base of Time Trek palaeoreconstructions and palaeobiomes as
proposed by Willis & McElwain (2002).
A
B
Late Cretaceous (c. 70 mya)
Warm temperate
Tropical summerwet
Cool temperate
Subtropical desert
Tropical palaeoflora
Mid-latitude desert
Nothofagus expansion
Early Eocene (c. 50 mya)
Warm temperate
Figure 5.4 Evolution model for the Chilean flora on
the base of Time Trek palaeoreconstructions and the
palaeobiomes sensu Willis & McElwain 2002) (see
also table 5.3)
122
Tropical everwet
Cool temperate
Subtropical desert
Subtropical summerwet
C
Early Oligocene (c. 30 mya)
D
Cold temperate / arctic
Tropical everwet
Glacial
Tropical palaeoflora
Warm / cool temperate
Cool / cold temperate
Antarctic palaeoflora
Mixed palaeoflora
E
Miocene (c. 10 mya)
Pliocene (5 - 2 mya)
Warm temperate
Tropical everwet
Cool temperate
Arctic
Winterwet
Glacial
Tropical everwet
Cool temperate
Arctic
Desert (hyperarid)
Glacial
Desert (arid)
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 2002) (see also table 5.3) (for the
Pliocene see also Dowsett et al. 1999 paleoreconstruction)
123
124
CampanianMaastrichtian
85-65
3. Tropical
Gondwanic (sensu
Hinojosa 2005)
Palaeocene
Late Cretaceous
(Cenomanian to
Coniacian)
99-85
2. Neotropical
(without
Nothofagus)
65-55
Early
Cretaceous
145-100
1. Primitive
angiosperms
(with
Nothofagus)
sensu Troncoso &
Romero 1998)
Period
Age (mya)
Palaeofloras
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.
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 30°S.
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.
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)
Composition
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.
Warm and tropical conditions dominated in W
Gondwana.
Six global biomes are recognized at the end
of the Cretaceous (Willis & McElwain 2002),
five of them to be found in S Am (figure 5.4
A)
Mild “icehouse” world. Cool temperate forests
covered the polar regions.
Palaeoenvironmental conditions
Table 5.3 Cretaceous to Neogene evolution of the floras of South America, mainly from Romero (1993), Hinojosa & Villagrán (1997), Troncoso & Romero (1998),
Willis & McElwain (2002), Hinojosa (2005).
125
23-5
5-2
7. Subtropical
Neogene (more
xerophytic)
40-23
5. Mixed (sensu
Hinojosa 2005)
6. Subtropical
Neogene
55-40
4.
Subtropical
Gondwanic (sensu
Hinojosa 2005)
Pliocene
Miocene
Oligocene
Eocene
Fragmentation of former continue subtropical forests. Origin of disjunctions
between central Chile and NE Brazil.
Australantarctic cold element is replaced by more mesic element.
Development of Mediterranean-type sclerophyllous vegetation in Central
Chile. First record (Cerro Los Litres, 33°18’) 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.
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.
New families like Araliaceae, Guttiferae, Meliaceae, Sterculiaceae,
Euphorbiaceae, Bignoniaceae, Malpighiaceae, Brassicacae, Solanaceae,
Asteraceae, Chenopodiaceae, and Ericaceae.
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).
Global mid-Miocene warm climatic optimum.
Well-defined climatic belts stretched from pole
to equator (figure 5.4 D)
General cooling. Separation of Australia
from Antarctica c. 37 mya, origin of the
circumantarctic ocean circulation and the E
Antarctic glaciation.
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).
Table 5.3 (continued) Cretaceous to Neogene evolution of the floras of South America, mainly from Romero (1993), Hinojosa & Villagrán (1997), Troncoso & Romero
(1998), Willis & McElwain (2002), Hinojosa (2005).
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 14-11 mya (Kvasov & Verbitski 1981), at ~30-21 mya
(Brown et al. 2006), or ~41 mya (Scher & Martin 2006). These last authors suggest that the 41
mya deepening of the passage coincides with increased biological productivity and abrupt climate
reversals. These authors agree with Brown et al. (2006) 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. (2006).
Antarctica was covered by forests from the Permian onwards (Taylor et al. 1992). 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. 2001). 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. 2003 and references therein). To this impressive
record it is possible to add Late Cretaceous fossil flowers related to extant Winteraceae (Eklund
2003).
These Antarctic forests were ultimately eradicated due to the global cooling of climate during
the Tertiary (Francis & Poole 2002). 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 2004). The extinction may have followed the mid-Miocene warm interval at c. 17 Ma or a
mid-Pliocene warm interval at about 3 Ma (Ashworth & Cantrill 2004). 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. 2002). In southern South America there was
a constant exchange of neotropical and Antarctic floras throughout the Cenozoic (Hinojosa &
Villagrán 1997, 2005; Troncoso & Romero 1998) (figure 5.4).
5.2 Oceanic transgressions over the continental surface
Riccardi (1988) proposed several phases of transgression-regression from the Late Jurassic
(Kimmeridgian [155 mya]) to the Late Cretaceous (65 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 155 mya, central-W Argentina and Central and N Chile
had gone through a regional uplift that established continental conditions. By ~150 mya, the
126
area was again downwarped. A marine back-arc basin extended in a NNW direction, from about
40°S to the latitude of Copiapó and perhaps Antofagasta (figure 5.5). Transgressive-regressive
sequences were deposited till ~140 mya, during a period of rapidly rising sea-level (Mitchum
& Uliana 1985, quoted by Riccardi 1988). 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
1986, as quoted by Riccardi 1988). 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 1988).
The transgressions since the Late Cretaceous [83 mya] have been revised by Donato et al. (2003)
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 & Villagrán (1997) considers this wide transgression having affected only
E South America. Martínez-Pardo (1990) suggested Central Chile could have been affected by
transgressions at 19-10 mya, in concordance with the global Miocene warming (Zachos et al.
2001).
30°
36°
42°
48°
54°
Figure 5.5 A marine back-arc basin extended from
about 40°S to the latitude of Copiapó/Antofagasta at
Early Cretaceous (145-130 mya) (from Riccardi 1988).
127
Figure 5.6 Late Oligocene-Early Miocene transgressions
that possibly affected south-central Chile between 35°48° (redrawn from Donato et al. 2003)
5.3 The Andean uplift
Stuessy & Taylor (1995) 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 65 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. 2003, Mutke &
Barthlott 2005). Several recent phylogenetic studies further support a direct association between
the diversification of Andean plants (e.g. Hughes & Eastwood 2006, Smith & Baum 2006) and the
major episodes of Andean uplift, from the early Miocene (about 20 mya) to the Pliocene (about 3
mya) (sensu Hooghiemstra & van der Hammen 1998).
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 1995).
5.3.1 Final uplift of the Andes and Atacama hyperaridity
Earth scientists have attempted to determine the elevation history of the Earth’s surface by 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. 2006). Analysing palaeobotanical data, Gregory-Wodzicki (2000) concluded that the Altiplano-Puna had attained no more than a third of its modern elevation of 3.700
m asl by 20 mya and no more than half its modern elevation by 10.7 Ma. These data imply surface
uplift on the order of 2.300–3.400 m asl since the late Miocene at uplift rates of 0.2–0.3 mm/yr.
Geomorphological evidence, i.e. lahar deposits in the Coastal Cordillera of central Chile (33°40’34°15’S) suggest an Oligocene–Miocene uplift of the Andes (Encinas et al. 2006).
Recently Ghosh et al. (2006) 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 1.03 +-0.12 mm per year between ~10.3 and ~6.7
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 2003, proposed a proto-Central Andean mountain range placed between 15 and 9 mya).
The rapid uplift model (from 0 to 4000 m asl since the Middle/Late Miocene) proposed by Ghost
et al. (2006) counters former geological evidence and is therefore criticized by Sempere et al.
(2006) (see the response of Eiler et al. 2006).
According to Villagrán & Hinojosa (1997), 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
128
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 (2002), 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 25 mya (Dunai et al.
2005), 14 mya (Alpers & Brimhall 1988) to 3 mya (Hartley & Chong 2002). Houston & Hartley
(2003), 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 15 to 10 mya (Middle Miocene).
On a recent paper, Hartley et al. (2005) further propose that “the sedimentary succession in the
Atacama Desert records deposition under an arid to semiarid climate from the late Jurassic (150
mya) to the present day. 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 (1) the continentality effect (enhanced by the Gondwanan
landmass), and (2) 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.
(2005) of erosional and depositional surfaces analysed via remote sensing. Ochsenius (1999) 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. (2003) suggest that western South America has been an arid region well over 35 000
years ago, and analysis of pollen assemblages of rodent middens also suggest that hyperariditiy
has prevailed during the Pleistocene-Holocene, the last 50 000 years (Maldonado et al. 2005).
5.4 The last 30 000 years: surviving the Ice Age
The Pleistocene is characterized by several cycles of glaciation/deglaciation. The last glaciation
cycle is known as the Llanquihue (LLG) glaciation. At the time of middle Llanquihue glaciation,
cool, humid interstades on Isla Grande de Chiloé with Subantarctic Evergreen Forest was
increasingly replaced by parkland under progressive cooling after 47,000 [14]C yr BP (Heusser
129
et al. 1999). A closed-canopy of North Patagonian Evergreen Forest established by 12,500 [14]C
yr BP. Later, after ca. 12,000 until 10,000 [14]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 [14]C years
of the Pleistocene at midlatitude in the Southern Hemisphere.
During the tardiglacial, between 29,400 and 14,450 [14]C yr BP, the advance of the glaciers seem
to have devasted 2/3 of the actual area of the southern forests, specially in the regions Aisén,
Magallanes, insular and continental Chiloé, and the central and southern Andes (Villagrán &
Hinojosa 2005). Periglacial effects like solifluction and glaciofluvial activity should have also
affected the Andes, longitudinal depression, and coastal Cordillera between 39°-43°, affecting
principally the Valdivian forest taxa (Veit 1994,
Denton et al. 1999, Heusser 2003) (figure 5.7).
36°
42°
The current disjunct range of several species in both
cordilleras is a relict of a formerly wider distribution
(before the cold period 30.000-14.000 yBP),
as shown by the relict presence of Fitzroya and
Pilgerodendron in the central depression, Villagrán
et al. 2004).
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. (2004) and Fickert et al. (2007). 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 5.7 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. 1999, Heusser 2002).
were considered. This hypothesis is concordant with
the results of Premoli et al. (2000), 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 2002).
Pleistocene and Holocene changes have disrupted species ranges, extirpated local populations, and
changed selective pressures (Premoli et al. 2000), but it is doubtful that they affected speciation
130
processes. Some authors have emphazised the role of last glaciations in speciation, but others call
this a ‘failed paradigm’ (e.g. Klicka & Zink 1997). 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 1962). Now the theory
of mantle plumes22 (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. (2005) 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 2003)23.
Figure 5.8 Mantle plumes, www.seismo.unr.edu/
ftp/pub/louie/class/plate/; www.seismo.unr.edu/.../
class/100/interior.html
131
The assessment of Pratt (2000) 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
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” 24. Dickins et al. (1992) 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 (2002) 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 (1992). 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 1992, p. 203). 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 (1999).
Cretaceous
has
also
been recently challenged
due to biogeographical
and geological reasons.
“Palaeomaps, it seems, are
like the weather: If you
Figure 5.9 Alternative palaeoreconstructions (Dobson 1992)
132
don’t like the alleged size or placement of a pre-Cenozoic ocean, just wait a while. It will change”
(McCarthy 2005 b).
Land bridges
The idea of former land bridges in the site of today’s oceans, contrary to the current paradigm of
plate tectonics, is not an old one, and like a circular idea, is constantly re-presented and re-rejected.
Probably the first one who proposed and illustrated former lands between today’s continents
was Hermann von Ihering (1907), 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
5.10). Also Croizat (1952) 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 2003), the pantropics (Zhou et al. 2006) and between Tertiary disjunct
relict floras from Eurasia – North America (Milne 2006).
Figure 5.10 Land bridges proposed by H. von Ihering (1907)
133
Figure 5.11 Land bridges as proposed by Leon Croizat (1958)
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 80’s, under the umbrella of vicariance biogeography the idea was revitalized
(Nur & Ben-Avraham 1977, 1981a; Kamp 1980). The hypothesis was firmly rejected by several
Figure 5.12 Former land areas in the present Pacific and Indian Oceans (from Dickins et al. 1992, 1996)
134
authors like Cox (1990). Dickins et al. (1992) 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 1988, as quoted by Grande 1994). 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 1994, p. 76).
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 (1927) and Hilgenberg (1933), the theory loosed attention during the 20
century, in spite of being still defended by some researchers (e.g. Carey 1988, Vogel 1990). The
theory was reanalysed specifically from a biogeographical point of view by Shields (1997, 1998)
and more recently by McCarthy (2003, 2005a). It has been very criticised by biogeographers
like Cox (1990) and Briggs (2004). Mainstream textbooks treat the idea in different forms: Cox
and Moore (2005), being consequent with Cox (1990), do not mention the hypothesis in their
Biogeography. But Lomolino et al. (2006) dedicate two pages to discuss the pros and cons of
the expanding Earth hypothesis (Lomolino et al. 2006, pp. 248-249). 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 (2007).
Figure 5.13 Expanding Earth hypothesis and
a close Pacific ocean (after Shields 1998, McCarthy 2003). See also http://www.expandingearth.org/
135
Bonus track: modeling expanding earth, Paola pregnant on 15.12.2004 (top); same on 15.03.2007 (bottom)
136
6 Phylogeny of the Chilean Plants / Conflicts in Systematics and Biogeography
137
March-December 2006
138
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 1997, Huelsenbeck et al. 2000, Britton et al. 2002, Sanderson 2002, Thorne &
Kishino 2002). There has been also an increasing number of taxonomically identifiable fossils
(Friis et al. 2005). In spite of the datings have given widely different results, more recent studies
tend to converge on similar ages (Sanderson et al. 2004, Bell & Donoghue 2005). Especially
datings of the lower nodes within the angiosperms, have given much older ages than those obtained
from the fossil record (e.g. Wikström et al. 2001).
Molecular dating methods currently in use are classified by Rutschmann (2006) in three main
classes:
1. methods using a molecular clock and one global rate of substitution (e.g. linear
regression, character-based maximum likelihood clock optimization);
2. methods that correct for rate heterogeneity (e.g. linearized trees, local rates methods);
3. 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 2006).
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:
1. 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 2005];
139
2. substitution numbers on branches are converted into measures of relative time within
the phylogeny, technique known as rate smoothing [Sanderson et al. 2004];
3. conversion of relative ages into actual ages, assigning an actual age to at least one node
of the phylogeny [e.g. Wikström et al. 2001]. 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. (2005), after discussing
the difficulties and uncertainties in obtaining a stem group age for the eudicots, finished their
paper asking if the results could not be rather an artifact from constraints or method: “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 [2001]. 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 2003, and references therein]. Other scientists suggest that
these events are real and that we should trust the fossil record [Benton et al. 2000]”.
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. (2005) 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
(2005b) did a deep revision of the whole molecular biogeographic program, and is one of the most
downloaded paper from the journal Cladistics. But in spite of being vastly cited, Heads’s concerns
are not yet seriously incorporated in most recent papers (Heads pers. comm.)
“Ultimately, we may well conclude that accurate divergence time estimates require multiple
140
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 2004).
In a recent opinion article Pulquério & Nichols (2007) 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.
(2006) propose a method that do not impose unproven assumptions about the pattern in clock-rate
variation among lineages. The answer to Pulquério & Nichols’ initial question but still seems to be
“we do not yet know”, and they further concluded “We await the more rigorous type of assessment
with some nervousness, given that we suspect they might reveal that many past studies placed too
much confidence in simple molecular clock analyses, and that their conclusions should thus be
revisited” (Pulquério & Nichols 2007).
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 20 years or so of the DNA revolution (Palmer
et al. 2004). 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 (2003), Palmer et al. (2004), Pryer et al. (2004), Davies
et al. (2004), and Stevens (2001 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. 2004; Palmer et al. 2004). Living euphyllophytes belong
to two major clades (figure 6.1): seed plants (spermatophytes) and monilophytes (Kenrick &
Crane 1997, Nickrent et al. 2000, Pryer et al. 2004).
Monilophytes, ferns s. str., include most former groups recognized as ‘fern and fern allies’.
Monilophytes have been dated back to the Late Devonian [ca. 370 mya]. Chilean representatives
are classified in 21 families and 47 genera (appendix A).
141
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 250.000 and 300.000 species (Thorne 2002; but see Scotland &
Wortley 2003). In Chile the seed plants are represented by 41 orders, 154 families and 763 genera
(table 2.2, 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. 300 mya] (Stewart and Rothwell 1993, Hill 2005). However, exact relationships
among these lineages and the pattern and chronology of divergence remain unclear, despite the
recent accumulation of molecular data sets to address the question (e.g. Magallón & Sanderson
2002, Burleigh & Mathews 2004). The earliest known seed plants have been reported from the
Late Devonian of West Virginia (Gillespie et al. 1981). Gnetales and modern conifer families
appeared in the Triassic to Jurassic, and angiosperms in the limit Jurassic/Cretaceous (Stewart and
Rothwell 1993, Crane 1996). From the Permian through the late Jurassic many seed plant lineages
went extinct, including lyginopterids, medullosans, Callistophytaceae, glossopterids, Cordaitales,
and Voltziales (Stewart & Rothwell 1993), and their relationships with extant groups remain
poorly resolved. During the Cretaceous and Cenozoic, the diversity of all surviving seed plant
lines except angiosperms decreased (Knoll 1984, Crane 1985).
The Chilean flora is lacking extant cycads or Ginkgoales, but several fossil taxa have been
described (Troncoso & Herbst 1999, Herbst & Troncoso 2000, Torres & Philippe 2002, Leppe &
Moisan 2003, Herbst et al. 2005, Nielsen 2005). Conifers are represented by one order, 3 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 2003). The fossil record of the angiosperms extends back at least to the early
Cretaceous, conservatively 130 mya (Crane et al. 2004). Molecular dating has but pushed this age
to [179-158 mya] the Early – Middle Jurassic (Wikström et al. 2001).
The most basal Chilean clades in the angiosperm tree are the following (proposed ages by
Wikström et al. 2001) (figure 6.1):
- Ceratophyllum [155 – 140 Mya] Late Jurassic, limit Jurassic/Cretaceous.
- Monocots [154 - 139 Mya] Late Jurassic, limit Jurassic/Cretaceous.
- Piperales [149 - 137 –Mya] Late Jurassic, limit Jurassic/Cretaceous, Piperaceae [96 – 90 Mya]
Late Cretaceous; Lactoris/Aristolochia [97 – 85 Mya] Late Cretaceous; Peperomia/Piper [43 – 41
Mya] Eocene.
- Laurales [142-133 Mya] Early Cretaceous, Monimiaceae splitting at 91-85 Mya (Late Cretaceous),
Lauraceae splitting from Gomortegaceae/Atherospermataceae at 87-80 Mya (Late Cretaceous),
Peumus/Hedycarya [78 – 71 Mya] Late Cretaceous, Gomortega/Laurelia [51-39 Mya] Eocene.
Winterales (alt. Canellales) split from Magnoliales at 131- 127 Mya (Early Cretaceous),
Winteraceae25 from Canellaceae at 99 Mya (limit Early/Late Cretaceous), Drimys/Belliolium [2429 Mya] Oligocene.
142
Jurassic
180
160
Early Cretaceous
140
120
Late Cretaceous
100
80
Paleogene
60
40
Neogene
20
P
Malpighiales
Oxalidales
Fabales
Fagales
Cucurbitales
eurosids I
Celastrales
Rosales
Zygophyllales
eurosids II
Malvales
Brassicales
eurosids
Myrtales
Sapindales
Tapiscia
Aphloia
Ixerba
Crossomatales
Geraniales
Vitaceae
eudicots
Saxifragales
Berberidopsidales
Dipsacales
Escallonia
Apiales
Lamiales
Gentianales
Oncotheca
Garryales
euasterids I
Solanales
asterids
Asterales
Aquifoliales
euasterids II
Eremosyce
Ericales
Cornales
Caryphyllales
Dilleniaceae
Santalales
Gunnerales
Trochodendraceae
Buxaceae
Sabiaceae
Proteales
Ranunculales
Piperales
Chlorantales
Austrobaileyaceae
Illiciaceae
Schisandraceae
Nymphaeaceae
Amborellaceae
180
160
140
120
100
80
60
40
20
143
P
magnoliids
Laurales
eumagnoliids
Ceratophyllum
Monocots
Magnoliales
Winterales
Figure 6.1 Angiosperm chronogram calibrated against the
geological time-scale (adapted
from Wikström et al. 2001).
In grey taxa not represented
in Chile.
Within the Monocots, results from Janssen & Bremer (2004) suggest that considerable monocot
diversification took place during the Early Cretaceous, with most families already present at the
Cretaceous–Paleogene boundary. The crown nodes of Araceae, Petrosaviaceae, Orchidaceae,
Arecaceae and Dasypogonaceae have been estimated to date back to the Early Cretaceous
(Janssen & Bremer 2004). Liliales diverged from their sister –group in the Early Cretaceous, 117
mya and the extant lineages diverged from each other ca. 82 mya; Poales at 115 +- 11 mya (MidCretaceous) (Bremer 2000). Bromeliaceae also appear to date back to the Cretaceous (Linder &
Rudall 2005), but the uncertainties in dating the Bromeliaceae are considerable (Bremer 2002).
Janssen & Bremer (2004) found an unexpected crown node age of 111 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 2004).
Basal Eudicots have been recently dated back to the Early Cretaceous: Ranunculales estimated at
131-147 (Wikström et al. 2001) or at 120 mya (Anderson et al. 2005), Proteales at 144-130 mya
(Wikström et al. 2001) or at 119 mya (Anderson et al. 2005).
Core Eudictos: Gunnerales has been dated at 127-116 mya, Santalales at 118-111 mya, Saxifragales
at 121-111 mya, Caryophyllales at 111-104. 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 124-114
mya (Wikström et al. 2001), the split between Berberidopsis and Aextoxicon at 100-90 mya (limit
Early/Late Cretaceous).
Rosids have been dated back at 109-100 mya, eurosids I at 101-95 mya, eurosids II at 109-100
mya.
For asterids, Wikström et al (2001) obtained consistently younger age estimates, mostly to 10 to
20 my for major groups and orders compared to estimates by Bremer et al. (2004). As example,
the crown node age in the analysis of Bremer et al. (2004), is 128 my whereas Wikström et al.
(2001) estimated it to 117 to 107 my. The crown node age of Asterales was estimated to 96 my by
Bremer & Gustafsson (1997) using rbcL sequences and calibration with fossil Asteraceae pollen,
and to 93 my (Bremer et al. 2004). 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
1997).
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6.3 Vicariance v/s dispersal in the Chilean Flora
As already mentioned in section 4.1, 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 1981, Humphries & Parenti 1999),
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. (2006) consequently found that the disjunct distribution of extant
species of Tiquilia is the result of at least four long-distance dispersal events from North America to
South America. Taken the dispersalist universe as framework, Moore et al. (2006) do not mention
alternative palaeogeographic hypothesis, as listed by Constance (1963). But taken seriously the
concerns already expresed in section 6.1, 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. 2005).
Recent geophysical research seems to support geographical scenarios suited for long-distance
dispersal upon the southern seas (Muñoz et al. 2004). 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. 2001). 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
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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 (1999) challenged the hypothesized long-distance
dispersal explanation proposed by Swenson & Bremer (1997) for Abrotanella (see also Wagstaff
et al. 2006).
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 1944, p. 305-306).
But researchers tend to ignore or minimize empirical evidence: “Hoffmannseggia fruits and seeds
have no obvious adaptations for external animal dispersal and no one has ever recorded their
being eaten by birds. Nevertheless, we believe that bird dispersal is the most likely explanation
for the repeated pattern of long-distance dispersal from South to North America” (Simpson et al.
2005).
“Hätten die Wandervögel die ihnen oft zugeschriebene Bedeutung für Verbreitung der
Pflanzen, so würden die Zugstraßen der Vögel als deren Fäcalstraßen sich floristisch
ebenso darstellen, wie etwa die prähistorischen Handelstraßen aus den Fundstücken
kartographisch rekonstruierbar sind. Von dem ist aber keine Rede.“ (von Ihering
1893).
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 6.1, 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 1994). 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.
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Table 6.1 Evolutionary processes (dispersal v/s vicariance) proposed for several Chilean taxa
Families or groups
Genera or group
Evolutionary process
Sources
Apocynaceae
-
Gondwanan origin for the family
Potgieter & Albert 2001
Apiaceae
Oreomyrrhis
Origin in Eurasia and subsequently
dispersal to North America and southern
Pacific Rim.
Chung et al. 2005
Araliaceae
Pseudopanax
Gondwanan vicariance between
Australasia-South America and LDD to
Hawaii
Mitchell & Wagstaff 2000
Araucariaceae
-
Gondwanan origin
Setoguchi et al. 1998
Asteraceae
Microseris
Colonization Aus from N Am
Vijverberg et al. 1999
Asteraceae
Microseris
Vicariance, widespread ancestor
Grehan 2007
Asteraceae
Abrotanella
Asteraceae
Abrotanella
Dispersal between Australasia – South
America
Vicariance
Swenson & Bremer 1997,
Wagstaff et al. 2006
Heads 1999
Asteraceae
Hypochaeris
Dispersal from NW Africa across the
Atlantic Ocean for the origin of the South
American taxa
Tremetsberger et al. 2005
Atherospermataceae
Laurelia
Initial diversification at 100–140 mya,
probably in West Gondwana, early
entry into Antarctica, and long-distance
dispersal to New Zealand and New
Caledonia
Renner et al. 2000
Berberidaceae
Berberis
South America-Old World disjunct
distribution due to Cretaceous vicariance
Kim et al. 2004
Boraginaceae
Tiquilia
three independent dispersals from North
to South American
Moore et al. 2006
Brassicaceae
Cardamine
Bleeker et al. 2002a
Brassicaceae
Rorippa
Brassicaceae
Lepidium
Cunoniaceae
Cyperaceae
Oreobolus
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
Ericaceae
-
Laurasian in origin.with following
dispersals
Kron & Luteyn 2005
Fabaceae
Hoffmannseggia
Simpson et al. 2005
Fabaceae
Sophora
Gentianaceae
Gentianella
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 2,7 mya
Gesneriaceae
Coronantheroid
Burtt 1998
Gunneraceae
Gunnera
Gondwanic origin, Coronantheroid
Gesneriaceae (Asteranthera, Mitraria,
Sarmienta) relicts
Early vicariance followed by recent
dispersals
Gunneraceae
Gunnera
Late Cretaceous vicariance
Fuller & Hickey 2005
147
Bleeker et al. 2002b
Mummenhoff et al. 2001
Bradford & Barnes 2001
Chacón et al. 2006
Hurr et al. 1999, Peña et
al. 2000
Von Hagen & Kadereit
2001
Wanntrop & Wanntrop
2003
Table 6.1 (continuation)
Lauraceae
Beilschmiedia/
Cryptocarya
Gondwanan vicariance (diverged from
ancestor about 90+- 20 mya.
Chanderbali et al. 2001
Lycopodiaceae
-
Vicariance, Permian origin of the genera
Wikström & Kenrick 2001
Monimiaceae
Peumus
Vicariance at 76 mya, former continuous
range from Chile across Antarctica and
the Kerguelen plateau to
Madagascar
Renner 2004
Monocots (major
groups)
Nothofagaceae
-
Gondwanan vicariance
Bremer & Janssen 2006
Nothofagus
Swenson et. al 2001,
vicariant main massings of the four
subgenera compatible with allopatric
differentiation and no substantial
dispersal
Heads 2006
Nothofagaceae
Nothofagus
Vicariance Am-Aus subgen Fuscospora,
dispersal in Lophozonia
Knapp et al. 2005, Cook &
Crisp 2005
Plantaginaceae
Hebe
at least two instances of transoceanic
LDD from Australasia to South America.
Godley 1967, Wagstaff &
Garnock-Jones 1998
Plantaginaceae
Ourisia
origin in the central Chilean Andes
followed by consecutive dispersal to the
southern Andes and dispersal(s) to the
northern Andes and Australasia
Meudt & Simpson 2006
Poales
-
Bremer 2002
Polemoniaceae
Gilia
Dispersal California to South America
Morrell et al. 2000
American origin and dispersals into
Australia
Applequist & Wallace
2001
Portulacaceae
Proteaceae
-
Gondwanan; Mid-Cretaceous divergence
between major groups
Hoot & Douglas 1998
Primulaceae
Primula
Ancestral haplotype gave origin to S Am,
Euro and N Am lineages.
Guggisberg et al. 2006
Ranunculaceae
Caltha
Schuettpelz & Hoot 2004
Restionaceae
Apodasmia
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
Linder et al. 2003
Rhamnaceae
Discaria
Probably Gondwanan relict
Richardson et al. 2000
Rubiaceae
Coprosma
Vicariance
Heads 1996
Rubiaceae
Coprosma
Africa as ancestral area followed by longdistance dispersal into the Pacific
Anderson et al. 2001
Tetrachondraceae
Tetrachondra
LDD New Zealand – South America
Wagstaff et al. 2000
Winteraceae
-
migration from tropical Gondwana to
Antarctica and Australia throughout
South America
Barreda & Archangelsky
2006
Zygophyllaceae
Fagonia
dispersals between South America and
southern Africa
Beier et al. 2004
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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 1981, Nelson 1981, Hinojosa 2005).
New analytical tools like DIVA analysis or parsimony-based tree fitting have the supposed capacity
to discriminate between vicariance and dispersal (e.g. Sanmartín & Ronquist 2004). Recently
Sanmartín et al. (2007) 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 Zealand–South
American events.
The authors concluded that “Only once we have a better understand of dispersal process will it
be possible to apply realistic estimates of dispersal frequency and asymmetry to biogeographical
reconstructions“ (Sanmartín et al. 2007). But this seems also valid for a better understanding of
vicariance processes, that frequently have been oversimplified as a several steps break-up model,
obviating the complex geological and biotic nature of a region (see next section).
6.4 Sloppy biogeography v/s harsh geology?
“Las hipótesis de relaciones entre áreas formuladas por la geología no tienen validez
intrínseca superior a las que se formulen a partir de datos biológicos, es decir, de los
cladogramas de áreas. A partir de este principio, de por sí totalmente aceptable, algunos
autores concluyen que las hipótesis geológicas no pueden utilizarse ni para corroborar
ni para refutar las hipótesis biogeográficas, en contradicción patente con uno de los
principios básicos del cladovicariancismo: la idea del paralelismo entre la evolución de
la vida y la de la Tierra” (Zunino & Zullini 2003, p. 290).
Recently some authors call for avoiding the circular logic in confronting biogeographical
hypothesis with geological evidence (e.g. Renner 2005, Waters & Craw 2006). “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 2005, p. 552).
The problem seems to be: how much geological evidence is necessary to accept a biogeographical
hypothesis?
149
In the words of Heads (2005b): “assuming a priori that any particular geological event, such as
the break-up of Gondwana, is relevant to biogeography is a fatal flaw of much biogeography, both
dispersalist and vicariance… In fact, a great deal of evidence suggests biogeographic patterns
involving New Zealand, New Guinea, New Caledonia etc. were determined by earth history
events both prior and subsequent to the break-up of Gondwana”. Croizat (1958) was the first to
call for this independent view of biogeography as a discipline with own methods and tools. He
proposed that biogeographers should not base all their study in geological “well established”
hypothesis. Biogeography, as a mature and independent discipline should be able to develop their
own theories and views to compare with the geological theories. That’s the way Wegener (1915)
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 2005b).
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 2005).
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
150
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 2001). 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
2001), New Zealand (Engler 1882, Craw 1988), the Subantarctic Islands (Michaux & Leschen
2005) Tasmania (Heads 1999), New Guinea (Heads 2002) and in southern South America (Crisci
et al. 1991, Katinas et al. 1999).
Back then to alternative palaeogeographies (section 5.6): “I am fully aware that my reconstruction
of Pacifica [continent] calls into question certain widely accepted dogmas of geology, and that for
my failure to accept them I am accused of naïveté. However, I suspect that in the final analysis it
will be the geologist who is proved naïve for his gullibility in swallowing the dogmas – hook, bait,
and sinker” (Melville 1981). Or in the words of Sluys (1994): “... 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 (1995) 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 1995, see a recent analysis of Nothofagus by
Knapp et al. 2005, and Heads 2006).
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 (1998) 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
151
seems to stem from an attachment to this Biological Species Concept (BSC) as canonical truth
and that knowledge of breeding compatibility has everything to do with reconstructing phylogeny.
The BSC states that a species is a population of organisms that can interbreed and produce fertile
offspring. It’s easy to see why it causes alarm, because it obviously can’t be applied to fossils. It’s
also invalid when applied to asexual organisms (of which there are not few…)26. 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 1990).
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 2003, p. 18). Other concepts seem to be more suited for the biogeographic analysis, like
the Evolutionary Species Concept (ESC), in the sense of Wiley (1978).
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 (1963). But researchers like Bush (1975, 1998) 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 1998). “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 1998, 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
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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
[2002] who included the following in his seven great questions of systematic biology: What is
a species?, How many species are there?, Where are the Earth’s species distributed?, and How
have species’ distributions changed over time?” Heads find equally myopic speciescentrism in
Hubbell’s (2001) Unified Neutral Theory of Biodiversity and Biogeography (Heads 2005a, p.
102). In spite of efforts made by authors like Salomon (2001), 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 (1973). 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 33°
to 42°S. On the other hand, Magellanica group shows a case of true speciation (i.e. the splitting
of an ancestral stock into several species by means of geographical isolation = allopatry). The
Magellanica group’s evolution is related to the last glaciations, in which the species suffered
reduction of the population size and the presence of barriers to gene flow. These results suggests
that the taxa living in higher elevations (above the timberline) have been pushed to rapidly
adapted to this new environments. But phyletic evolution sounds paradoxical, since it considers
evolutionary stasis for a long period, with an abrupt change conducent to speciation only during
the Pleistocene (sensu Simpson 1973 for Prenanthoides group).
Hershkovitz et al. (2006) 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. 2006, p. 11).
In support of Simpson’s findings are the results with another Asteraceae genus, Hypochaeris,
which suggest that populations from the coastal Cordillera are older than the Andean ones
(Muellner et al. 2005).
The speciation mode described as phyletic evolution by Simpson (1973) has been more recently
treated as anagenetic speciation by Stuessy et al. (2006). In anagenetic speciation “a founder
population arrives on an island and simply diverges through time without further specific
differentiation” (Stuessy et al. 2006). 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 1998). Carlquist (1974) mentioned Juan Fernández as one of the best scenarios for
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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 1974, p. 206). Sanders et al. (1987) attempted to characterize this habitat differences
(primarily edaphic factors) but were not able to demonstrate clear-cut differences. On the contary,
Stuessy et al. (1990) 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. 2006).
Noteworthy, Hughes & Eastwood (2006) 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 (2006) 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 2001,
p. 235).
As proposed by Smith & Baum (2006) for a clade of Andean Solanaceae, the modes of speciation
could be a continue mixture of (micro)allopatric and sympatric episodes of hybridization. It
appears that these events have patially obscured the underlying divergent phylogenetic history,
having clouded the branching pattern in some parts of the tree. Some authors like Dickerman
(1998) have developed modified parsimony methods to detect hybridization or horizontal gene
transfer. Descent patterns in a phylogenetic system with a single hybrid event can be described as
the sum of two gene trees, each describing the history of part of the genetic material composing
the system. Systems with more than one hybrid event will require a larger set of trees. This
set of gene trees is called a phylogenetic forest. Dickerman’s (1998) method returns a reticulate
hypertree in which some taxa have more than one immediate ancestor, as one would expect if
hybridization or horizontal transfer has occurred. Ané and Sanderson (2005) developed another
method that returns either a single tree or a collection of trees. In the opinion of Dickerman (1998),
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
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1998). In the same line, Posada & Crandall (2001) and Winkworth et al. (2005) 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 (2006) 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 2004]”. Hershkovitz (2006) found for the Andean Portulacaceae, “the unexpected
similarity of the marker sequences in morphologically divergent taxa may itself result from
hybridization”. He therefore suggests that “…interpretation of molecular data derived from one
or a few samples per taxon and few DNA sequences must allow for the possibility of past gene
flow…[]…If ecological instability and novelty of habitats favour hybrid success, then current and
historical environmental parameters must be considered favorable to hybridization of Andean
Portulacaceae”.
The existence of x Margyracaena, an hybrid genus between an endemic species (Margyricarpus
digynus) and the introduced genus Acaena in Juan Fernández, is a good example of the real
possibilities of the hybridization process.
6.5.2 Monophyletic v/s polyphyletic
L. Constance, while analizing the antitropical (amphitropical) disjunct pattern recognized “on the
common assumption that each species, genus, or natural family has subsequently radiated in all
directions to occupy suitable territory, we should be led to expect a series of relatively continuous
distributions of such taxa over the earth’s surface, much in the sense of Willi’s concept of Age
and Area” (Constance 1963, p. 109-110). He continues “…Attempts to explain the broken ranges
of living or extinct organisms have led to postulate polytopic origins, parallel and convergent
evolution, orthogenesis, shifting poles and an oscillating equator, long-distance dispersal, land
bridges, and continental drift, among others”. Constance ruled out some explanations: “…
modern bicentric or vicarious species can, then, scarcely be the relicts of widely and continuously
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distributed ancestral populations. Nor am I able to accept the idea of a polyphyletic or polytopic
origin of these, although such an explanation presented no difficulty to the pre-Darwinian Special
Creationists. In their more modern forms, postulated polytopic origins simple push the problem
backward to the presumed existence of a more continuously distributed ancestor, which was
able to give rise to “look-alike” descendents in several or many portions of its range [Briquet,
Süssenguth, Pachosky, Rosa, Schröter, Du Rietz, Wulff]… (Constance 1963, p. 111)27 . 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 1921, p. 4). Guppy advanced the theory of differentiation and pointed out: “if
the differentiation hypothesis is correct, no natural order could have been developed on the lines
implied by the Darwinian theory, which, as interpreted in recent works, begins with the variety
and terminated with the order, a process that reverses the usual methods of Nature…Yet such
a process, as is there implied is common enough in the plant world, but it accounts not for the
natural orders but for the oddities of plant forms …It is here termed a specializing process in
contrast with that of differentiation; but it is the differentiation process that has been the principal
determining cause of diversification in plants” (Guppy quoted by Bews 1921, p. 3).
In this sense, Hu (1950) 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. 2002) and
Nair’s Triphyletic theory (Nair 1979). While carpels, double fertilization, and flowers are viewed
by some to be too complex to evolve more than once from totally unrelated seed plant lineages,
evidence from studies of phenotypes of homeotic genes suggest otherwise. The activities of insects
may have hastened and stimulated the development and evolution of seed plant reproductive
organs multiple times, in many evolutionary lines (Miller 2006). 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
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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 1936, p. 102).
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 2005b). Furthermore, Stuessy do not ruled out the
possibility of a “polyphyletic origin of life on Earth” (Stuessy 2006, p. 37).
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 2004, McCourt et
al. 2004). Megafossil evidence for the land plant crown group comprises taxa from the Middle
Silurian, about 420–430 mya (Kenrick & Crane 1997) or early Late Silurian (Wellman & Gray
2000). However, microscopic spores from the Ordovician have supposedly derived from parent
plants that were bryophyte-like if not in fact bryophytes (Wellman & Gray 2000, Wellman et al.
2003). The fossil-based age estimate for these microfossils is mid-Ordovician, about 475 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. (2001) challenged the more traditional view: analysing 50 protein sequences, the
authors estimated a Precambrian age for land plants of 703 +/- 45 mya. This result contrasts
sharply with palaeobotanical estimates (Kenrick & Crane 1997, Wellman et al. 2003). Sanderson
(2003) applied a penalized likelihood approach with two alternative and internal calibration
points, and different genes than Heckman et al. (2001). Sanderson (2003) results suggest the land
plant crown group to be of Ordovician age (483 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. (2001) results, pushing the land plant origin forward into
the Early Silurian (435 and 425 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. 2004). During the past 20 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 2001, Pryer et al. 2004)
(Fig. 1). 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 1993).
The most diverse of the monilophytes are the leptosporangiate ferns, a group of more than
11000 extant species. The earliest known occurrence of fossil leptosporangiate ferns is in the
Early Carboniferous (Galtier & Philips 1996); by the end of the Carboniferous six families were
present. In subsequent major radiations in the Permian, Triassic, and Jurassic, several families
with extant representatives replaced these Carboniferous families (e.g. Osmundaceae, Schizaceae,
Matoniaceae, and Dipteridaceae) (Rothwell 1987). The more derived polypod ferns dramatically
diversified in the Cretaceous, acompanying the great angiosperm diversification (Schneider et al.
2004).
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 370 mya) of West Virginia (Rothwell et al.
1989, Kenrick & Crane 1997).
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 150 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 2005).
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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
“Darwin’s abominable mystery” of botany (e.g. Crepet 1998, Bowe et al. 2000, Davies et al.
2004). Janssen & Bremer (2004) wrote: “From fossil evidence, a major radiation of angiosperms
is obvious in the mid-Cretaceous 130–90 mya [Lidgard & Crane 1990, 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. (2005) 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 (2005)
results indicate that widely divergent age estimates can result from the different methods (198–
139 mya), different sources of data (275–122 mya), and the inclusion of temporal constraints to
topologies. Most dates, however, are between 180–140 million years ago, suggesting a Middle
Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil
angiosperms by about 45–5 million years. These dates are consistent with other recent studies
that propose the hypothesis that the angiosperms may be older than the fossil record indicates.
Wikström et al. (2001) recognize “the calibrated phylogeny here is a working hypothesis and
should be viewed as such”. In Bremer’s opinion (2002): “The analysis by Wikström et al. (2001)
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 2002).
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. 2004, Friis et al. 2005, 2006). 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. 2005): “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. 2005). Contrasts among different morphology-based analyses can
159
be ascribed to different character selection, different sized data matrices, and disputes over
homology assessment (Crepet 1998). 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 1998).
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. 2002). 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. (2003) 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. (2003), specially correctly coding the leave form of Cabomba, still resolves the genus as the
most basal in the angiosperm phylogeny (Nixon pers. comm.). In Crepet’s words “I have learned
that the discovery of a few specimens of a new fossil taxon is seldom a unique event--there will
be new specimens of Archaefructus and the kinds of characters critically needed are also the
kinds likely to be preserved. Given the potential informative value of this taxon and the recent
pace of innovation in studies of angiosperm systematics and paleobotany today, I predict that the
great “abominable mystery,” [the origin of the angisperms] with us for over 100 years, will not
last another 10” (Crepet 1998). Crepet also recognized (2000): “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 (2005): “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 56 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 2005).
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In view of the difficulty of finding ancestors for the angiosperms, yet also considering their
sudden appearance and explosive evolutionary success, Stuessy (2004) 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
100 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
Figure 6.2 Traditional view of vascular plant evolution
(from Cornet 2002)
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angiosperms
undoubtedly
originated long before the Cretaceous
period. The specialised character and
astonishingly modern facies of many
Cretaceous angiosperms confirm our
belief in an antiquity of angiosperms
antedating by many millions of years,
probably by several geologic periods,
the first appearance of recognisable
pioneers of the present ruling dynasty
in the modern world (Seward 1933,
cited by Takhtajan 1981, p. 122).
Arldt (1938) and Berry (1920) 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 1947, Thomas 1947, Axelrod
1961, Zimmermann 1959). Others like
Golenkin (1927, quoted by Takhtajan
1981) 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 1960, p. 1735).
In concordance with Takhtajan (1961) 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. (2006), 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 6.3 Alternative view of angiosperm origins (from Cornet 2002)
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
162
between unrelated seed plants, over more than 200 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 2006).
Results obtained by Martin et al. (1989, 1993) suggest that separation of monocotyledonous and
dicotyledonous lineages took place in Late Carboniferous (300 mya), and Wolfe et al. (1989)
place the age of angiosperm origin at 230 mya. Sanmiguelia, a Late Triassic fossil described by
Brown (1956) has been interpreted by Cornet (1989) 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 (2002) asks “why is there still so much resistance to the recognition of pre-Cretaceous fossils
that could belong to the angiosperms or to their non-gymnospermous precursors if angiosperms
did not evolve from any known group of gymnosperms (i.e., Gnetales, Bennettitales, Cycadales,
Pteridospermales, and Coniferales)? Has the Cretaceous origin of angiosperms become a myth
perpetuated by fear of uncertainty? Why can’t paleobotanists be more honest and say that they
don’t have the answer yet to angiosperm origins, rather than bias the subject by saying “early”
angiosperms and “primary” radiation in the Cretaceous - or worse by shifting the age of possible
Jurassic strata to conform to a Cretaceous origin? If angiosperms evolved in the Triassic, Early
Cretaceous angiosperms would be middle age. And if the genetic roots of angiospermy extend all
the way back to the Silurian (see diagram above), that would make Cretaceous angiosperms “late
bloomers”.
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164
7 Back to Postmodern Biogeography
165
From the exhibition: Impressionen der Flora von Chile, A.M.M., Botanical Garden Erlangen, March-December 2006
166
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 1993, p. 321).
7.1 Biogeography as a social science
The monoboreal relic hypothesis was summarized by Thiselton-Dyer (1909, p. 311) 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 (1940, p. 244),
the monoboreal relic hypothesis was worked out in great detail with
regard to animals by Matthew (1915), who came to the conclusion that
“the principal lines of migration in later geological epochs have been
radial from Holarctic centres of dispersal” (Matthew 1915, p. 172), 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 1915, p.
W.D. Matthew
130). 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 1915, p. 263). Matthew’s views on biogeography were replicated and
advanced by G.G. Simpson and the fellows of the so called New York School of biogeography.
These ideas were nicely illustrated by means of caricatures accompanying Simpson’s book
Evolution and Geography (box 7.1). A discussion on Matthew’s biogeographic program has been
done by Nelson & Ladiges (2001) (see also an early review by Barbour 1916). A critique is to
167
Box 7.1 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
168
Figure 7.1 Griffith Taylor‘s map: Zones of migration showing the evolution of the races (1921).
varieties of the human race occupied the tropical zones… That Taylor’s map was part of a more
general cartographic discourse relegating the tropics to cultural insignificance is clear from two
related cartographic ventures. The projection that Taylor called upon to depict his racialised
conception of evolutionary history has earlier been used by his mentor William Diller Matthew in
1915 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
1915, p. 43). Livingstone
continues noting the
reprint of Taylor’s map
in Huntington’s book The
character of races, author
who otherwise wrote “the
people of the tropics are
in reality the children of
the human race. They
represent our primitive
ancestors… It is not to be
expected that such people
Figure 7.2 Matthew‘s map: migration and the distribution of the human
races (1915).
should ever rise very high
169
in the scale of civilisation” (Huntington 1924, as quoted by Livingstone 2002). 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
1925, p. ix). But in Livingstone’s opinion: “the cartography accompanied a discourse of climate’s
moral economy that flourished because it facilitated the elaboration of a moral topography that
was crucial to the project of the racial ideology, and because it employed scientific language to
diagnose and treat the sickness of a colonial regime” (Livingstone 1994). 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 (1980), Hettlage
(1982) and others as reviewed by Müller (1996).
Let’s just briefly revise the role of Darwin and his Origin of Species in the evolution of modern
geography. Livingstone (1993, p. 179) begins these reflections asking why Darwin is mostly
ignored in the history of geography, with the notable exception of Stoddart (1986) (we can add
Livingstone himself, and more recently Kennedy 2006). On the contrary, Darwin is a permanent
theme in biogeography, as the obligate reference point (e.g. Davies et al. 2004) or as the main
receptor of the critics (Croizat 1962). Darwinismus entered into geography through Friedrich
Ratzel (1844-1904), who visited Ernst Haeckel’s courses at Jena during 1869. Haeckel is known
as one of the most convinced defender of Darwin’s ideas (Livingstone 1992, p. 198). Ratzel’s
work was also greatly influenced by the naturalist, explorer and evolutionary theorist Moritz
Wagner. He was an early devotee of Darwin’s theory, but he believed that Darwin had failed to
appreciate the significance of migration and geographical isolation in the processes of speciation
(Wagner 1868). “Migration, isolation, space, and environmental determinism were all part and
parcel of the Wagnerian scheme of things. And it was precisely these themes that dramatically
surfaced in Ratzel’s new anthropogeographie… Ratzel’s Anthtropogeographie can best be read
as an attempt to situate the new science of human geography within the naturalistic framework of
Wagner’s Migratiosgesetz, which he portrayed as the [most] fundamental law of world history”
(Livingstone 1992, pp. 199-200). But Wagner was convinced that direct organic modifications in
response to environmental conditions could be affected and their benefits transmitted to succeeding
generations, adopting rather a Neo-Lamarckian position, that provided “the theoretical sustenance
on with the Ratzelian programme could thrive” (Livingstone 1992, p. 202). This Ratzelian
program has been vastly and differently reinterpreted and misunderstood (see conclusions chapter
in Müller 1996 and his claims against Stoddart 1986). Let’s allow Ratzel to speak himself about
his relation to Darwinismus, and we will find many elements that will be later crucial in the
development of biogeography28:
“Aber wenn man nun die wirkliche Verbreitung der Lebewesen ansieht, kann man
doch allen diesen Hilfsmitteln der passiven Wanderung nicht die große Wirksamkeit
zusprechen, die seit Darwin und Wallace so viele ihnen zugeschrieben haben“
170
„Da sich Länder und Meere auf der Erde ununterbrochen verschoben haben und noch
heute unter unseren Augen sich verschieben, so verändert sich also beständig der
Lebensraum für die wasserlebenden und ebenso die landlebenden Organismen“29.
„Das Suchen nach einen Mittelpunkt kann uns dabei offenbar nur verwirren. Besonders
bei der Erforschung des Ursprungs eines Volkes oder eine Rasse darf man nicht vergessen,
daß man weder nach Punkten (Ursprung) noch nach Linien (Wegen), sondern nach
Räumen oder Gebieten zu fragen hat“.
Ratzel’s (1901) 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 (1892),
as one of the founders of the discipline (see Ebach & Goujet 2006, 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 2005, p. 121).
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 1998, reviewed by Dawkins 1998). 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 (1989) 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 1995, p. 76). Dawkin’s 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 1995, 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
171
and the cooperative behavior of social animals, such as ants and monkeys (a vision similar to the
early one of Kropotkin 1902). 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 1972, Eldredge 2004). 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 2006).
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 1973, as
quoted by Livingstone 1992, p. 185).
In the same line are Gieryn’s (1999) Cultural Boundaries of Science, or Kohler’s (2002) Lanscapes
and Labscapes, in which the former explores how boundaries of science are sought out in complex
entanglements with political and cultural forces, and the latter questions if Clement’s famous field
ecology quadrats might be the reflection of Nebraska’s cultural landscape in the mind of the
ecologist (Anker 2003).
“a better explanation for the cultural authority of science lies downstream, when scientific
claims leave laboratories and enter courtrooms, boardrooms, and living rooms. On such
occasions, we use maps to decide who to believe—cultural maps demarcating science
from pseudoscience, ideology, faith, or nonsense…[]…Was phrenology good science?
How about cold fusion? Is social science really scientific? Is organic farming?” (Gieryn
1999)
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 1999).
In this respect also fits Cornet’s claims regards the neglecting of pre-Cretaceous angiosperm fossils
by the scientific community (section 6.6): “Funding is paramount for survival in the academic
172
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 2000]. 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
(2001) regards the media coverage gained by the APG group (section 1.3) and by Hamilton (2002)
regards the assumptions in current geologic paradigms like plate tectonics (section 5.5).
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 (1919) proposed the
genus Pereskia, due to its similarity to other woody flowering plants, the nearest cactus relative
to the other families. The most derived are the Cactoideae 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 1997, Applequist & Wallace 2001). 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 1961).
173
Box 7.2 Some notes on the biogeography and evolution of the Chilean Cactaceae
Chilean Cactaceae genera are classified under the following subfamilies and tribes.
Subfamily
Tribe
Genera
Maihuenioideae
Maihuenia
Opuntioideae
Cumulopuntia, Maihueniopsis, Miqueliopuntia, Pterocactus, Tunilla
Cactoideae
Trichocereeae
Echinopsis, Haageocereus,Oreocereus
Notocacteae
Austrocactus, Copiapoa, Eriosyce, Eulychnia, Neowerdermannia
Browningieae
Browningia
Pachycereeae
Corryocactus
As Edwards et al. (2005) 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 (1969) 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 (2002) 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
(1986) suggested a late Cretaceous origin of the group. Recent studies (Hershkovitz &
Zimmer 1997, Nyffeler 2002, Edwards et al. 2005), suggest that the cacti can’t be that
old because sequence divergence between major clades is limited. Edwards et al. 2005
assert that both Opuntioideae and Cactoideae originated in the central Andean region of
Peru, Bolivia, and northern Argentina. They appeal to the Andean orogeny as the cause of
diversification for many plant lineages, as early authors already recognized (e.g. Raven &
Axelrod 1974). Early uplift in the central Andean region (25–20 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 (1986), 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
2001) have their highest diversity on continents of the former Gondwana (Hershkovitz &
Zimmer 1997). This is generally taken as circumstantial evidence that the family Cactaceae
originated in South America (e.g. Buxbaum 1969). 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 1986). The
presence of a Rhipsalis species in tropical Africa, Madagascar, and Sri Lanka (Barthlott
1983, Barthlott & Taylor 1995) led some authors to propose that this distribution indicates an
old vicariance between South America and Africa (e.g. Backeberg 1942) or even an origin
of the cacti in the Old World (Croizat 1952). This would imply that the cacti originated
before the split of the southern continents during the late Cretaceous and that all other cacti
174
Box 7.2 (continuation)
Notocacteae
Neowerdermannia
Copiapoa
Eriosyce
Browningieae
! Browningia
Maihuenioideae
Pachycereae
$ Maihuenia
$ Corryocactus
Eulychnia
Austrocactus
$
$
$
Opuntioideae
Trichocereeae
Echinopsis
Tunilla
Cumulopuntia
# Miqueliopuntia
Maihueniopsis
Pterocactus
Haageocereus
Oreocereus
175
#
#
#
$
!
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 1983),
has been explained as the result of relatively recent long-distance dispersal by birds (Gibson &
Nobel 1986, Barthlott & Hunt 1993). Maxwell (1998) 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 1998).
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, 65–90 mya following the breakup of the western part of the
Gondwana supercontinent, has been favoured (e.g. Gibson & Nobel 1986, Mauseth 1990). 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 1997). Hershkovitz & Zimmer (1997) proposed a much more
recent origin of cacti in the Late Paleogene (30 mya). Nyffeler (2002) 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°
!
"
#"
"
#!
# "
! Copiapoa
$ Eriosyce
# Eulychnia
$ Neowerdermannia
! Austrocactus
30°
!
!
#
18°
$
!$
#!
#
#
#
#
#
#
30°
!
!
#
Trichocereeae
!
Echinopsis
$ Haageocereus
# Oreocereus
!
24° !
18°
!
!
24°
!
!!
!
!
!
!
!
!
30°
36°
36°
42°
42°
48°
48°
48°
54°
54°
54°
36°
$
42°
$
!
Edwards et al. 2005 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. 2005 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.
176
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 1997,
Applequist & Wallace 2001).
2. Some studies still consider Pereskia as a deep lineage (Nyffeler 2002, 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 2005).
3. 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).
5. Maihueniopsis appears to circumscribe a suite of early Opuntioid characters including
diminutiveness, early deciduous leaves and geophytism (Griffith 2004a).
6. 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. 5 in Edwards et al. 2005)
177
7.2 Biogeography toward a science of qualities
“If accepted, these views of language and science have tremendous implications. They
demolish religion, science, and any other means of “knowing” imaginable. Each person
is consigned to play their own language “game,” never truly understanding the extent
to which all thought is shaped and determined by the prison of syntax and grammar.
Nevertheless, despite its unattractiveness, Deconstruction and its cousin Postmodernism
do have a certain logic and appeal. They are, without a doubt, carefully arranged and
skillful critiques of the current means of knowing. Either way—whether Postmodernism
is nonsense or “truth”—the dominance of Modernist thinking seems severely challenged,
and there is little doubt that the reverence this culture has had for science will necessarily
change as a result of this opposition” (Kau 2001).
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 2004). 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. 2003, Morrone 2005) serve more for feeding the critics about the supposed
identity crisis of biogeography (Riddle 2005). Several attempts are currently under development
to find a more integrative research program in biogeography (e.g. Salomon 2001, Donoghue
& Moore 2003), 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 2004, p. 2). 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 1.2), showing real
possibilities for its renewal... just under a new name?
This is in concordance with Hengeveld’s Dynamic Biogeography (Hengeveld 1990). 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.
178
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 (1994) and Reason & Goodwin (1999) 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 1999). 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 1994).
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 (1995).
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 2000, García Azkonobieta 2005). This development will soon encompass systematic
problems in plant biology (e.g. Frohlich 2006 in the recent volume on monocots evolution).
This is a good sign toward the better integration between evolutionary biology, cladistics and
systematics. Goodwin classify this advances as a new biology: “The new biology is biology in the
form of an exact science of complex systems concerned with dynamics and emergent order. Then
everything in biology changes. Instead of the metaphors of conflict, competition, selfish genes,
climbing peaks in fitness landscapes, what you get is evolution as a dance. It has no goal. As
Stephen Jay Gould says, it has no purpose, no progress, no sense of direction. It’s a dance through
morphospace, the space of the forms of organisms” (Goodwin in Brockman 1995, p. 97)30
179
Did this development in evolutionary biology (i.e the new biology) already reach the new
biogeography? (in the sense of Lomolino & Heaney 2004). Are evolutionary scenarios been taken
explicitly in the biogeographical discussion? And if not, will this happen during the next decade?
Some authors are still skeptic about the recent developments: “…contemporary biogeography,
especially phylogeography, is simply boring to read because the mindless slaves that produce it
ignore Horace’s dictum sapere aude (Epistle I, 2) –”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 2000 years. Ancient
teleology is replayed every night on television, and mysterious means of dispersal and perfect
adaptations are extolled as wonders of nature” (Heads 2005a, p. 112). One possibility for the
sceptical is just turn off the switch (figure 7.3)… We just can, with all these concerns and conflicts
in mind, keep going, asking and (re)searching. Let’s take Reason & Goodwin’s positive view,
together with Gyerin pragmatic one: “…may the best science win…” (Gieryn 1999, title of
chapter 3).
Figure 7.3 Valparaíso (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 don’t want to speculate on wether this is good or
bad, but I believe that without discovery of more repeating patterns in systematics, those
of us interested in evolutionary process and its connection to phylogenetic pattern will
be stuck in a rut… Of course it is possible that we may never discover such constants in
biology, and a smoother synthesis of pattern and process will never be made, but I remain
an optimistic. I also unambiguously identify my optimism as a result of my beliefs rather
than of scientific evidence. Distinguishing between one’s beliefs and one’s scientific
evidence may be one of the most difficult aspects of evolutionary studies” (Grande 1994,
p. 79).
180
8 Conclusions: toward a biogeographic synthesis of the Chilean Flora
181
March-December 2006
182
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 1944, pp. 210-211).
•
The extant vascular flora of Chile is composed, under current knowledge (updated
09 February 2007) of 59 orders, 179 families, 813 genera, and about 4 333 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 29,6°S, while
the south-temperate and australasiatic elements show the southernmost average at
42,3°S and 41,5°S respectively. The other elements show an average distribution in
Central Chile between 32,2° and 36°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.
183
•
North and Central Chile belong to the neotropical floristic realm, while Central Chile
represents a core area of endemism or a panbiogeographic node.
•
Integrationg palaeobotanical and geological data under the current plate tectonics
paradigm, the most logic model for the Chilean flora is an interchange of tropical
floras with australasiatic floras, with later development of a subtropical more xeric
flora.
•
The geographic ranges of many Chilean plants have been reduced by the glaciations
of the Pleistocene, but it is doubtfull that this relative recent events had an impact on
speciation processes.
•
The Pacific offshore islands have been traditionally classified as oceanic islands,
but the floristic composition of Juan Fernández and its floristic similarity with the
Magallanes region, is a strong sign to consider it as a continental flora, maybe the
remnant of an ancient flora of an older terrestrial landmass to the West of current
South America (recognized by some geologists as the Pacifica lost continent).
•
Alternative palaeogeographic scenarios for the islands and for the continent as well,
should not be rouled out till much of the evidence is confronted.
•
Also alternative views in the evolution of plants (specially angiosperms) can change
dramatically our understanding of biogeography and the relation to palaeogeographic
events.
•
Conflicts in tectonic hypotheses, systematics, and molecular dating, directly affect any
proposal in biogeography. Being biogeography fragmented in the last decades into a
plethora of methods, a synthetical biogeography will be still elusive and murky for
decades. This synthesis will not come from the scientific consensus but from a deep
reanalysis of the conceptual and epistemological basis of the discipline, in connection
with new developments and debates in complexity theory and postmodern science.
•
The biogeography and evolution of the southern hemisphere’s flora will continue
fascinating naturalists as it has been the case for more than 250 years. The view from
the top of Cerro La Campana in Central Chile and from El Camote in Juan Fernández
releases the inspiration for an intuitive (re)search in qualitative biogeography.
•
The plant geography of Chile is (and will be ever) a work in progress.
184
“Trapped between fundamentalists, who believe they have found truth, and
relativists, who refuse to pin it down, the bewildered majority in between
continues to hope there is a truth worth looking for, without knowing how to go
about it or how to answer the voices from either extreme. We need a new Guide
for the Perplexed – a way of understanding and identifying truth which can
survive the postmodern era” (Fernández-Armesto 1997, p. 3).
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 2006).
“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 1989, p. 2).
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.
185
Footnotes
Hugh M. Raup (a botanist) did a review of the subject for the Annals of the Association of American
Geographers. Aside the title of the article, he widely use the term plant geography… Raup’s case as a
botanist publishing in a geographical journal shows graphically that the differences in the field are very
permeable.
1
2
Areology is also the study of planet Mars, as counterpart to Geology (www.duden.de).
Note that von Wettstein’s work can be seen as the precursor of the modern concept of allopatric
speciation.
3
Many people use the terms ‘systematics’ and ‘taxonomy’ as synonyms, but Stuessy (1979, 2006: 32)
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 2006, p. 37).
5
news in the English newspaper The Independent of 23rd November 1998, as quoted by
Endersby (2001)
6
„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 21 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 Croizat’s critic
on Hennig’s explanation of a progression rule while confounding the islands upon the example rests
(Croizat 1976).
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/~geo337/
crap.html]
10
“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”.
11
12
http://www.mnhn.cl/botanica/Herbario/index.html
13
Muñoz Pizarro (1966) considered also many naturalized taxa.
186
14
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.
15
“I thus hold divisions of the sphere by areas, suggested until now, for artificial systems…. they just
harmed science”
16
“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”
17
18
Croizat apparently took the term from Van Steenis (Croizat 1962)
19
Due to human pressure, Santalum fernandezianum is all but extinct from Juan Fernández (Islands).
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).
20
21
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. 1999).
22
23
Fischer, J.M. (2003) Beyond the Plume Myth www.newgeology.us
Also the website: “Problems with Plate Tectonics” http://ourworld.compuserve.com/homepages/dp5/
lowman.htm
24
Winteraceae have often been regarded as the ‘‘most primitive’’ extant family of angiosperms (Cronquist
1981, Endress 1986). The phylogenetic position based on molecular data indicates that the family is
nested in the Canellales (APG II 2003, Soltis & Soltis 2004).
25
26
Brazeau, M. (2005) http://lancelet.blogspot.com/
27
Constance does not give the reference to these authors’ papers, but their work is worth of attention.
It is a pity that Müller (1996) in his excellent revision of Ratzel’s program does not explicit analyze this
great piece: Der Lebensraum: eine biogeographische Studie (Ratzel 1901).
28
29
Note that this was written 15 years before the proposal of continental drift by Alfred Wegener.
30
Also available at http://www.edge.org/3rd_culture/bios/goodwin.html
187
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Appendix A: List of Chilean genera, geographic distribution, floristic elements
221
222
Asterids
Asterids
Monocots
Asterids
Monocots
Eudicots
Eudicots
Asterids
Asterids
Rosids
Monocots
Monocots
Alonsoa
Alopecurus
Aloysia
Alstroemeria
Alternanthera
Amaranthus
Amblyopappus
Ambrosia
Amomyrtus
Amphibromus
Amphiscirpus
Asterids
Agalinis
Alona
Eudicots
Aextoxicon
Eudicots
Filicopsida
Adiantum
Allionia
Rosids
Adesmia
Monocots
Rosids
Adenopeltis
Rosids
Asterids
Adenocaulon
Alisma
Asterids
Acrisione
Alchemilla
Asterids
Achyrocline
Monocots
Asterids
Acantholippia
Agrostis
Rosids
Acaena
Asterids
Rosids
Acacia
Agoseris
Asterids
Abrotanella
Asterids
Monocots
Aa
Ageratina
CLASE A
GENUS
Poales
Poales
Myrtales
Asterales
Asterales
Caryophyllales
Caryophyllales
Liliales
Lamiales
Poales
Lamiales
Solanales
Caryophyllales
Alismatales
Rosales
Poales
Asterales
Asterales
Lamiales
Berberidopsidales
Pteridales
Fabales
Malpighiales
Asterales
Asterales
Asterales
Lamiales
Rosales
Fabales
Asterales
Asparagales
ORDEN
Cyperaceae
Poaceae
Myrtaceae
Asteraceae
Asteraceae
Amaranthaceae
Amaranthaceae
Alstroemeriaceae
Verbenaceae
Poaceae
Scrophulariaceae
Solanaceae
Nyctaginaceae
Alismataceae
Rosaceae
Poaceae
Asteraceae
Asteraceae
Orobanchaceae
Aextoxicaceae
Pteridaceae
Fabaceae
Euphorbiaceae
Asteraceae
Asteraceae
Asteraceae
Verbenaceae
Rosaceae
Fabaceae
Asteraceae
Orchidaceae
FAMILIA
1
12
2
43
1
60
100
75
30
36
16
6
3
10
250
220
11
290
40
1
150
230
1
5
2
30
6
100
1200
20
27
N° SP
Canada, USA, Arg, Chile
Aus, NZ, S Am
Temp S Am: Arg, Chile
Canada, Mexico, SA, West Indies
1 sp. Calif, NW Mexico, Chile
Subtrop & temp, Euro, S Am, N Am, 16 sp Russia,
Tropics & subtrop, S Am, N Am, Aus, 1 sp Caucasus
Chile, S Brasil, Perú, Arg
Am: SW US to Chile & Arg
warm temp & subtrop regions & montane tropics
Trop Am Mexico, C Am, Chile, Bol, 2 sp S Afr
Chile
SW & C US, Mexico to Chile & Arg
temp N Hemishp, Euro, Middle asia, Australia, S Am
temp N hemis, Middle Asia, montane tropics, S Afr
temp & warm regions, montane tropics, JF, IP
9 sp W N Am, Mexico, 2 sp Chile, Arg
E USA, C & W S Am, West Indies
Trop, warm C & S Am, USA
Chile, Arg
Cosmop, esp Neotrops, Madag
Peru, Chile, Arg, S Brasil, TFuego
Chile
W NA, Mexico, Hawaii, Guatemala, Chile, Arg,
Himalayas, E Asia
Central Chile
Tropics Afr, Am, Madagascar,
Arid S Am, Arg, Chile, Bol
S Afr (1), New Guinea, Aus, Tasmania, NZ, subantarctic Is, Polynesia, Hawaii, California to SA, JF
Tropics, Subtrop, esp Afr y Aus
New Guinea, Aus, Tasmania, NZ, S Am (TFuego), JF,
Falkland Is.
Andes Colombia to Argentina
DISTRIB
ANT-P
AUS
W-NT
ANT-P
S-AMZ
W-NT
W-ANT
W-NT
ANT-P
W-ANT
W-ANT
ANT-P
W-NT
ANT-P
S-AMZ
W-ANT
AND
SPT
AND
TRACK
4
2
5
3
4
7
1
3
3
4
3
6
4
7
4
4
4
3
4
5
7
3
6
4
6
1
3
4
1
2
3
ELEM
1
1
2
2
1
2
2
33
3
4
1
6
1
1
3
24
2
1
1
1
6
132
1
1
2
1
3
19
1
4
1
SP
CHILE
-27,00
-35,40
-35,33
-17,58
-22,08
-33,23
-18,50
-20,33
-20,50
-31,50
-29,50
-18,83
-20,17
-33,43
-18,20
-31,00
-30,95
-25,00
-29,00
-30,67
-21,27
-18,20
-30,50
-36,83
-31,62
-18,23
-18,42
-22,00
-27,25
-43,00
-17,58
N_MAX
-55,00
-43,00
-46,83
-42,72
-30,65
-40,35
-28,50
-54,00
-30,50
-53,67
-36,67
-32,00
-23,50
-40,00
-18,23
-56,00
-56,00
-36,67
-30,00
-43,67
-45,40
-54,83
-36,83
-55,00
-45,50
-18,23
-30,00
-56,00
-37,90
-55,75
-24,50
S_MAX
-41,00
-39,20
-41,08
-30,15
-26,37
-36,79
-23,50
-37,17
-25,50
-42,58
-33,08
-25,42
-21,83
-36,72
-18,22
-43,50
-43,48
-30,83
-29,50
-37,17
-33,33
-36,52
-33,67
-45,92
-38,56
-18,23
-24,21
-39,00
-32,58
-49,38
-21,04
MittelW
28,00
7,60
11,50
25,13
8,57
7,12
10,00
33,67
10,00
22,17
7,17
13,17
3,33
6,57
0,03
25,00
25,05
11,67
1,00
13,00
24,13
36,63
6,33
18,17
13,88
0,00
11,58
34,00
10,65
12,75
6,92
Range
223
Asterids
Monocots
Magnoliids
Rosids
Eudicots
Eudicots
Asterids
Filicopsida
Filicopsida
Monocots
Asterids
Aristeguietia
Aristida
Aristolochia
Aristotelia
Arjona
Armeria
Artemisia
Arthropteris
Asplenium
Astelia
Aster
Asterids
Apium
Asterids
Asterids
Aphyllocladus
Argylia
Rosids
Aphanes
Eudicots
Eudicots
Antidaphne
Eudicots
Monocots
Anthoxanthum
Argemone
Monocots
Anthochloa
Arenaria
Asterids
Antennaria
Pinopsida
Eudicots
Anisomeria
Araucaria
Eudicots
Anemone
Rosids
Asterids
Androsace
Monocots
Rosids
Andeimalva
Araeoandra
Monocots
Anatherostipa
Arachnitis
Rosids
Anarthrophyllum
Monocots
Asterids
Anagallis
Apodasmia
Asterids
Amsinckia
Asterales
Asparagales
Aspleniales
Davalliales
Asterales
Caryophyllales
Santalales
Oxalidales
Piperales
Poales
Asterales
Lamiales
Ranunculales
Caryophyllales
Pinales
Geraniales
Liliales
Poales
Apiales
Asterales
Rosales
Santalales
Poales
Poales
Asterales
Caryophyllales
Ranunculales
Ericales
Malvales
Poales
Fabales
Ericales
Unplaced
Asteraceae
Asteliaceae
Aspleniaceae
Oleandraceae
Asteraceae
Plumbaginaceae
Santalaceae
Elaeocarpaceae
Aristolochiaceae
Poaceae
Asteraceae
Bignoniaceae
Papaveraceae
Caryophyllaceae
Araucariaceae
Vivianiaceae
Corsiaceae
Restionaceae
Apiaceae
Asteraceae
Rosaceae
Santalaceae
Poaceae
Poaceae
Asteraceae
Phytolaccaceae
Ranunculaceae
Primulaceae
Malvaceae
Poaceae
Fabaceae
Myrsinaceae
Boraginaceae
250
25
650
15
390
90
10
5
300
290
20
12
28
160
18
1
1
10
27
5
20
8
20
1
50
2
90
150
4
11
15
20
15
temp Eurasia, S Afr, Madag, Am, Hawaii
Masc, New Guinea, Aus, NZ, Polinesia, Chile
Subcosmop, Madag, JF
New Guinea, NZ, Juan Fernandez, Madag
temp Eurasia & W N Am, N + S Afr, S Am
SAm Andes to Tfuego, Euro, Russia
Temp S Am, Brazil
E Aus, Tasman, NZ, Arg, Chile
trop, subtrop, some in temp regions
N Am, Medit, temp Asia, tropics & subtropics
Colombia, Ecuador, Peru, Chile
S Peru, N C Chile & Arg, A. uspallatensis S Bol
N & S AM, WI, Hawaii
America, temp Eurasia, N Afr
New Guinea, Aus, New Caledonia, S Chile, SE Brasil
Chile
Chile, Arg
SE Asia, Malesia, New Guinea, Aus, Tasm, NZ, Chile,
Arg
Euro, Medit, N & S Afr, Madag, W & E Asia, Malesia,
Aus, N, C & S Am, JF, IP
Andes S Bolivia, N Chile, NW Arg
Am, Euro, Medit, Ethiop, Australia, Caucasus
W trop SA: S Mexico, Guatem, Costa Rica, Colom, Venez, Ecuad, Peru, Bol, Bras, Chile, Cuba, Haiti, P.Rico
temp & warm Eurasia, N Afr, montane tropics
Andes Chile Peru
Arctic & temp Euras, Andes S Am
Chile
Euras, Sumatra, S & E Afr, NZ, A multifida disj N Am
to Chile
N Hemisphere, China, few sp SA (Tfuego)
Chile, Peru, Bol
Arg, Bol, Chile, Ecuador, Peru
Andes Chile Arg
Euro, Russia, N & S Afr, Madag, W Asia, Himalayas, N
Aus, 2 sp. S Am
W US, W temp S Am
W-ANT
SPT
SPT
W-ANT
W-ANT
S-AMZ
AUS
AND
AND
W-NT
W-ANT
SPT
SPT
AND
W-NT
AND
W-ANT
W-ANT
W-ANT
AND
AND
ANT-P
4
2
7
2
4
4
3
2
1
7
3
3
3
4
2
6
5
2
7
3
7
3
7
3
4
6
4
4
3
3
5
7
4
4
1
7
1
2
1
3
1
2
3
1
11
4
7
1
1
1
1
8
1
3
1
7
1
1
2
6
1
1
4
5
2
2
-30,65
-40,17
-18,00
-17,75
-30,67
-31,00
-30,83
-27,50
-18,83
-29,50
-20,82
-18,48
-18,00
-37,50
-30,17
-33,00
-36,83
-19,80
-18,82
-33,40
-37,67
-36,50
-18,38
-31,00
-24,83
-30,83
-53,17
-28,58
-19,83
-30,47
-25,00
-22,25
-56,00
-56,00
-55,92
-53,00
-56,00
-54,50
-44,00
-33,48
-37,83
-36,45
-34,47
-35,35
-55,00
-40,05
-30,83
-50,00
-51,00
-54,93
-27,78
-37,17
-42,37
-56,00
-21,50
-53,13
-35,97
-53,15
-53,17
-36,90
-25,50
-52,67
-55,00
-53,00
-43,33
-48,08
-36,96
-35,38
-43,33
-42,75
-37,42
-30,49
-28,33
-32,98
-27,64
-26,92
-36,50
-38,78
-30,50
-41,50
-43,92
-37,37
-23,30
-35,28
-40,02
-46,25
-19,94
-42,07
-30,40
-41,99
-53,17
-32,74
-22,67
-41,57
-40,00
-37,63
25,35
15,83
37,92
0,00
35,25
25,33
23,50
13,17
5,98
19,00
6,95
13,65
16,87
37,00
2,55
0,67
17,00
14,17
35,13
8,97
3,77
4,70
19,50
3,12
22,13
11,13
22,32
0,00
8,32
5,67
22,20
30,00
30,75
224
Asterids
Asterids
Rosids
Eudicots
Eudicots
Pinopsida
Rosids
Monocots
Rosids
Filicopsida
Asterids
Asterids
Asterids
Asterids
Eudicots
Rosids
Rosids
Eudicots
Asterids
Magnoliids
Asterids
Asterids
Eudicots
Eudicots
Asterids
Monocots
Filicopsida
Asterids
Rosids
Asterids
Rosids
Eudicots
Asterids
Monocots
Asterids
Eudicots
Filicopsida
Asteranthera
Asteriscium
Astragalus
Atriplex
Austrocactus
Austrocedrus
Avellanita
Axonopus
Azara
Azolla
Azorella
Baccharis
Bacopa
Bahia
Bakerolimon
Balbisia
Balsamocarpon
Barneoudia
Bartsia
Beilschmiedia
Belloa
Benthamiella
Berberidopsis
Berberis
Bidens
Bipinnula
Blechnum
Blennosperma
Blepharocalyx
Blumenbachia
Boehmeria
Boerhavia
Bolax
Bomarea
Boopis
Boquila
Botrychium
Ophioglossales
Ranunculales
Asterales
Liliales
Apiales
Caryophyllales
Rosales
Cornales
Myrtales
Asterales
Blechnales
Asparagales
Asterales
Ranunculales
Berberidopsidales
Solanales
Asterales
Laurales
Lamiales
Ranunculales
Fabales
Geraniales
Caryophyllales
Asterales
Lamiales
Asterales
Apiales
Salviniales
Malpighiales
Poales
Malpighiales
Pinales
Caryophyllales
Caryophyllales
Fabales
Apiales
Lamiales
Ophiglossaceae
Lardizabalaceae
Calyceraceae
Alstroemeriaceae
Apiaceae
Nyctaginaceae
Urticaceae
Loasaceae
Myrtaceae
Asteraceae
Blechnaceae
Orchidaceae
Asteraceae
Berberidaceae
Berberidopsidaceae
Solanaceae
Asteraceae
Lauraceae
Orobanchaceae
Ranunculaceae
Fabaceae
Ledocarpaceae
Plumbaginaceae
Asteraceae
Gratiolaceae
Asteraceae
Apiaceae
Azollaceae
Salicaceae
Poaceae
Euphorbiaceae
Cupressaceae
Cactaceae
Amaranthaceae
Fabaceae
Apiaceae
Gesneriaceae
35
1
13
100
5
35
50
12
17
3
175
8
240
470
2
12
1
230
60
3
1
8
1
10
60
400
70
6
10
110
1
1
5
250
2000
8
1
temp, polar, trop montane
Arg, Chile
Andes, Arg, S Brasil
Mex to trop Am
Temp SA
tropics & subtropics
Pantrop, JF
Brasil, Uruguay, Paraguay, Argentina, Chile
trop & subtro S Am, WI
2 California, 1 Chile
Temp S Am: Chile, Arg
cosmop
Euras, N Afr, N & S AM
B. beckleri en Aus: Queensland & New S Wales, B.
corallina: Chile
S Chile-Arg, Patagonia
Andes Arg, Chile
pantropics, Mex to S Bras, Chile, Aus, NZ, Afr, Madag
Circumboreal, Eur, Medit, Afr, Andes
Arg, Chile
Chile
Peru, Bol, Chile, Arg
Chile
SW US, Mex, Chile
Trop & Subtrop, Hawaii, Madag, Aus
AM, esp S AM
Andes to temp S Am, Malvinas, Antarctic Islands
trop & warm, Euro, Asia, Madag, Tristan da Cunha,
Hawaii
8 temp Chile-Arg, s subtrop NW Arg, Bol, SE Bra,
Uruguay, JF
pantrops, IP
Chile
Arg, Chile
Arg, Chile
cosmop, Eurasia, Aus, ID
N Am, Andes, temp Eurasia, montane trop Afr
Mexico to Patagonia
Chile, Arg
W-ANT
S-AMZ
W-NT
S-AMZ
AND
ANT-P
W-ANT
AUS
W-ANT
AND
AND
ANT-P
W-NT
AUS
S-AMZ
W-ANT
W-NT
4
5
3
3
5
1
1
3
3
4
7
5
7
4
2
5
5
1
4
3
6
3
6
4
1
3
2
7
3
1
6
5
5
7
4
3
5
1
1
8
3
2
2
1
3
1
1
10
4
7
47
1
4
1
2
3
3
1
3
1
1
1
43
14
1
8
1
1
1
2
20
46
4
1
-50,50
-35,83
-36,00
-18,00
-49,15
-20,50
-33,75
-32,50
-33,42
-30,67
-25,00
-17,58
-25,02
-37,13
-46,58
-34,83
-32,55
-18,00
-29,82
-27,37
-18,00
-23,50
-21,27
-20,00
-17,58
-18,00
-18,18
-30,00
-33,83
-32,65
-33,78
-18,00
-17,58
-25,20
-35,60
-55,67
-42,70
-54,00
-39,00
-55,52
-37,00
-45,50
-41,30
-36,60
-55,97
-40,50
-37,17
-55,97
-37,67
-53,67
-40,75
-36,67
-36,50
-34,25
-29,83
-30,00
-30,00
-36,62
-30,50
-56,00
-55,50
-42,55
-46,00
-34,10
-40,22
-46,50
-53,33
-53,50
-37,83
-47,00
-53,08
-39,27
-45,00
-28,50
-52,33
-28,75
-39,63
-36,90
-35,01
-43,32
-32,75
-27,38
-40,49
-37,40
-50,13
-37,79
-34,61
-27,25
-32,03
-28,60
-24,00
-26,75
-28,94
-25,25
-36,79
-36,75
-30,37
-38,00
-33,97
-36,43
-40,14
-35,67
-35,54
-31,52
-41,30
5,17
6,87
18,00
21,00
6,37
16,50
0,00
11,75
8,80
3,18
25,30
15,50
19,58
30,95
0,53
7,08
5,92
4,12
18,50
4,43
2,47
12,00
6,50
15,35
10,50
38,42
37,50
24,37
16,00
0,00
0,27
7,57
12,72
35,33
35,92
12,63
11,40
225
Rosids
Asterids
Rosids
Eudicots
Monocots
Monocots
Eudicots
Asterids
Monocots
Asterids
Camissonia
Campsidium
Cardamine
Cardionema
Carex
Carpha
Carpobrotus
Castilleja
Catabrosa
Centaurea
Rosids
Caldcluvia
Asterids
Asterids
Calceolaria
Calystegia
Eudicots
Calandrinia
Monocots
Monocots
Calamagrostis
Calydorea
Asterids
Caiophora
Asterids
Rosids
Caesalpinia
Eudicots
Rosids
Bulnesia
Calycera
Monocots
Bulbostylis
Caltha
Asterids
Buddleja
Monocots
Eudicots
Browningia
Calotheca
Monocots
Bromus
Asterids
Monocots
Bromidium
Asterids
Rosids
Bridgesia
Calopappus
Monocots
Brachystele
Callitriche
Asterids
Brachyclados
Rosids
Asterids
Bowlesia
Calliandra
Monocots
Bouteloua
Asterales
Poales
Lamiales
Caryophyllales
Poales
Poales
Caryophyllales
Brassicales
Lamiales
Myrtales
Solanales
Asparagales
Asterales
Ranunculales
Poales
Asterales
Lamiales
Fabales
Oxalidales
Lamiales
Caryophyllales
Poales
Cornales
Fabales
Zygophyllales
Poales
Lamiales
Caryophyllales
Poales
Poales
Sapindales
Asparagales
Asterales
Apiales
Poales
Asteraceae
Poaceae
Orobanchaceae
Aizoaceae
Cyperaceae
Cyperaceae
Caryophyllaceae
Brassicaceae
Bignoniaceae
Onagraceae
Convolvulaceae
Iridaceae
Calyceraceae
Ranunculaceae
Poaceae
Asteraceae
Plantaginaceae
Fabaceae
Cunoniaceae
Calceolariaceae
Portulacaceae
Poaceae
Loasaceae
Fabaceae
Zygophyllaceae
Cyperaceae
Scrophulariaceae
Cactaceae
Poaceae
Poaceae
Sapindaceae
Orchidaceae
Asteraceae
Apiaceae
Poaceae
500
4
200
30
15
2000
6
200
1
62
25
15
17
10
1
1
17
200
11
300
13
230
56
150
8
90
90
11
150
5
1
22
3
15
40
Eurasia, N China, N & tro Afr, Aus, NZ, N Am
temp Eurasia, N Am, Chile
Euras, E N Am, C & S Am
S Afr, Aus, Tasman, NZ, Calif, Chile
trop & S Afr, Madag, japan, New Guinea, Aus, Tasm, NZ,
temp S Am
cosmop, JF
W N Am to Chile
Cosmop, temp N hemis & montane tropics, JF
Arg, Chile: C. valdivianum
W N Am, Mex, S Am temp
Cosmop, temp & trop hemis N & S, JF
Trop & Subtrop Am
Temp S Am, Brazil
Arctic & temp N Hemis, NZ, temp S AM
Arg, Bras, Chile, Uruguay
Chile
Cosmop
C & S Am, Afr, Madag, trop Asia
Philippines, Sulawesi, Moluccas, New Guinea, trp Aus,
S Chile Arg
AM from Mex to Tfuego
Am W USA to Chile
temp & subtrop reg & montane tropics
Peru, Bol, Arg, Chile, 1 sp Urug, Bras, Ecuad
Pantropics, subtropics, Arg, Madag, S Afr, Arabia
S Am: Venezuela Colombia disjunct to Chile-Arg, Brazil
E Asia, tro & subtrop AM, Afr
Peru, Bol, Chile
Eurasia, N & S Afr, montane trop Asia, Am, JF
S US & subtrop S Am
Chile
trop Am esp. Brasil
Temp SA, Chile-Arg
N & S Am, B. incana S US
Am from Canada to Argentina
W-ANT
W-ANT
ANT-P
W-ANT
ANT-P
W-NT
S-AMZ
W-ANT
S-AMZ
SPT
W-NT
W-NT
S-AMZ
AND
AND
W-ANT
ANT-P
S-AMZ
W-NT
W-NT
7
4
4
4
1
7
4
4
5
4
7
3
3
4
3
6
7
1
2
3
3
1
3
1
3
1
1
3
4
4
6
3
5
3
3
7
2
4
1
1
55
3
20
1
1
2
1
11
3
1
1
5
1
1
50
11
24
13
2
1
1
3
1
11
2
1
1
1
6
1
-22,00
-18,33
-31,50
-23,50
-40,50
-21,00
-18,00
-30,00
-35,83
-27,37
-32,88
-31,00
-22,00
-29,88
-34,40
-32,50
-25,00
-29,00
-35,33
-18,17
-18,00
-18,33
-17,58
-19,47
-27,33
-37,33
-27,28
-18,67
-18,20
-31,67
-28,57
-30,67
-30,83
-18,17
-18,37
-37,80
-55,63
-39,27
-40,00
-56,00
-56,00
-37,83
-56,00
-52,25
-43,17
-40,00
-36,67
-40,00
-55,97
-37,83
-35,50
-56,00
-30,50
-49,42
-55,00
-54,50
-55,00
-38,40
-30,67
-30,60
-39,67
-42,10
-19,17
-55,00
-40,00
-32,88
-43,62
-30,95
-52,00
-24,00
-29,90
-36,98
-35,38
-31,75
-48,25
-38,50
-27,92
-43,00
-44,04
-35,27
-36,44
-33,83
-31,00
-42,93
-36,12
-34,00
-40,50
-29,75
-42,38
-36,58
-36,25
-36,67
-27,99
-25,07
-28,97
-38,50
-34,69
-18,92
-36,60
-35,83
-30,73
-37,14
-30,89
-35,08
-21,18
15,80
37,30
7,77
16,50
15,50
35,00
19,83
26,00
16,42
15,80
7,12
5,67
18,00
26,08
3,43
3,00
31,00
1,50
14,08
36,83
36,50
36,67
20,82
11,20
3,27
2,33
14,82
0,50
36,80
8,33
4,32
12,95
0,12
33,83
5,63
226
Asterids
Monocots
Filicopsida
Eudicots
Asterids
Asterids
Asterids
Asterids
Rosids
Monocots
Monocots
Eudicots
Eudicots
Asterids
Monocots
Asterids
Asterids
Rosids
Eudicots
Eudicots
Asterids
Rosids
Rosids
Rosids
Monocots
Rosids
Rosids
Asterids
Chaptalia
Chascolytrum
Cheilanthes
Chenopodium
Chersodoma
Chevreulia
Chiliophyllum
Chiliotrichum
Chiropetalum
Chloraea
Chloris
Chorizanthe
Chrysosplenium
Chuquiraga
Chusquea
Cicendia
Ciclospermum
Cissarobryon
Cissus
Cistanthe
Citronella
Clarkia
Cleome
Cliococca
Codonorchis
Colletia
Colliguaja
Collomia
Ericales
Malpighiales
Rosales
Asparagales
Malpighiales
Brassicales
Myrtales
Unplaced
Caryophyllales
Vitales
Geraniales
Apiales
Gentianales
Poales
Asterales
Saxifragales
Caryophyllales
Poales
Asparagales
Malpighiales
Asterales
Asterales
Asterales
Asterales
Caryophyllales
Pteridales
Poales
Asterales
Asterales
Asterids
Chaetanthera
Caryophyllales
Solanales
Eudicots
Cerastium
Asterales
Asterids
Asterids
Centipeda
Apiales
Asterales
Cestrum
Asterids
Centella
Ceratophyllales
Asterids
Centaurodendron
Gentianales
Ceratophyllum
Asterids
Centaurium
Polemoniaceae
Euphorbiaceae
Rhamnaceae
Orchidaceae
Linaceae
Cleomaceae
Onagraceae
Icacinaceae
Portulacaceae
Vitaceae
Vivianiaceae
Apiaceae
Gentianaceae
Poaceae
Asteraceae
Saxifragaceae
Polygonaceae
Poaceae
Orchidaceae
Euphorbiaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Amaranthaceae
Pteridaceae
Poaceae
Asteraceae
Asteraceae
Solanaceae
Ceratophyllaceae
Caryophyllaceae
Asteraceae
Apiaceae
Asteraceae
Gentianaceae
15
5
5
3
1
275
42
20
49
200
1
3
2
110
23
64
50
55
50
20
7
3
5
9
125
150
6
55
44
175
20
100
6
40
2
60
W N Am, Bol to Patagonia
temp S Am, S Brasil
subtrop & temp S Am, Brazil, JF
S trop & temp S Am, Brazil
temp S Am
tropics & subtropics, spec. Am
N Am spec. California, 1 sp Chile/Arg
Malesia, E Aus, New Cale, Fiji, Samoa, C & trop S Am
N Am, Calif, Arizona, N México & S Am, Peru, Bol, Arg,
Chile
tropics incl Australia
Chile
Arg, Bol, Brasil, Chile, Paraguay, Uruguay
1 Euro & Medit, 1 Calif, W S Am
C & S Am, WI, JF
Andes S Am, Patagonia
N & temp S Am, Euro, N Afr, Asia,
arid & semiarid W Am, Chile
Cosmop
Andes Chile Perú, Arg, Brazil
Mexico 2, Peru, S Bras, Arg, Chile
S Andes Chile, Argentina
S Andes Chile, Argentina
S Brasil, Bol, Paraguay, Urug, Chile, N C Arg, Falkland
S Peru, Bol, Chile, Arg
cosmop, JF, ID
Cosmop, esp Neotrops & S Afr, Madag
Central & S Am, USA
trop & subtrop Am, S USA
trop Am, esp Brasil & Andes
cosmop
cosmop
Madag, Mascarene Is, Asia, Aus, NZ, New Cale, Tahiti,
Chile
Pantrop to Chile, NZ, JF
JF
Cosmop exc trop & S Afr, N hemisp 1 ext. to Aus, 1 to
Chile
ANT-P
S-AMZ
S-AMZ
S-AMZ
ANT-P
SPT
ANT-P
S-AMZ
W-ANT
W-NT
AND
W-ANT
ANT-P
S-AMZ
ANT-P
S-AMZ
AND
ANT-P
W-NT
AND
W-NT
4
3
3
3
5
1
4
2
4
1
6
3
4
3
3
4
4
7
3
4
5
5
3
3
7
7
4
3
3
3
7
7
1
1
6
7
2
4
2
1
1
1
1
1
18
1
1
1
1
13
5
2
24
1
29
4
2
2
3
3
12
9
1
1
37
1
1
2
1
1
2
1
-32,90
-23,50
-30,25
-35,00
-34,88
-20,22
-29,67
-30,67
-18,58
-30,00
-33,00
-20,82
-25,00
-30,67
-17,72
-52,00
-18,00
-18,47
-30,50
-22,50
-36,00
-52,62
-33,03
-17,72
-18,00
-18,00
-32,13
-33,03
-17,58
-18,75
-33,42
-23,00
-30,00
-35,12
-25,00
-53,00
-46,67
-46,50
-56,00
-40,32
-26,33
-40,33
-38,75
-37,93
-43,00
-37,80
-40,00
-38,00
-46,67
-35,75
-56,00
-37,22
-18,47
-55,33
-38,00
-56,00
-53,25
-43,00
-23,58
-54,00
-46,55
-43,83
-36,67
-40,25
-40,33
-36,82
-33,33
-41,83
-43,00
-41,50
-42,95
-35,08
-38,38
-45,50
-37,60
-23,28
-35,00
-34,71
-28,26
-36,50
-35,40
-30,41
-31,50
-38,67
-26,73
-54,00
-27,61
-18,47
-42,92
-30,25
-46,00
-52,93
-38,02
-20,65
-36,00
-32,28
-37,98
-34,85
-28,92
-29,54
-35,12
-28,17
-35,92
-39,06
-33,25
20,10
23,17
16,25
21,00
5,43
6,12
10,67
8,08
19,35
13,00
4,80
19,18
13,00
16,00
18,03
4,00
19,22
0,00
24,83
15,50
20,00
0,63
9,97
5,87
36,00
28,55
11,70
3,63
22,67
21,58
3,40
10,33
11,83
7,88
0,00
16,50
227
Eudicots
Asterids
Monocots
Asterids
Asterids
Eudicots
Asterids
Asterids
Asterids
Rosids
Eudicots
Eudicots
Monocots
Rosids
Asterids
Eudicots
Rosids
Asterids
Rosids
Rosids
Rosids
Asterids
Asterids
Magnoliids
Filicopsida
Asterids
Asterids
Eudicots
Asterids
Monocots
Asterids
Monocots
Asterids
Filicopsida
Rosids
Colobanthus
Combera
Conanthera
Convolvulus
Conyza
Copiapoa
Coprosma
Cordia
Coreopsis
Coriaria
Corrigiola
Corryocactus
Cortaderia
Corynabutilon
Cotula
Crassula
Cremolobus
Cressa
Crinodendron
Cristaria
Croton
Cruckshanksia
Cryptantha
Cryptocarya
Cryptogramma
Cuatrecasasiella
Cuminia
Cumulopuntia
Cuscuta
Cynodon
Cynoglossum
Cyperus
Cyphocarpus
Cystopteris
Dalea
Fabales
Dryopteridales
Asterales
Poales
Unplaced
Poales
Solanales
Caryophyllales
Lamiales
Asterales
Pteridales
Laurales
Unplaced
Gentianales
Malpighiales
Malvales
Oxalidales
Solanales
Brassicales
Saxifragales
Asterales
Malvales
Poales
Caryophyllales
Caryophyllales
Cucurbitales
Asterales
Unplaced
Gentianales
Caryophyllales
Asterales
Solanales
Asparagales
Solanales
Caryophyllales
Fabaceae
Dryopteridaceae
Campanulaceae
Cyperaceae
Boraginaceae
Poaceae
Convolvulaceae
Cactaceae
Lamiaceae
Asteraceae
Pteridaceae
Lauraceae
Boraginaceae
Rubiaceae
Euphorbiaceae
Malvaceae
Elaeocarpaceae
Convolvulaceae
Brassicaceae
Crassulaceae
Asteraceae
Malvaceae
Poaceae
Cactaceae
Caryophyllaceae
Coriariaceae
Asteraceae
Boraginaceae
Rubiaceae
Cactaceae
Asteraceae
Convolvulaceae
Tecophilaeaceae
Solanaceae
Caryophyllaceae
165
12
3
400
75
9
155
20
1
2
2
230
100
7
1250
75
5
5
7
200
55
7
25
12
13
15
114
320
90
25
60
150
5
2
20
Am, Canada to Arg
temp & warm regions, montane tropics
Chile
trop, subtrop, temp regions, JF
temp & subtrop reg
cosmop
Bol, S Peru, N Chile
JF
Ecuador, Peru, Chile, Arg
temperate N & S
tropics & subtropics, Madag, S Afr, S India, Ceylon,
Malaysia, Philippines, N Australia
W US, W temp S Am
Chile, Arg
Chile, Arg, Bol, Perú, ID
Bol, N Arg, C Chile, Brasil
trop & subtrop
Andes Colombia to Chile
cosmop, Euro, A & S Afr, Madag, Aus, Am
S Afr, NZ, New Guinea, Aus, S Am
Chile, Arg
S Am, 4 NZ, 1 New Guinea
Bol, S Peru, N Chile
S Am Andes Colom to Chile, Euro, Medit, Afr, Madag
Mex to Chile, W Medit, Himal to Japan, New Guinea,
NZ. C ruscifolia disj NZ
N & S Am
trop & subtropics
E Malesia, Aus, NZ, Pacific, JF, 12 sp Hawaii
N Chile
cosmop esp. temp, trop & subtrops
Chile
Andes Chile Arg
S Pacific, Aus, NZ, temp S Am, Ecuador
W-NT
W-ANT
AND
AND
W-ANT
ANT-P
AND
AND
S-AMZ
AND
AUS
AND
W-NT
SPT
SPT
3
4
6
7
7
1
7
3
6
3
4
1
4
3
1
3
3
1
3
7
1
5
2
3
7
7
3
1
2
6
1
7
6
5
2
3
1
3
26
1
5
10
2
1
1
1
1
39
7
1
19
2
1
1
7
1
7
6
1
3
1
1
1
2
25
27
5
5
2
3
-18,00
-18,00
-27,18
-20,00
-32,95
-18,47
-19,25
-18,00
-17,72
-31,30
-30,33
-18,83
-22,08
-24,90
-18,17
-32,23
-20,45
-17,58
-18,00
-20,25
-30,67
-18,47
-19,75
-30,00
-35,02
-18,17
-27,82
-22,00
-17,58
-25,00
-22,08
-37,00
-17,58
-25,40
-55,00
-32,35
-43,00
-39,98
-33,43
-40,35
-33,00
-24,50
-37,40
-40,25
-44,50
-33,83
-25,33
-35,63
-42,77
-35,33
-18,50
-56,00
-22,00
-42,53
-56,00
-20,00
-37,00
-43,83
-18,17
-31,83
-31,00
-55,00
-40,00
-37,95
-44,00
-56,00
-21,70
-36,50
-29,77
-31,50
-36,47
-25,95
-29,80
-25,50
-21,11
-34,35
-35,29
-31,67
-27,96
-25,12
-26,90
-37,50
-27,89
-18,04
-37,00
-21,13
-36,60
-37,23
-19,88
-33,50
-39,43
-18,17
-29,83
-26,50
-36,29
-32,50
-30,02
-40,50
-36,79
7,40
37,00
5,17
23,00
7,03
14,97
21,10
15,00
0,00
6,78
6,10
9,92
25,67
11,75
0,43
17,47
10,53
14,88
0,92
38,00
1,75
11,87
37,53
0,25
7,00
8,82
0,00
4,02
0,00
9,00
37,42
15,00
15,87
7,00
38,42
228
Monocots
Monocots
Monocots
Rosids
Asterids
Asterids
Asterids
Filicopsida
Asterids
Rosids
Rosids
Dissanthelium
Distichia
Distichlis
Dodonaea
Domeykoa
Donatia
Doniophyton
Doodia
Downingia
Draba
Drapetes
Rosids
Dinemagonum
Rosids
Monocots
Dielsiochloa
Asterids
Asterids
Dicliptera
Discaria
Filicopsida
Dicksonia
Diposis
Asterids
Dichondra
Asterids
Monocots
Dichanthelium
Diplostephium
Monocots
Deuterocohnia
Asterids
Eudicots
Desmaria
Filicopsida
Asterids
Desfontainia
Diplolepis
Rosids
Descurainia
Diplazium
Monocots
Deschampsia
Asterids
Filicopsida
Dennstaedtia
Diostea
Asterids
Dendroseris
Monocots
Filicopsida
Davallia
Dioscorea
Asterids
Daucus
Rosids
Asterids
Dasyphyllum
Dinemandra
Monocots
Danthonia
Malvales
Brassicales
Asterales
Blechnales
Asterales
Asterales
Apiales
Sapindales
Poales
Poales
Poales
Rosales
Apiales
Asterales
Gentianales
Dryopteridales
Lamiales
Dioscoreales
Malpighiales
Malpighiales
Poales
Lamiales
Dennstaedtiales
Solanales
Poales
Poales
Santalales
Unplaced
Brassicales
Poales
Dennstaedtiales
Asterales
Davalliales
Apiales
Asterales
Poales
Thymelaeaceae
Brassicaceae
Campanulaceae
Blechnaceae
Asteraceae
Stylidiaceae
Apiaceae
Sapindaceae
Poaceae
Juncaceae
Poaceae
Rhamnaceae
Apiaceae
Asteraceae
Apocynaceae
Dryopteridaceae
Verbenaceae
Dioscoreaceae
Malpighiaceae
Malpighiaceae
Poaceae
Acanthaceae
Dicksoniaceae
Convolvulaceae
Poaceae
Bromeliaceae
Loranthaceae
Desfontainiaceae
Brassicaceae
Poaceae
Dennstaedtiaceae
Asteraceae
Davalliaceae
Apiaceae
Asteraceae
Poaceae
1
350
11
12
2
2
4
68
9
3
16
8
3
90
4
300
4
400
1
1
1
200
20
7
58
13
1
1
40
35
45
11
90
22
40
20
temp S Am
Arctic & temp n Hemis, Asia, montane C & S Am
W N Am, 1 Chile
Australia, NZ, Pacific islands, Hawaii, Easter Island, N
Guinea
Andes Chile Arg
Tasman, NZ, subantarctic S Am
Peru Chile
tropics & subtrop, espec Australia
Am from Canada to Argentina, 1 sp Australia
Andes S Am
Peru, Chile, Bol to California
Aus, NZ, S Am
temp S Am
trop Andes from Colombia to Bol & N Chile, 1 sp Costa
Rica
Chile, Arg
cosmop
temperate S Am, Arg, Chile
trop, subtrop, warm-temp regions
Chile
Chile
Peru, Bol, Arg, Chile
trop Am, Malesia, New Guinea, Aus, New Caled, NZ
tropics & subtropics, JF
Toda America
Peru, Bol, SW Bra, N Chile, N Arg
Chile
Costa Rica to Cape Horn
S Am, N Am, Canary Is.
temp & cool, espec N Euro, montane tropics
trop to warm temp, Madag
JF
W Medit, Himal, Japan, Aus, Tahiti, Afr, Madag, IP
Euro, Medit, SW & S Asia, trop Afr. Aus, NZ, & Am
Chile, Andes, SE Brazil
N & S Am, Euro, Caucasus
W-ANT
ANT-P
SPT
AND
AUS
AND
C-PAC
AND
ANT-P
AUS
AND
AND
SPT
W-NT
S-AMZ
AND
W-ANT
W-ANT
S-AMZ
W-ANT
5
4
4
2
3
2
3
1
4
3
4
2
5
3
5
7
5
1
6
6
3
1
2
1
3
3
6
3
4
4
1
6
1
7
3
4
1
7
1
1
2
1
2
1
3
2
1
4
1
3
4
1
1
41
1
1
1
1
2
1
1
1
1
1
14
16
1
11
1
2
2
8
-46,33
-18,00
-31,50
-21,50
-37,80
-22,00
-31,00
-18,47
-18,00
-17,58
-28,72
-31,97
-17,58
-33,35
-31,00
-23,50
-21,50
-27,50
-18,00
-24,50
-25,00
-36,00
-25,00
-35,60
-35,33
-18,20
-28,80
-32,45
-25,38
-32,88
-32,00
-56,00
-55,00
-38,00
-30,67
-56,00
-25,43
-31,17
-46,50
-21,00
-18,20
-55,00
-34,47
-23,00
-47,00
-43,83
-42,00
-27,42
-31,50
-18,20
-25,50
-40,83
-38,00
-26,17
-40,18
-53,00
-53,15
-56,00
-35,85
-46,30
-42,47
-42,62
-51,17
-36,50
-34,75
-26,08
-46,90
-23,72
-31,08
-32,48
-19,50
-17,89
-41,86
-33,22
-20,29
-40,18
-37,42
-32,75
-24,46
-29,50
-18,10
-25,00
-32,92
-37,00
-25,58
-37,89
-44,17
-35,68
-42,40
-34,15
-35,84
-37,68
-37,31
9,67
37,00
6,50
0,00
9,17
18,20
3,43
0,17
28,03
3,00
0,62
26,28
2,50
5,42
13,65
0,00
12,83
18,50
5,92
4,00
0,20
1,00
0,00
15,83
0,00
1,17
4,58
17,67
34,95
27,20
3,40
0,00
0,00
20,92
9,58
10,62
229
Asterids
Asterids
Rosids
Asterids
Asterids
Eudicots
Rosids
Rosids
Asterids
Asterids
Rosids
Rosids
Eremocharis
Eremodraba
Eriachaenium
Erigeron
Eriosyce
Erodium
Errazurizia
Eryngium
Escallonia
Eucryphia
Eudema
Asterids
Encelia
Erechtites
Asterids
Empetrum
Eudicots
Eudicots
Embothrium
Monocots
Asterids
Elytropus
Ercilla
Monocots
Elymus
Eragrostis
Monocots
Elodea
Sphenopsida
Monocots
Eleocharis
Equisetum
Rosids
Elatine
Monocots
Filicopsida
Elaphoglossum
Epipetrum
Eudicots
Echinopsis
Rosids
Asterids
Eccremocarpus
Epilobium
Rosids
Dysopsis
Gnetopsida
Asterids
Dunalia
Monocots
Filicopsida
Dryopteris
Ephedra
Eudicots
Drymaria
Enneapogon
Magnoliids
Eudicots
Drimys
Drosera
Brassicales
Oxalidales
Unplaced
Apiales
Fabales
Geraniales
Caryophyllales
Asterales
Asterales
Brassicales
Apiales
Asterales
Caryophyllales
Poales
Equisetales
Dioscoreales
Myrtales
Ephedrales
Poales
Asterales
Ericales
Proteales
Gentianales
Poales
Alismatales
Poales
Malpighiales
Blechnales
Caryophyllales
Lamiales
Malpighiales
Solanales
Dryopteridales
Caryophyllales
Caryophyllales
Canellales
Brassicaceae
Cunoniaceae
Escalloniaceae
Apiaceae
Fabaceae
Geraniaceae
Cactaceae
Asteraceae
Asteraceae
Brassicaceae
Apiaceae
Asteraceae
Phytolaccaceae
Poaceae
Equisetaceae
Dioscoreaceae
Onagraceae
Ephedraceae
Poaceae
Asteraceae
Ericaceae
Proteaceae
Apocynaceae
Poaceae
Hydrocharitaceae
Cyperaceae
Elatinaceae
Lomariopsidaceae
Cactaceae
Bignoniaceae
Euphorbiaceae
Solanaceae
Dryopteridaceae
Caryophyllaceae
Droseraceae
Winteraceae
6
6
40
240
4
75
35
200
1
2
9
10
2
300
30
3
164
60
35
15
3
1
1
150
9
200
12
400
100
5
1
5
150
50
115
11
Mexico, C & S Am, JF
Andes Ecuador to Arg
SE Aus, Tasm, Chile
Andes S Am, SE Brasil, JF
trop & temp, Eurasia, Am, Afr, Malesia, Aus, JF
SW US, Mex, 1 sp Chile
Cosmop
Chile, Arg, S Peru
Eurasia, N & S Am, Aus, JF
Tierra del Fuego
Andes Chile Peru
Andes Chile Peru
N & S Am
Chile
temp & trop, ID
almost cosmop, Madag
Chile
temp Euro, N Am, arctic, montane trop
Eurasia, AM
temp Eurasia, Am, trop & subtrop
SW USA, Mexico, Chile, Peru, Galapagos
N temp & Arctic, Europa, S Andes, Falkland, JF
Chile y Argentina, 1 spp.
Chile, Arg
temp & subtrop, montane tropics
AM temp
cosmop, JF
temp, trop & subtrop regions
pantrops & subtrop, Madag, IP
S Am
Peru, Chile, Arg
Andes Costa Rica, Ecuador, Chile, Arg, JF
Andes Colombia to Argentina
cosmop
pantrop
cosmop, espec Aus, Tasm & NZ
AND
AUS
AND
ANT-P
AND
W-ANT
AND
AND
W-NT
W-ANT
W-ANT
ANT-P
W-ANT
W-NT
W-ANT
AND
AND
AND
AND
W-NT
3
2
3
7
4
7
3
4
5
3
3
3
6
7
7
6
4
4
7
4
4
5
5
7
3
7
4
1
3
3
3
3
7
1
7
3
2
2
13
10
1
1
28
12
1
1
1
1
2
6
2
3
12
7
1
1
1
1
1
6
1
19
1
3
9
1
1
1
1
3
1
1
-18,23
-36,08
-18,50
-25,20
-25,00
-32,75
-18,42
-29,00
-48,83
-19,80
-21,50
-23,53
-33,50
-18,17
-18,40
-25,00
-18,00
-18,17
-19,87
-19,28
-32,83
-35,25
-33,45
-22,00
-19,25
-17,67
-34,80
-39,83
-18,50
-32,93
-30,67
-18,20
-22,00
-37,83
-30,33
-22,95
-45,50
-53,00
-40,60
-30,50
-33,38
-36,72
-56,00
-53,50
-19,80
-27,07
-25,92
-44,58
-38,00
-44,35
-37,67
-55,50
-53,00
-23,00
-30,67
-56,00
-55,67
-43,83
-53,00
-36,50
-55,00
-39,63
-43,83
-35,00
-40,25
-55,00
-18,83
-25,40
-56,00
-55,97
-20,59
-40,79
-35,75
-32,90
-27,75
-33,07
-27,57
-42,50
-51,17
-19,80
-24,28
-24,73
-39,04
-28,08
-31,38
-31,33
-36,75
-35,58
-21,43
-24,98
-44,42
-45,46
-38,64
-37,50
-27,88
-36,33
-37,22
-41,83
-26,75
-36,59
-42,83
-18,52
-23,70
-46,92
-43,15
4,72
9,42
34,50
15,40
5,50
0,63
18,30
27,00
4,67
0,00
5,57
2,38
11,08
19,83
25,95
12,67
37,50
34,83
3,13
11,38
23,17
20,42
10,38
31,00
17,25
37,33
4,83
4,00
16,50
7,32
24,33
0,63
0,00
3,40
18,17
25,63
230
Eudicots
Rosids
Asterids
Asterids
Asterids
Asterids
Asterids
Rosids
Monocots
Monocots
Pinopsida
Asterids
Asterids
Asterids
Rosids
Rosids
Eudicots
Rosids
Rosids
Monocots
Asterids
Asterids
Asterids
Asterids
Asterids
Asterids
Monocots
Rosids
Asterids
Asterids
Rosids
Rosids
Monocots
Rosids
Eudicots
Asterids
Eulychnia
Euphorbia
Euphrasia
Evolvulus
Exodeconus
Fabiana
Facelis
Fagonia
Fascicularia
Festuca
Fitzroya
Flaveria
Flourensia
Fonkia
Fragaria
Francoa
Frankenia
Fuchsia
Fuertesimalva
Gaimardia
Galinsoga
Galium
Gamocarpha
Gamochaeta
Gamochaetopsis
Gaultheria
Gavilea
Gayophytum
Gentiana
Gentianella
Geoffroea
Geranium
Gethyum
Geum
Gevuina
Gilia
Ericales
Proteales
Rosales
Asparagales
Geraniales
Fabales
Gentianales
Gentianales
Myrtales
Asparagales
Ericales
Asterales
Asterales
Asterales
Gentianales
Asterales
Poales
Malvales
Myrtales
Caryophyllales
Geraniales
Rosales
Lamiales
Asterales
Asterales
Pinales
Poales
Poales
Zygophyllales
Asterales
Solanales
Solanales
Solanales
Lamiales
Malpighiales
Caryophyllales
Polemoniaceae
Proteaceae
Rosaceae
Alliaceae
Geraniaceae
Fabaceae
Gentianaceae
Gentianaceae
Onagraceae
Orchidaceae
Ericaceae
Asteraceae
Asteraceae
Calyceraceae
Rubiaceae
Asteraceae
Centrolepidaceae
Malvaceae
Onagraceae
Frankeniaceae
Francoaceae
Rosaceae
Gratiolaceae
Asteraceae
Asteraceae
Cupressaceae
Poaceae
Bromeliaceae
Zygophyllaceae
Asteraceae
Solanaceae
Solanaceae
Convolvulaceae
Orobanchaceae
Euphorbiaceae
Cactaceae
25
3
50
2
400
3
256
360
9
15
140
1
40
6
350
15
3
14
105
50
1
15
1
32
21
1
500
1
30
3
15
6
100
300
2000
7
Peru, Chile
W N Am & S Am
New Guinea, Aus, Chile, Arg
Arctic & temp regions
Chile
cosmop, espec tem regs & tropic montane
S Am from Colombia & Venez to Arg, Brazil
Eurasia, NW Afr, N, C & S Am, Aus, NZ
Euro, NW Afr, Asia, E Aus, Am
temp W N Am & W S Am
Temp S Am, JF
E & SE Asia, Malesia, Aus, NZ, N, C, S Am, WI, JF
S Arg, Chile
N & S Am, Brazil, JF
temp S Am
cosmop, JF
temp & subtrop C & S Am
New Guinea, NZ, Tasmania, TFuego, Falkland
Mexico, C Andes, Galapagos, ID
C & S Am, NZ, Tahiti
temp Euro, Afr, Aus, Am, ID
Chile
N & S Am, temp Euras
Chile, Arg
SW USA, Mexico, Peru, Chile, Arg
W N Am, Mexico, S Am, 1 sp Aus
Chile Arg
cosmop
Chile
Medit, Euro, SW Asia, NE Afr, SW N Am, Peru, Chile
S Brasil, Peru, Bol, Paraguay, Urug, Chile, Arg
S Am, Peru, Ecuador, Galapagos, N Chile
trops & subtrops
temp N Hemishp, Euro, Middle asia, Australia, NZ, temp
S Am, JF
subcosmop
ANT-P
SPT
AND
W-ANT
ANT-P
C-PAC
W-NT
W-NT
SPT
W-NT
AUS
W-ANT
ANT-P
C-PAC
W-ANT
S-AMZ
AND
AND
W-ANT
AND
4
2
7
6
7
3
7
4
4
5
4
5
3
5
7
3
2
3
2
7
6
4
5
4
4
5
7
6
4
3
3
3
1
4
7
3
4
1
4
2
5
1
3
6
2
12
11
1
22
6
17
2
1
3
2
4
1
1
1
1
1
1
24
1
2
2
7
2
1
15
19
6
-18,00
-35,40
-33,95
-29,50
-17,67
-18,47
-21,00
-18,00
-28,50
-30,67
-30,67
-37,48
-17,58
-36,83
-18,00
-17,58
-39,88
-18,30
-28,50
-20,63
-32,23
-35,00
-40,75
-29,45
-19,00
-40,17
-17,67
-33,58
-18,83
-25,40
-18,20
-18,38
-35,33
-30,00
-18,47
-18,50
-51,00
-45,42
-55,00
-34,25
-55,50
-31,50
-56,00
-55,00
-37,00
-55,33
-56,00
-37,48
-55,25
-46,00
-56,00
-42,62
-56,00
-30,90
-55,00
-53,00
-43,50
-46,43
-40,75
-33,83
-36,60
-42,58
-56,00
-44,00
-31,67
-39,82
-45,77
-24,00
-37,83
-55,00
-52,67
-32,25
-34,50
-40,41
-44,48
-31,88
-36,58
-24,98
-38,50
-36,50
-32,75
-43,00
-43,33
-37,48
-36,42
-41,42
-37,00
-30,10
-47,94
-24,60
-41,75
-36,82
-37,87
-40,72
-40,75
-31,64
-27,80
-41,38
-36,83
-38,79
-25,25
-32,61
-31,98
-21,19
-36,58
-42,50
-35,57
-25,38
33,00
10,02
21,05
4,75
37,83
13,03
35,00
37,00
8,50
24,67
25,33
0,00
37,67
9,17
38,00
25,03
16,12
12,60
26,50
32,37
11,27
11,43
0,00
4,38
17,60
2,42
38,33
10,42
12,83
14,42
27,57
5,62
2,50
25,00
34,20
13,75
231
Monocots
Eudicots
Eudicots
Asterids
Rosids
Asterids
Asterids
Asterids
Monocots
Asterids
Asterids
Habenaria
Haloragis
Hamadryas
Haplopappus
Haplorhus
Hebe
Hedyotis
Helenium
Helictotrichon
Heliotropium
Helogyne
Asterids
Griselinia
Eudicots
Asterids
Grindelia
Haageocereus
Monocots
Greigia
Asterids
Asterids
Gratiola
Asterids
Rosids
Grammosperma
Gypothamnium
Filicopsida
Grammitis
Gymnophyton
Asterids
Grabowskia
Monocots
Eudicots
Gomphrena
Gymnachne
Magnoliids
Gomortega
Asterids
Asterids
Gochnatia
Asterids
Asterids
Gnaphalium
Guynesomia
Rosids
Glycyrrhiza
Gutierrezia
Monocots
Glyceria
Eudicots
Eudicots
Glinus
Gunnera
Filicopsida
Gleichenia
Rosids
Asterids
Glandularia
Guindilia
Monocots
Gilliesia
Asterales
Unplaced
Poales
Asterales
Gentianales
Lamiales
Sapindales
Asterales
Ranunculales
Saxifragales
Asparagales
Caryophyllales
Asterales
Apiales
Poales
Asterales
Asterales
Gunnerales
Sapindales
Apiales
Asterales
Poales
Lamiales
Brassicales
Polypodiales
Solanales
Caryophyllales
Laurales
Asterales
Asterales
Fabales
Poales
Caryophyllales
Gleicheniales
Lamiales
Asparagales
Asteraceae
Boraginaceae
Poaceae
Asteraceae
Rubiaceae
Plantaginaceae
Anacardiaceae
Asteraceae
Ranunculaceae
Haloragaceae
Orchidaceae
Cactaceae
Asteraceae
Apiaceae
Poaceae
Asteraceae
Asteraceae
Gunneraceae
Sapindaceae
Griseliniaceae
Asteraceae
Bromeliaceae
Gratiolaceae
Brassicaceae
Grammitidaceae
Solanaceae
Amaranthaceae
Gomortegaceae
Asteraceae
Asteraceae
Fabaceae
Poaceae
Molluginaceae
Gleicheniaceae
Verbenaceae
Alliaceae
8
260
50
32
150
80
1
75
5
27
600
20
1
6
1
1
27
40
3
6
60
27
25
1
400
4
100
1
66
100
20
45
9
110
100
3
Peru, Chile, Bol, Arg
temp, subtrop, trop regs, arid
temp N Hemis, montane trops, S Afr
W N Am, Mexico, C America, Andes
tropics espec Asia & Malesia, JF
New Guinea, Aus, Tasm, NZ, temp S Am, Falkland
Chile, Peru
Am espec Chile
Antarctic S Am
Aus, N. Caled, NZ, Rapa, JF
pantrop & subtrop
Peru, Chile
N Chile
Andes Chile Arg
Chile
Chile
W N Am & South S Am
Am from Mexic to Chile, trop & S Afr, Madag, Malesia,
Tasm, NZ, Hawaii, JF
Chile Arg
NZ, Chile, Paraguay, S Brasil
W N Am & S Am, Brazil
C & S Am, JF
Patagonia
New Guinea, Asia, Am, Aus, Madag, warm & temp, JF
Mexico, Galapagos, temp S Am
trop & subtrop Am, Afr, Indochina, Aus
Chile
Trop & Subtrop Am, S USA, SE Asia
cosmop
Eurasia, Aus, N Am & temp S Am
pantrop
pantrop, Madag, JF
disjunct N Am S Am
Chile, Arg
AND
W-NT
SPT
AND
W-NT
AUS
AND
AND
ANT-P
AND
AUS
W-NT
W-NT
W-NT
C-PAC
W-ANT
ANT-P
3
7
7
3
1
2
3
3
5
2
1
3
6
3
6
6
4
1
3
2
3
3
7
5
1
3
1
6
4
7
4
7
1
1
4
5
2
24
1
7
1
2
1
64
4
2
1
2
1
5
1
1
6
7
1
4
4
2
1
1
3
1
2
1
1
30
1
1
1
4
6
2
-17,58
-18,20
-36,83
-22,00
-34,77
-46,67
-18,75
-21,18
-50,62
-28,50
-18,33
-25,00
-23,50
-32,88
-29,75
-21,00
-29,83
-31,00
-24,93
-18,42
-35,00
-34,77
-51,00
-37,42
-25,40
-18,00
-35,62
-30,67
-17,58
-34,08
-33,25
-31,50
-37,23
-21,27
-32,37
-23,87
-38,63
-36,83
-40,33
-42,50
-56,00
-19,25
-45,50
-54,92
-52,00
-20,00
-26,13
-34,55
-40,00
-31,50
-52,50
-56,00
-35,97
-49,20
-38,78
-43,50
-42,50
-51,00
-55,87
-25,40
-22,50
-37,67
-37,80
-54,50
-37,58
-41,67
-37,17
-55,83
-38,00
-38,25
-20,73
-28,42
-36,83
-31,17
-38,63
-51,33
-19,00
-33,34
-52,77
-40,25
-19,17
-25,57
-29,03
-36,44
-30,63
-36,75
-42,92
-33,48
-37,07
-28,60
-39,25
-38,63
-51,00
-46,64
-25,40
-20,25
-36,64
-34,23
-36,04
-35,83
-37,46
-34,33
-46,53
-29,63
-35,31
6,28
20,43
0,00
18,33
7,73
9,33
0,50
24,32
4,30
0,00
23,50
1,67
1,13
11,05
7,12
1,75
31,50
26,17
4,97
24,27
20,37
8,50
7,73
0,00
18,45
0,00
4,50
2,05
7,13
36,92
3,50
8,42
5,67
18,60
16,73
5,88
232
Asterids
Asterids
Lycopsida
Rosids
Asterids
Monocots
Asterids
Monocots
Monocots
Monocots
Isoetes
Ivania
Jaborosa
Jarava
Jovellana
Juania
Jubaea
Juncus
Asterids
Hydrocotyle
Ipomopsis
Asterids
Hydrangea
Ipomoea
Rosids
Hybanthus
Monocots
Lycopsida
Huperzia
Imperata
Asterids
Huidobria
Asterids
Asterids
Huanaca
Filicopsida
Monocots
Hordeum
Hypsela
Asterids
Homalocarpus
Hypolepis
Rosids
Hollermayera
Asterids
Rosids
Hoffmannseggia
Rosids
Filicopsida
Histiopteris
Hypochaeris
Asterids
Hippuris
Hypericum
Asterids
Hieracium
Filicopsida
Monocots
Heterozostera
Hymenophyllum
Asterids
Heterosperma
Filicopsida
Monocots
Herreria
Hymenoglossum
Monocots
Herbertia
Poales
Arecales
Arecales
Lamiales
Poales
Solanales
Brassicales
Isoetales
Ericales
Solanales
Poales
Asterales
Dennstaedtiales
Asterales
Malpighiales
Hymenophyllales
Hymenophyllales
Apiales
Cornales
Malpighiales
Lycopodiales
Cornales
Apiales
Poales
Apiales
Brassicales
Fabales
Dennstaedtiales
Lamiales
Asterales
Alismatales
Asterales
Asparagales
Asparagales
Juncaceae
Arecaceae
Arecaceae
Calceolariaceae
Poaceae
Solanaceae
Brassicaceae
Isoetaceae
Polemoniaceae
Convolvulaceae
Poaceae
Campanulaceae
Dennstaedtiaceae
Asteraceae
Hypericaceae
Hymenophyllaceae
Hymenophyllaceae
Apiaceae
Hydrangeaceae
Violaceae
Lycopodiaceae
Loasaceae
Apiaceae
Poaceae
Apiaceae
Brassicaceae
Fabaceae
Dennstaedtiaceae
Plantaginaceae
Asteraceae
Zosteraceae
Asteraceae
Agavaceae
Iridaceae
275
1
1
5
59
23
1
150
26
650
9
5
40
55
370
250
1
130
29
130
350
2
4
30
6
1
28
5
1
200
1
5
8
5
Cosmop, JF, IP
Chile
JF
NZ, Chile
Arg, Bol, Chile, Colombia, CosRica, Ecuador, El Salvador,
Guatemala, Mexico, Peru, USA, Uruguay, Venezuela
temp S Am. S Peru, Bol, Bras, Parag, Chile, Arg, Uru
N Chile
cosmop, Madag
W N Am & Florida, 1 Chile Arg
pantrop
trop, subtrop & temp regions
E Aus, NZ, Andes S Am
trop & subtrop, Madag
temp S Am, Eurasia Medit, N Afr
temp regs, montane tropics
trop & S temp, Euro, Japan, Madag, JF
Chile, JF
cosmop
Himalayas, Asia, Philippines, N, C Am, Andes to Chile
trops & subtrops
Subcosmop, Madag
Chile
Patagonia
temp & subtrop regs, montane tropics
Chile
Chile
25 SW US, Chile, Arg
pantrop, Madag
circumboreal, S Am, Australia
temp regions & montane tropics, excl Australasia
coastal temp Aus, Tasm, N Chile
SW USA, Mexico, C Am, Peru, Arg, Chile
Brasil, Uru, Parag, Arg, Chile
temp S Am, Chile, Brazil
AUS
W-NT
S-AMZ
ANT-P
AUS
W-ANT
W-ANT
W-ANT
ANT-P
W-ANT
W-ANT
AUS
W-NT
S-AMZ
S-AMZ
7
6
6
2
3
3
6
7
4
1
7
2
1
4
7
7
6
7
4
1
7
6
5
4
6
6
4
1
4
4
2
3
3
3
28
1
1
3
22
9
1
2
1
1
4
1
1
33
2
19
1
8
1
1
1
2
4
11
6
1
7
1
1
6
1
2
1
1
-17,58
-30,50
-35,75
-17,67
-18,18
-27,40
-37,80
-21,00
-30,60
-18,00
-30,67
-17,58
-34,77
-30,67
-37,60
-18,83
-33,50
-36,50
-53,67
-19,00
-36,60
-18,18
-25,20
-38,67
-18,00
-42,05
-37,00
-34,65
-30,20
-17,58
-35,33
-33,00
-56,00
-35,50
-40,00
-53,50
-53,83
-27,40
-53,53
-26,93
-40,33
-54,00
-49,15
-54,00
-42,50
-56,00
-49,15
-51,27
-46,67
-37,08
-54,67
-29,50
-52,95
-55,00
-33,75
-41,00
-33,45
-54,07
-55,00
-55,00
-30,30
-23,00
-37,00
-40,00
-36,79
-33,00
-37,88
-35,58
-36,01
-27,40
-45,67
-23,97
-35,47
-36,00
-39,91
-35,79
-38,63
-43,33
-43,38
-35,05
-40,08
-36,79
-54,17
-24,25
-44,78
-36,59
-29,48
-39,83
-25,73
-48,06
-46,00
-44,83
-30,25
-20,29
-36,17
-36,50
38,42
5,00
0,00
4,25
35,83
35,65
0,00
15,73
5,93
0,00
9,73
36,00
18,48
36,42
7,73
25,33
11,55
32,43
13,17
0,58
1,00
10,50
16,35
36,82
8,55
2,33
15,45
12,02
18,00
20,35
0,10
5,42
1,67
7,00
233
Asterids
Rosids
Monocots
Rosids
Asterids
Monocots
Magnoliids
Asterids
Asterids
Monocots
Monocots
Eudicots
Asterids
Rosids
Eudicots
Asterids
Rosids
Asterids
Magnoliids
Magnoliids
Asterids
Rosids
Asterids
Rosids
Monocots
Eudicots
Monocots
Asterids
Rosids
Eudicots
Asterids
Pinopsida
Asterids
Asterids
Junellia
Kageneckia
Koeleria
Krameria
Kurzamra
Kyllinga
Lactoris
Lagenophora
Lampaya
Landoltia
Lapageria
Lardizabala
Laretia
Larrea
Lastarriaea
Lasthenia
Lathyrus
Latua
Laurelia
Laureliopsis
Lebetanthus
Lecanophora
Legenere
Legrandia
Lemna
Lenzia
Leontochir
Lepechinia
Lepidium
Lepidoceras
Lepidophyllum
Lepidothamnus
Leptinella
Leptocarpha
Asterales
Asterales
Pinales
Asterales
Santalales
Brassicales
Lamiales
Liliales
Caryophyllales
Alismatales
Myrtales
Asterales
Malvales
Ericales
Laurales
Laurales
Solanales
Fabales
Asterales
Caryophyllales
Zygophyllales
Apiales
Ranunculales
Liliales
Alismatales
Lamiales
Asterales
Piperales
Poales
Lamiales
Zygophyllales
Poales
Rosales
Lamiales
Asteraceae
Asteraceae
Podocarpaceae
Asteraceae
Santalaceae
Brassicaceae
Lamiaceae
Alstroemeriaceae
Portulacaceae
Lemnaceae
Myrtaceae
Campanulaceae
Malvaceae
Ericaceae
Atherospermataceae
Atherospermataceae
Solanaceae
Fabaceae
Asteraceae
Polygonaceae
Zygophyllaceae
Apiaceae
Lardizabalaceae
Philesiaceae
Lemnaceae
Verbenaceae
Asteraceae
Lactoridaceae
Cyperaceae
Lamiaceae
Krameriaceae
Poaceae
Rosaceae
Verbenaceae
1
33
3
1
2
180
40
1
1
14
1
2
7
1
1
2
1
155
18
3
5
1
1
1
1
3
15
1
40
1
18
45
4
47
Chile
New Guinea, Aus, NZ, Subantarctic Is, S Am
2 NZ, 1 S Chile/Arg
Patagonia Chile-Arg.
S Peru, Chile
Cosmop, ID
California, Andes, Mexico to S Brasil, Arg, Chile
Chile
Chile Arg
Cosmop
Chile
1 Calif, 1 Chile
Arg, Chile
Patagonia
Chile Arg
NZ & Chile
Chile
temp N hemis, Medit, montane tropics, E Afr, S Am
SW US, Chile-Arg
California, Chile
SW N Am, S Am
Chile, Argentina
Chile and adjacent Argentina
Chile
cosmop
Bol, Chile, Arg
SE Asia, Malesia, Australia, Tasmania, NZ, Central and
South America, JF, Falkland, Tristan da Cunha.
JF
pantrops
Chile / Arg
SW US, Mexico, Ecuador, Chile, Arg, Brazil
temp & subtrops, montane trops
Chile, Peru, Bolivia, Arg
SPT
AUS
AND
W-NT
AND
ANT-P
AUS
W-ANT
ANT-P
ANT-P
ANT-P
AND
AND
SPT
AND
W-NT
AND
AND
6
2
2
5
3
7
3
6
3
7
6
4
5
5
5
2
6
4
4
4
4
3
5
6
7
3
2
6
1
3
3
7
3
3
1
1
1
1
1
22
3
1
1
3
1
1
1
1
1
1
1
11
1
1
3
1
1
1
1
1
3
1
1
1
2
2
2
31
-35,78
-40,00
-40,17
-52,33
-36,33
-18,00
-29,83
-27,90
-23,00
-18,00
-35,27
-40,00
-52,10
-43,42
-38,37
-34,73
-40,17
-29,47
-31,25
-27,83
-28,80
-30,17
-32,50
-30,67
-55,50
-18,17
-33,67
-28,72
-17,58
-31,50
-30,33
-18,00
-40,82
-56,00
-55,00
-53,00
-43,00
-55,00
-40,00
-28,08
-30,25
-46,50
-36,68
-40,00
-52,10
-56,00
-46,67
-41,17
-43,00
-53,00
-51,00
-35,97
-36,75
-35,50
-39,87
-40,83
-55,50
-23,50
-56,00
-30,50
-33,50
-54,83
-38,67
-52,50
-38,30
-48,00
-47,58
-52,67
-39,67
-36,50
-34,92
-27,99
-26,63
-32,25
-35,98
-40,00
-52,10
-49,71
-42,52
-37,95
-41,58
-41,23
-41,13
-31,90
-32,78
-32,83
-36,18
-35,75
-55,50
-20,83
-44,83
-29,61
-25,54
-43,17
-34,50
-35,25
5,03
16,00
14,83
0,67
6,67
37,00
10,17
0,18
7,25
28,50
1,42
0,00
0,00
12,58
8,30
6,43
2,83
23,53
19,75
8,13
7,95
5,33
7,37
10,17
0,00
5,33
22,33
0,00
0,00
1,78
15,92
23,33
8,33
34,50
234
Monocots
Monocots
Asterids
Eudicots
Asterids
Monocots
Asterids
Monocots
Eudicots
Monocots
Asterids
Eudicots
Asterids
Asterids
Asterids
Rosids
Asterids
Rosids
Asterids
Rosids
Asterids
Asterids
Eudicots
Asterids
Filicopsida
Rosids
Asterids
Asterids
Rosids
Rosids
Rosids
Monocots
Monocots
Asterids
Asterids
Asterids
Lycopsida
Leptochloa
Leptophyllochloa
Leptostigma
Lepuropetalon
Leucheria
Leucocoryne
Leunisia
Libertia
Ligaria
Lilaea
Lilaeopsis
Limonium
Limosella
Linanthus
Lindernia
Linum
Lippia
Lithrea
Littorella
Llagunoa
Loasa
Lobelia
Lomatia
Lophopappus
Lophosoria
Lotus
Lucilia
Luciliocline
Ludwigia
Luma
Lupinus
Luzula
Luzuriaga
Lycapsus
Lycium
Lycopersicon
Lycopodium
Lycopodiales
Solanales
Solanales
Asterales
Liliales
Poales
Fabales
Myrtales
Myrtales
Asterales
Asterales
Fabales
Dicksoniales
Asterales
Proteales
Asterales
Cornales
Sapindales
Lamiales
Sapindales
Lamiales
Malpighiales
Lamiales
Ericales
Lamiales
Caryophyllales
Apiales
Alismatales
Santalales
Asparagales
Asterales
Asparagales
Asterales
Saxifragales
Gentianales
Poales
Poales
Lycopodiaceae
Solanaceae
Solanaceae
Asteraceae
Luzuriagaceae
Juncaceae
Fabaceae
Myrtaceae
Onagraceae
Asteraceae
Asteraceae
Fabaceae
Lophosoriaceae
Asteraceae
Proteaceae
Campanulaceae
Loasaceae
Sapindaceae
Plantaginaceae
Anacardiaceae
Verbenaceae
Linaceae
Linderniaceae
Polemoniaceae
Gratiolaceae
Plumbaginaceae
Apiaceae
Juncaginaceae
Loranthaceae
Iridaceae
Asteraceae
Alliaceae
Asteraceae
Saxifragaceae
Rubiaceae
Poaceae
Poaceae
55
10
80
1
3
75
220
4
81
12
12
100
1
6
12
330
36
3
3
4
200
200
100
35
15
250
13
1
2
10
1
11
46
1
6
1
17
trop & temp
S Am Colombia to Chile, Brazil
N Am, tem S Am, Eurasia, S Afr, Aus, IP
Desventuradas Islands (Chile)
1 NZ, 3 Chile, Arg to T Fuego
cosmop, JF
N & S Am, Medit, montane trops, E Afr
Chile Arg
Cosmop
Andes S Am, Chile, Arg, Peru
Andes S Am, Chile, Arg, Peru
Macaronesia, temp eurasia, Afr, Aus, S Am
trop Am
Aus, Tas, S Am
trops & subtrops
Chile, Arg, 1 sp coastal Peru
Andean trop S Am
1 N Am, 1 S Am, 1 Euro
S Am: Arg, Bol, Paraguay, SE Brazil, Chile.
Am trop, many in Afr trop
temp & subtrops
trops, subtrops, temp, Madag
W N Am, Chile
Cosmop, Madag
N temperate, circumboreal, austral
Am Canada to TFuego, Aus, NZ
Pacific Am Canada to Chile
C Peru, E Bol, C Chile, Urug, n Arg, S Brasil
New Guinea, E Aus, NZ, Andes S Am, JF
C Chile
Chile
Peru, Bol, Arg, Chile, Falkland
SE USA, Mexico, Chile, Arg Uruguay
SE Asia, NZ, W S Am
Arg Chile, JF
tropics & subtrop
AND
AUS
W-ANT
AND
AND
W-NT
AND
AUS
AND
AND
W-ANT
S-AMZ
ANT-P
W-ANT
C-PAC
ANT-P
S-AMZ
SPT
AND
ANT-P
SPT
7
3
7
6
2
7
4
5
7
3
3
7
3
3
2
1
3
3
4
3
1
7
7
4
7
4
4
4
3
2
6
6
3
4
2
5
1
5
1
9
1
3
6
6
2
4
2
3
1
1
2
3
8
36
1
1
1
2
5
1
1
1
2
1
1
1
4
1
11
43
1
1
1
3
-37,00
-18,27
-18,00
-34,77
-30,00
-18,00
-29,50
-28,50
-17,58
-33,00
-24,83
-35,22
-18,00
-30,28
-25,22
-21,03
-29,73
-39,67
-30,00
-23,50
-23,00
-38,60
-31,00
-18,00
-27,07
-18,25
-33,22
-30,50
-32,23
-30,80
-20,83
-17,58
-32,57
-36,75
-38,00
-18,42
-56,00
-28,88
-35,33
-56,00
-56,00
-40,00
-45,42
-40,60
-23,50
-38,67
-40,00
-46,42
-20,08
-50,00
-43,83
-50,00
-34,13
-55,00
-38,25
-30,50
-37,58
-38,60
-37,67
-55,00
-32,55
-53,67
-42,05
-37,00
-49,15
-32,75
-40,00
-55,00
-40,00
-42,47
-40,00
-24,00
-46,50
-23,58
-26,67
-45,38
-43,00
-29,00
-37,46
-34,55
-20,54
-35,83
-32,42
-40,82
-19,04
-40,14
-34,53
-35,52
-31,93
-47,33
-34,13
-27,00
-30,29
-38,60
-34,33
-36,50
-29,81
-35,96
-37,63
-33,75
-40,69
-31,78
-30,42
-36,29
-36,28
-39,61
-39,00
-21,21
19,00
10,62
17,33
0,00
21,23
26,00
22,00
15,92
12,10
5,92
5,67
15,17
11,20
2,08
19,72
18,62
28,97
4,40
15,33
8,25
7,00
14,58
0,00
6,67
37,00
5,48
35,42
8,83
6,50
16,92
1,95
19,17
37,42
7,43
5,72
2,00
5,58
235
Asterids
Monocots
Asterids
Asterids
Rosids
Eudicots
Eudicots
Eudicots
Asterids
Rosids
Rosids
Rosids
Rosids
Filicopsida
Monocots
Asterids
Monocots
Rosids
Eudicots
Asterids
Monocots
Filicopsida
Monocots
Asterids
Asterids
Rosids
Asterids
Rosids
Filicopsida
Eudicots
Asterids
Asterids
Filicopsida
Asterids
Monocots
Asterids
Lysimachia
Machaerina
Macrachaenium
Madia
Magallana
Maihuenia
Maihueniopsis
Maireana
Malacothrix
Malesherbia
Malvella
Mancoa
Margyricarpus
Marsilea
Marsippospermum
Marticorenia
Mastigostyla
Mathewsia
Maytenus
Mecardonia
Megalachne
Megalastrum
Melica
Melosperma
Menodora
Menonvillea
Mentzelia
Metharme
Microlepia
Microphyes
Micropsis
Microseris
Microsorum
Microsteris
Miersia
Mikania
Asterales
Asparagales
Ericales
Polypodiales
Asterales
Asterales
Caryophyllales
Dennstaedtiales
Zygophyllales
Cornales
Brassicales
Lamiales
Lamiales
Poales
Dryopteridales
Poales
Lamiales
Celastrales
Brassicales
Asparagales
Asterales
Poales
Marsileales
Rosales
Brassicales
Malvales
Malpighiales
Asterales
Caryophyllales
Caryophyllales
Caryophyllales
Brassicales
Asterales
Asterales
Poales
Ericales
Asteraceae
Alliaceae
Polemoniaceae
Polypodiaceae
Asteraceae
Asteraceae
Caryophyllaceae
Dennstaedtiaceae
Zygophyllaceae
Loasaceae
Brassicaceae
Oleaceae
Plantaginaceae
Poaceae
Dryopteridaceae
Poaceae
Gratiolaceae
Celastraceae
Brassicaceae
Iridaceae
Asteraceae
Juncaceae
Marsileaceae
Rosaceae
Brassicaceae
Malvaceae
Malesherbiaceae
Asteraceae
Amaranthaceae
Cactaceae
Cactaceae
Tropaeolaceae
Asteraceae
Asteraceae
Cyperaceae
Myrsinaceae
430
1
1
40
15
8
3
45
1
80
26
25
2
80
40
2
10
200
7
16
1
3
70
3
7
4
24
16
58
18
2
2
10
1
45
150
pantrops
Chile
W N Am & S Am
trop & warm, Madag
W N Am, 1 Chile, 1 Aus, NZ
Brazil, Arg, Urug, Chile
Chile
paleotrop, 1 Afr, Madag, Japan, NZ
N Chile
Chile, Arg to Canada, WI
Chile, Arg
trop & subtrop Am, S Afr
Chile, Arg
temp regs & montane trops
Trop Am, 1 Afr to Mascarenes (no Madag), JF
JF
Am S US to Arg
trops & subtrops
Peru, Chile
Peru, Arg
Chile
NZ, S S Am, Tfuego, Falkland
trop & temp, Madag
S Andes to Chile, JF, Brazil
Mexico, Andes Peru, Bol, Arg, Chile
3 sp W USA, Mexico, 1 sp Peru to Uruguay, 1 sp. Medit
Andes Peru, Chile, Arg
W N Am, Chile
Australia, Chile
Andes Peru, Bol, Chile, Arg
Chile Arg
Chile, Arg
W N Am, Chile-Arg
Patagonia & Tfuego
trops esp Australia, Hawaii, JF
Cosmop
ANT-P
C-PAC
S-AMZ
W-NT
W-ANT
W-NT
AND
AND
AUS
S-AMZ
ANT-P
W-ANT
AND
ANT-P
AUS
AND
ANT-P
1
6
4
1
4
3
6
1
6
3
5
4
5
4
1
6
3
1
3
3
6
2
7
3
4
4
3
4
2
3
5
5
4
5
1
7
1
1
1
1
1
1
3
1
1
7
19
1
2
7
1
2
1
4
6
1
1
3
1
1
2
1
18
2
1
11
1
1
2
1
1
1
-34,50
-31,00
-18,00
-25,00
-31,33
-24,67
-20,08
-22,08
-22,08
-32,05
-31,00
-25,10
-30,67
-35,33
-28,50
-22,00
-18,17
-32,72
-37,00
-30,63
-25,00
-18,00
-25,00
-18,50
-22,00
-27,68
-19,42
-35,55
-46,53
-30,00
-38,27
-36,33
-37,92
-35,33
-30,50
-37,83
-36,50
-37,50
-20,63
-33,33
-51,17
-32,63
-36,50
-39,88
-46,68
-35,47
-55,00
-42,00
-23,00
-33,60
-56,00
-32,15
-43,67
-25,00
-33,68
-35,00
-30,67
-27,68
-30,00
-37,50
-46,53
-55,00
-55,00
-41,67
-36,21
-33,17
-24,25
-31,42
-33,92
-31,08
-20,36
-27,71
-36,63
-32,34
-33,75
-32,49
-38,68
-35,40
-41,75
-32,00
-20,58
-33,16
-46,50
-31,39
-34,33
-21,50
-29,34
-26,75
-26,33
-27,68
-24,71
-36,53
-46,53
-42,50
-46,63
-39,00
3,42
4,33
12,50
0,00
12,83
5,17
12,83
0,00
0,55
11,25
29,08
0,58
5,50
14,78
16,02
0,00
0,13
26,50
20,00
4,83
0,88
19,00
1,52
18,67
7,00
8,68
16,50
8,67
0,00
10,58
1,95
0,00
25,00
16,73
0,00
5,33
236
Rosids
Rosids
Eudicots
Rosids
Asterids
Eudicots
Asterids
Asterids
Asterids
Monocots
Asterids
Asterids
Eudicots
Asterids
Myriophyllum
Myrteola
Nama
Nanodea
Nardophyllum
Nasa
Nassauvia
Nassella
Nastanthus
Navarretia
Neowerdermannia
Nertera
Monocots
Muhlenbergia
Myrcianthes
Eudicots
Muehlenbeckia
Myrceugenia
Asterids
Moschopsis
Eudicots
Asterids
Moscharia
Myosurus
Rosids
Morella
Asterids
Asterids
Monttea
Eudicots
Eudicots
Montiopsis
Myosotis
Eudicots
Montia
Myoschilos
Asterids
Mniodes
Asterids
Asterids
Mitraria
Mutisia
Eudicots
Misodendrum
Monocots
Eudicots
Mirabilis
Munroa
Eudicots
Miqueliopuntia
Asterids
Eudicots
Minuartia
Mulinum
Asterids
Mimulus
Gentianales
Caryophyllales
Ericales
Asterales
Poales
Asterales
Cornales
Asterales
Santalales
Unplaced
Myrtales
Saxifragales
Myrtales
Myrtales
Ranunculales
Unplaced
Santalales
Asterales
Poales
Apiales
Poales
Caryophyllales
Asterales
Asterales
Fagales
Lamiales
Caryophyllales
Caryophyllales
Asterales
Lamiales
Santalales
Caryophyllales
Caryophyllales
Caryophyllales
Lamiales
Rubiaceae
Cactaceae
Polemoniaceae
Calyceraceae
Poaceae
Asteraceae
Loasaceae
Asteraceae
Santalaceae
Boraginaceae
Myrtaceae
Haloragaceae
Myrtaceae
Myrtaceae
Ranunculaceae
Boraginaceae
Santalaceae
Asteraceae
Poaceae
Apiaceae
Poaceae
Polygonaceae
Calyceraceae
Asteraceae
Myricaceae
Plantaginaceae
Portulacaceae
Portulacaceae
Asteraceae
Gesneriaceae
Misodendraceae
Nyctaginaceae
Cactaceae
Caryophyllaceae
Phrymaceae
13
2
30
9
79
39
100
7
1
45
3
60
50
40
15
100
1
60
5
20
150
25
8
2
40
3
18
20
4
1
8
54
1
120
150
Madag, SE Asia, Malesia, Aus, NZ, Hawaii, S Am, JF
W N Am, Chile, Arg
Chile, Arg, Falkland Is.
C & S Am
Bol, Arg, Chile
Colombia, Ecuador, Peru, few in Chile, Bolivia, Venezuela and C Am (to S Mexico).
Bol, Arg, Chile
temp S Am
N Am from California,Mex, C Am to Brasil, WI, Hawaii
Colombia to Chile, JF, Brazil
subcosmop
trop Am, Andes, Brazil, WI
Chile, Arg, SE Brasil, JF
temp regs hemis N & S, M apetalus disj N Am
Eurasia, montane trops, S Afr, N Guinea, Aus, NZ
Chile, Arg
Andes Colombia to Chile, Arg, Parag, SE Brasil, Urug
1 W US, 4 SW S Am
S Andes
temp & subtrop Am, Asia
New Guinea, Aus, Tasm, NZ, W S Am
Perú, Chile, Arg, Patagonia
Chile
Sudam Andes Peru Ecuador, Africa, E Asia, Filipinas y
Malasia
Chile 1, Arg. 2
Chile, Argentina, Bolivia y Perú
temp Eurasia, Medit, montane tropics, Afr, A Aus, A & S
Am,
Peru, Chile
Chile, Arg.
S Am, south of 33
Am trop, Himal, China
Chile
Arctic & temp regions, Mexico, Ethiopia, Himalayas,
Chile
Am, S Afr, Madag, E Asia, JF
C-AUS
AND
ANT-P
W-NT
AND
W-NT
AND
W-NT
AND
AND
S-AMZ
W-ANT
W-ANT
AND
ANT-P
W-ANT
SPT
AND
AND
AND
AND
W-ANT
W-ANT
2
3
4
5
3
3
3
3
5
3
3
7
3
3
4
4
5
3
4
5
4
2
3
6
1
3
3
7
3
5
5
4
6
4
7
1
1
1
5
26
25
1
5
1
2
1
2
1
12
2
2
1
23
2
7
3
1
4
2
1
1
18
1
1
1
8
4
1
1
8
-30,67
-18,50
-33,25
-25,00
-17,67
-30,00
-20,00
-23,00
-40,33
-21,50
-36,23
-19,50
-29,43
-30,67
-29,80
-48,47
-31,62
-17,58
-18,18
-20,98
-19,50
-19,65
-17,67
-29,72
-18,83
-25,00
-19,25
-29,50
-17,58
-30,67
-33,00
-18,33
-27,50
-33,00
-22,00
-56,00
-18,50
-40,53
-46,00
-50,00
-55,67
-25,00
-53,00
-56,00
-25,50
-56,00
-55,00
-30,00
-46,43
-53,38
-54,88
-55,00
-48,92
-24,00
-45,48
-41,00
-45,50
-55,00
-36,50
-19,12
-31,67
-40,75
-55,00
-18,17
-49,88
-56,00
-36,00
-30,00
-36,75
-45,50
-43,33
-18,50
-36,89
-35,50
-33,83
-42,83
-22,50
-38,00
-48,17
-23,50
-46,12
-37,25
-29,72
-38,55
-41,59
-51,68
-43,31
-33,25
-21,09
-33,23
-30,25
-32,58
-36,33
-33,11
-18,98
-28,33
-30,00
-42,25
-17,88
-40,28
-44,50
-27,17
-28,75
-34,88
-33,75
25,33
0,00
7,28
21,00
32,33
25,67
5,00
30,00
15,67
4,00
19,77
35,50
0,57
15,77
23,58
6,42
23,38
31,33
5,82
24,50
21,50
25,85
37,33
6,78
0,28
6,67
21,50
25,50
0,58
19,22
23,00
17,67
2,50
3,75
23,50
237
Asterids
Rosids
Asterids
Asterids
Eudicots
Rosids
Asterids
Eudicots
Rosids
Filicopsida
Monocots
Asterids
Rosids
Monocots
Rosids
Monocots
Eudicots
Rosids
Filicopsida
Asterids
Monocots
Eudicots
Asterids
Asterids
Eudicots
Asterids
Monocots
Asterids
Rosids
Asterids
Rosids
Rosids
Monocots
Asterids
Eudicots
Nesocaryum
Neuontobothrys
Nicotiana
Nierembergia
Nitrophila
Noccaea
Nolana
Notanthera
Nothofagus
Notholaena
Nothoscordum
Noticastrum
Nototriche
Ochagavia
Oenothera
Olsynium
Ombrophytum
Onuris
Ophioglossum
Ophryosporus
Oreobolus
Oreocereus
Oreomyrrhis
Oreopolus
Orites
Orobanche
Ortachne
Osmorhiza
Otholobium
Ourisia
Ovidia
Oxalis
Oxychloë
Oxyphyllum
Oxytheca
Caryophyllales
Asterales
Poales
Oxalidales
Malvales
Lamiales
Fabales
Apiales
Poales
Lamiales
Proteales
Gentianales
Apiales
Caryophyllales
Poales
Asterales
Ophioglossales
Brassicales
Santalales
Asparagales
Myrtales
Poales
Malvales
Asterales
Asparagales
Pteridales
Fagales
Santalales
Solanales
Brassicales
Caryophyllales
Solanales
Solanales
Brassicales
Unplaced
Polygonaceae
Asteraceae
Juncaceae
Oxalidaceae
Thymelaeaceae
Plantaginaceae
Fabaceae
Apiaceae
Poaceae
Orobanchaceae
Proteaceae
Rubiaceae
Apiaceae
Cactaceae
Cyperaceae
Asteraceae
Ophiglossaceae
Brassicaceae
Balanophoraceae
Iridaceae
Onagraceae
Bromeliaceae
Malvaceae
Asteraceae
Alliaceae
Pteridaceae
Nothofagaceae
Loranthaceae
Solanaceae
Brassicaceae
Amaranthaceae
Solanaceae
Solanaceae
Brassicaceae
Boraginaceae
7
1
6
600
2
25
36
10
2
130
9
1
23
7
15
37
26
5
4
20
121
4
100
20
26
40
37
1
77
20
8
20
67
2
1
W N Am, Chile, Arg
N Chile
Andes S Am
Cosmop
temp S Am
Tasm, NZ, Andes S Am
E & S Afr, temp S Am
N Am, E Asia, temp S Am
Arg, Chile
temp, subtrop, N Hemis
6 temp E Aus, 3 Andes S Am
Chile, Arg, Patagonia
E Asia, Borneo, N Guinea, SE Aus, NZ, Mex, C Am,
Andes, Tfuego, Falkland
Malesia, Aus, Tasm, NZ, Hawaii, JF, trop & temp Am,
Chile, Tfuego, Falkland
trop & subtrop S Am, Brazil
Chile, Patagonia
Peru, W Brasil, N Arg, Chile
Andean and temp S Am and NW N Am
temp & subtrop N C & S Am
Chile, JF
Ecuador, Peru, Bol, Chile, Arg
Peru, Bol, Chile, Arg, Ecuador, Brazil
Am
warm to trop Am, SW US, Mex, Madag, JF
New Guinea, Aus, Tasm, NZ, New Caled, temp S Am
Chile temperate, JF
Peru to Chile, Galapagos
N temp, few S Am
US, Mex, Chile, Arg
Mex to Arg, Bras
Am, USA to Chile, Aust, New Caledonia, Namibia, JF
Chile
ANT-P
AND
AUS
W-ANT
AUS
SPT
AND
SPT
S-AMZ
AND
W-NT
W-NT
AND
S-AMZ
W-NT
SPT
AND
W-ANT
ANT-P
W-NT
4
6
3
7
5
2
1
4
5
7
2
5
2
3
2
3
7
5
3
3
3
6
3
3
3
1
2
6
3
4
4
3
1
6
6
1
1
1
117
2
13
2
3
2
2
1
1
1
3
1
5
3
5
1
16
14
3
24
5
6
2
10
1
36
1
1
1
10
2
1
-24,00
-24,83
-18,00
-18,00
-37,80
-18,23
-29,83
-30,83
-39,48
-18,00
-35,83
-33,00
-52,00
-18,17
-36,50
-17,58
-33,10
-37,58
-18,20
-23,50
-18,00
-31,67
-17,72
-31,92
-29,90
-17,58
-32,95
-32,57
-18,25
-33,50
-22,50
-34,83
-18,47
-20,13
-31,50
-26,15
-34,25
-56,00
-45,93
-56,00
-40,00
-54,93
-56,00
-33,50
-38,25
-54,50
-54,00
-22,50
-56,00
-36,67
-54,83
-54,67
-22,12
-55,00
-52,00
-38,25
-34,67
-38,73
-40,25
-27,28
-55,97
-43,00
-40,17
-54,83
-24,50
-51,00
-43,62
-20,13
-27,75
-25,49
-26,13
-37,00
-41,87
-37,12
-34,92
-42,88
-47,74
-25,75
-37,04
-43,75
-53,00
-20,33
-46,25
-27,13
-43,97
-46,13
-20,16
-39,25
-35,00
-34,96
-26,19
-35,33
-35,08
-22,43
-44,46
-37,78
-29,21
-44,17
-23,50
-42,92
-31,04
-20,13
7,50
1,32
16,25
38,00
8,13
37,77
10,17
24,10
16,52
15,50
2,42
21,50
2,00
4,33
19,50
19,08
21,73
17,08
3,92
31,50
34,00
6,58
16,95
6,82
10,35
9,70
23,02
10,43
21,92
21,33
2,00
16,17
25,15
0,00
0,00
238
Asterids
Eudicots
Monocots
Monocots
Asterids
Monocots
Asterids
Monocots
Eudicots
Asterids
Rosids
Pinopsida
Rosids
Philippiella
Phleum
Phragmites
Phrodus
Phycella
Phyllachne
Phylloscirpus
Phytolacca
Picrosia
Pilea
Pilgerodendron
Pilostyles
Magnoliids
Peperomia
Philibertia
Monocots
Pennisetum
Monocots
Asterids
Pelletiera
Philesia
Filicopsida
Pellaea
Monocots
Asterids
Pectocarya
Asterids
Monocots
Patosia
Phalaris
Rosids
Passiflora
Phacelia
Monocots
Paspalum
Magnoliids
Monocots
Pasithea
Peumus
Eudicots
Paronychia
Magnoliids
Rosids
Parietaria
Persea
Asterids
Parastrephia
Asterids
Monocots
Panicum
Asterids
Rosids
Palaua
Perityle
Asterids
Pachylaena
Perezia
Monocots
Oziroe
Malpighiales
Pinales
Rosales
Asterales
Caryophyllales
Poales
Asterales
Asparagales
Solanales
Poales
Poales
Caryophyllales
Gentianales
Liliales
Poales
Unplaced
Laurales
Laurales
Asterales
Asterales
Piperales
Poales
Ericales
Pteridales
Unplaced
Poales
Malpighiales
Poales
Asparagales
Caryophyllales
Rosales
Asterales
Poales
Malvales
Asterales
Asparagales
Rafflesiaceae
Cupressaceae
Urticaceae
Asteraceae
Phytolaccaceae
Cyperaceae
Stylidiaceae
Amaryllidaceae
Solanaceae
Poaceae
Poaceae
Caryophyllaceae
Apocynaceae
Philesiaceae
Poaceae
Boraginaceae
Monimiaceae
Lauraceae
Asteraceae
Asteraceae
Piperaceae
Poaceae
Myrsinaceae
Pteridaceae
Boraginaceae
Juncaceae
Passifloraceae
Poaceae
Hemerocallidaceae
Caryophyllaceae
Urticaceae
Asteraceae
Poaceae
Malvaceae
Asteraceae
Hyacinthaceae
25
1
250
2
25
2
4
3
1
4
15
1
40
1
20
200
1
120
63
32
1000
115
2
35
13
1
500
330
1
100
15
5
600
15
2
5
Am US to Magallanes, Asia, 2 W Aus
Chile Arg
trop & warm excl Australasia
subtrop S Am, Chile, Peru, Arg, Brazil
trop & warm
Arg, Chile
Tasm, NZ, temp S Am
Chile, Arg
Chile (Atacama & Coquimbo)
Cosmop
temperate Eurasia, America
Patagonia Chile-Arg.
Andes Perú, Chile, Bolivia, Argentina
Chile, Arg
temperate Eurasia, America
W N Am, S Am
Chile
trops & subtrops Am & Asia, Macaronesia
SW N Am, 1 Chile Peru
Andes from Colombia to Chile, SE Brazil
trop & warm esp Am, JF, Hawaii
trop & temp regs
1 subtrop S Am, 1 Macaronesia
trop & warm-temp, Madag
W N Am, S Am
Andes Chile Arg Bol
trop & subtrop Am, 1 Madag, 20 trop E Asia, Malesia,
3 Aus, NZ
temp & trop regs, IP
Peru, Chile
North temperate, austral
subcosmop, JF, ID
Andes Bol, Chile, Perú, Arg
temp, subtrop, trop regs
Coastal Peru and Chile
Andes N Chile Arg
Chile, Arg, Perú, Bol, Paraguay
S-AMZ
AUS
AND
W-ANT
AND
W-ANT
ANT-P
W-ANT
ANT-P
AND
W-ANT
ANT-P
AND
AND
W-ANT
AND
AND
AND
AND
1
5
1
3
1
5
2
3
6
7
4
5
3
5
4
4
6
4
4
3
1
4
1
1
4
3
1
7
3
4
7
3
7
3
3
3
1
1
2
1
1
2
1
3
1
1
1
1
1
1
2
6
1
2
1
23
4
1
1
2
4
1
1
5
1
5
2
5
2
4
2
4
-18,18
-39,90
-37,62
-34,50
-37,13
-22,00
-45,00
-29,67
-26,43
-19,67
-29,50
-48,00
-18,33
-39,83
-30,83
-18,00
-30,67
-33,08
-20,80
-17,58
-24,83
-20,00
-32,50
-18,27
-23,58
-24,00
-31,97
-20,50
-22,00
-18,18
-20,80
-17,58
-33,50
-20,38
-28,00
-18,40
-36,90
-54,25
-45,42
-36,72
-41,88
-51,00
-56,00
-36,62
-31,33
-40,00
-55,17
-50,00
-22,27
-55,50
-39,88
-54,83
-40,50
-43,18
-29,90
-56,00
-42,50
-33,53
-40,00
-40,32
-35,97
-38,00
-32,50
-42,50
-40,00
-37,80
-35,62
-25,00
-37,67
-25,20
-35,80
-36,00
-27,54
-47,08
-41,52
-35,61
-39,51
-36,50
-50,50
-33,14
-28,88
-29,83
-42,33
-49,00
-20,30
-47,67
-35,36
-36,42
-35,58
-38,13
-25,35
-36,79
-33,67
-26,77
-36,25
-29,29
-29,78
-31,00
-32,23
-31,50
-31,00
-27,99
-28,21
-21,29
-35,58
-22,79
-31,90
-27,20
18,72
14,35
7,80
2,22
4,75
29,00
11,00
6,95
4,90
20,33
25,67
2,00
3,93
15,67
9,05
36,83
9,83
10,10
9,10
38,42
17,67
13,53
7,50
22,05
12,38
14,00
0,53
22,00
18,00
19,62
14,82
7,42
4,17
4,82
7,80
17,60
239
Filicopsida
Asterids
Rosids
Monocots
Rosids
Asterids
Filicopsida
Monocots
Asterids
Asterids
Asterids
Asterids
Asterids
Filicopsida
Rosids
Filicopsida
Asterids
Eudicots
Monocots
Monocots
Asterids
Pinopsida
Monocots
Asterids
Asterids
Eudicots
Rosids
Eudicots
Rosids
Filicopsida
Monocots
Filicopsida
Rosids
Eudicots
Monocots
Asterids
Pilularia
Pinguicula
Pintoa
Piptochaetium
Pitavia
Pitraea
Pityrogramma
Placea
Plagiobothrys
Plantago
Plazia
Plectritis
Pleocarphus
Pleopeltis
Pleurophora
Pleurosorus
Pluchea
Plumbago
Poa
Podagrostis
Podanthus
Podocarpus
Podophorus
Polemonium
Polyachyrus
Polycarpon
Polygala
Polygonum
Polylepis
Polypodium
Polypogon
Polystichum
Porlieria
Portulaca
Potamogeton
Pouteria
Ericales
Alismatales
Caryophyllales
Zygophyllales
Dryopteridales
Poales
Polypodiales
Rosales
Caryophyllales
Fabales
Caryophyllales
Asterales
Ericales
Poales
Pinales
Asterales
Poales
Poales
Caryophyllales
Asterales
Aspleniales
Myrtales
Polypodiales
Asterales
Dipsacales
Asterales
Lamiales
Unplaced
Asparagales
Pteridales
Lamiales
Sapindales
Poales
Zygophyllales
Lamiales
Marsileales
Sapotaceae
Potamogetonaceae
Portulacaceae
Zygophyllaceae
Dryopteridaceae
Poaceae
Polypodiaceae
Rosaceae
Polygonaceae
Polygalaceae
Caryophyllaceae
Asteraceae
Polemoniaceae
Poaceae
Podocarpaceae
Asteraceae
Poaceae
Poaceae
Plumbaginaceae
Asteraceae
Aspleniaceae
Lythraceae
Polypodiaceae
Asteraceae
Valerianaceae
Asteraceae
Plantaginaceae
Boraginaceae
Amaryllidaceae
Pteridaceae
Verbenaceae
Rutaceae
Poaceae
Zygophyllaceae
Lentibulariaceae
Marsileaceae
200
95
40
5
160
13
150
15
90
500
16
7
40
1
100
2
4
350
20
80
4
9
10
1
5
3
270
70
5
16
1
1
30
1
85
6
trop. Am, Afr, Asia, Aus, NZ, New Caled
Cosmop
trop, subtrop & temp regions
S USA, Mexico, S Am
Cosmop, Madag, IP
Cosmop
Cosmop, JF
Andes Venezuela to Arg, Chile
Cosmop
subcosmop
Cosmop
Peru, Chile
temp Eurasia, N Am, Chile
JF
Afr, Madag, Asia, Aus, Tasm, NZ, Mex, C & S Am, WI
Chile
Arg, Chile, Canadá, USA
temp & cold & montane trops
trop & warm-temp regs
Asia, N & S Am, Afr, Aus
España, Marruecos, Aus, Tasm, NZ, Chile, Arg
S Am, Brazil
trop Am, 1 Afr-Madag, India, Sri Lanka
N Chile
4 W N Am, 1 Chile
Peru, Bol, Arg, Chile
Cosmop, JF, ID
W N Am, South Am, E Asia, Aus
Chile
Am trop desde S US, Afr, Madag
S Am, Chile, Peru, Arg
Chile
Am US to Arg, JF
Chile
Am, Medit, circumboreal
Euro, Aus, NZ, Ethiopia, W S Am
ANT-P
AND
AND
W-ANT
ANT-P
W-ANT
W-ANT
S-AMZ
ANT-P
AND
C-PAC
AND
W-NT
W-ANT
1
7
7
4
7
7
7
3
7
7
7
3
4
6
1
6
4
4
7
4
7
3
1
6
4
3
7
4
6
1
3
6
3
6
4
7
1
6
2
1
5
4
3
2
1
8
1
7
1
1
2
2
1
61
1
1
1
3
1
1
1
2
18
19
5
1
1
1
7
1
2
1
-32,10
-21,50
-18,33
-30,00
-31,30
-18,00
-23,52
-17,72
-30,00
-29,00
-30,00
-17,58
-30,83
-35,83
-30,28
-35,25
-18,00
-24,75
-18,42
-32,25
-25,00
-24,00
-29,15
-32,50
-18,18
-18,17
-29,83
-31,00
-18,50
-18,47
-35,33
-29,50
-27,33
-35,50
-32,15
-33,03
-55,00
-28,80
-34,28
-55,00
-46,67
-52,88
-22,00
-46,75
-54,00
-30,00
-34,73
-54,00
-50,38
-38,25
-40,33
-56,00
-31,93
-19,12
-45,58
-34,42
-42,40
-33,50
-37,33
-21,95
-56,00
-54,00
-34,55
-18,67
-33,83
-37,53
-40,00
-28,88
-56,00
-40,28
-32,57
-38,25
-23,57
-32,14
-43,15
-32,33
-38,20
-19,86
-38,38
-41,50
-30,00
-26,16
-42,42
-43,11
-34,27
-37,79
-37,00
-28,34
-18,77
-38,92
-29,71
-33,20
-31,33
-34,92
-20,07
-37,08
-41,92
-32,78
-18,58
-26,15
-36,43
-34,75
-28,11
-45,75
-36,22
0,93
33,50
10,47
4,28
23,70
28,67
29,37
4,28
16,75
25,00
0,00
17,15
23,17
0,00
14,55
7,97
5,08
38,00
7,18
0,70
13,33
9,42
18,40
4,35
4,83
3,77
37,83
24,17
3,55
0,17
15,37
2,20
10,50
1,55
20,50
8,13
240
Asterids
Asterids
Asterids
Rosids
Asterids
Pinopsida
Asterids
Asterids
Psilotopsida
Filicopsida
Eudicots
Rosids
Monocots
Monocots
Eudicots
Monocots
Rosids
Eudicots
Eudicots
Eudicots
Monocots
Rosids
Asterids
Rosids
Asterids
Monocots
Monocots
Monocots
Eudicots
Asterids
Rosids
Monocots
Monocots
Rosids
Eudicots
Filicopsida
Pozoa
Pratia
Primula
Prosopis
Proustia
Prumnopitys
Pseudopanax
Psilocarphus
Psilotum
Pteris
Pterocactus
Pteromonnina
Puccinellia
Puya
Pycnophyllum
Pycreus
Quillaja
Quinchamalium
Ranunculus
Reicheella
Relchela
Retanilla
Reyesia
Rhamnus
Rhaphithamnus
Rhodophiala
Rhombolytrum
Rhynchospora
Ribes
Robinsonia
Rorippa
Rostkovia
Rostraria
Rubus
Rumex
Rumohra
Dryopteridales
Caryophyllales
Rosales
Poales
Poales
Brassicales
Asterales
Saxifragales
Poales
Poales
Asparagales
Lamiales
Rosales
Solanales
Rosales
Poales
Caryophyllales
Ranunculales
Santalales
Fabales
Poales
Caryophyllales
Poales
Poales
Fabales
Caryophyllales
Pteridales
Psilotales
Asterales
Apiales
Pinales
Asterales
Fabales
Ericales
Asterales
Apiales
Dryopteridaceae
Polygonaceae
Rosaceae
Poaceae
Juncaceae
Brassicaceae
Asteraceae
Grossulariaceae
Cyperaceae
Poaceae
Amaryllidaceae
Verbenaceae
Rhamnaceae
Solanaceae
Rhamnaceae
Poaceae
Caryophyllaceae
Ranunculaceae
Santalaceae
Quillajaceae
Cyperaceae
Caryophyllaceae
Bromeliaceae
Poaceae
Polygalaceae
Cactaceae
Pteridaceae
Psilotaceae
Asteraceae
Araliaceae
Podocarpaceae
Asteraceae
Fabaceae
Primulaceae
Campanulaceae
Apiaceae
7
200
500
2
2
75
7
150
250
2
30
2
160
4
3
1
1
500
25
3
70
17
190
80
130
9
280
2
8
10
8
3
44
400
15
2
circumaustral, J Fernandez, Madag
cosmop
cosmop, JF
Arg, Caribe, Chil, Mexico, Peru, USA ,Urug
NZ, S S Am, Tfuego, Falkland
cosmop
JF
N Hemis, N Afr, S Am Canada to Tfuego
cosmp
S Brasil, Urug, Chile
Chile, Arg, Bol, Bras
Chile, Arg, JF
N Hemisp, S Afr, S Am
Chile Arg
Chile, Arg
Chile Arg
Chile
Cosmop, R acaulis disj NZ, Falkland, R bonariensis
disj N Am S Am
Andes Bol, Chile, Perú, Arg
temp S Am, Arg, Chile, Peru, Brazil
pantrops
Andes Peru, Chile, Arg, Bol
C & S Am, Brazil
N temp: Asia, N Am, S temp: S Afr, Aus, S Am
Am SW US to Chile
Patagonia Arg, Chile
Cosmop, Madag, JF
pantrop, Hawaii, Australia, NZ, Madag, IP
W US, temp S Am
Tasm, NZ, Chile
Costa Rica to Chile, New Caled, NZ, E Aus
Bol, Chile, Arg, Peru
trop & subtrop Am, Afr, W Asia
N Hemisp, trop Asia, temp S Am
SE Asia tropical, Aus, NZ, Am Austral
Andes Chile Arg
C-AUS
W-NT
AUS
W-ANT
S-AMZ
S-AMZ
W-ANT
AND
AND
S-AMZ
AND
W-NT
W-NT
ANT-P
AUS
AUS
AND
W-ANT
SPT
2
7
7
3
2
7
6
4
7
3
3
5
4
3
5
5
6
7
3
3
1
3
3
7
3
5
7
1
4
2
2
3
1
4
2
5
1
10
2
1
1
6
6
13
1
1
9
1
1
4
2
1
1
17
16
1
1
5
8
8
4
1
2
1
2
2
1
3
6
1
1
2
-30,67
-30,00
-36,90
-25,00
-46,43
-33,60
-29,83
-37,83
-33,00
-25,00
-30,67
-35,83
-19,10
-31,62
-35,85
-18,00
-17,78
-18,18
-30,00
-18,17
-22,50
-20,35
-29,00
-46,50
-32,55
-30,00
-35,33
-35,83
-18,00
-19,00
-38,50
-18,83
-31,50
-55,08
-55,00
-55,67
-26,38
-56,00
-43,83
-56,00
-39,23
-35,62
-40,00
-45,42
-41,00
-30,20
-37,80
-42,08
-22,58
-54,92
-51,00
-38,23
-24,73
-37,80
-56,00
-37,83
-46,50
-49,70
-38,00
-52,33
-39,50
-40,33
-34,33
-55,83
-56,00
-38,58
-42,88
-42,50
-46,28
-25,69
-51,22
-38,72
-42,92
-38,53
-34,31
-32,50
-38,04
-38,42
-24,65
-34,71
-38,97
-20,29
-36,35
-34,59
-34,12
-21,45
-30,15
-38,18
-33,42
-46,50
-41,13
-34,00
-43,83
-37,67
-29,17
-26,67
-47,17
-37,42
-35,04
24,42
25,00
18,77
1,38
9,57
10,23
0,00
26,17
1,40
2,62
15,00
14,75
5,17
11,10
6,18
6,23
4,58
37,13
32,82
8,23
0,00
6,57
15,30
35,65
8,83
0,00
17,15
0,00
8,00
17,00
3,67
22,33
15,33
17,33
37,17
7,08
241
Monocots
Monocots
Eudicots
Monocots
Rosids
Asterids
Asterids
Asterids
Asterids
Eudicots
Asterids
Eudicots
Eudicots
Asterids
Asterids
Pinopsida
Eudicots
Eudicots
Eudicots
Rosids
Filicopsida
Asterids
Asterids
Rosids
Asterids
Monocots
Monocots
Asterids
Asterids
Asterids
Asterids
Asterids
Rosids
Filicopsida
Rosids
Asterids
Ruppia
Rytidosperma
Sagina
Sagittaria
Salix
Salpichroa
Salpiglossis
Salvia
Samolus
Sanctambrosia
Sanicula
Santalum
Sarcocornia
Sarmienta
Satureja
Saxe-gothaea
Saxifraga
Saxifragella
Saxifragodes
Schinus
Schizaea
Schizanthus
Schizeilema
Schizopetalon
Schkuhria
Schoenus
Scirpus
Scutellaria
Scyphanthus
Selkirkia
Selliera
Senecio
Senna
Serpyllopsis
Sicyos
Sigesbeckia
Asterales
Cucurbitales
Hymenophyllales
Fabales
Asterales
Asterales
Unplaced
Cornales
Lamiales
Poales
Poales
Asterales
Brassicales
Apiales
Solanales
Schizaeales
Sapindales
Saxifragales
Saxifragales
Saxifragales
Pinales
Lamiales
Lamiales
Caryophyllales
Santalales
Apiales
Caryophyllales
Ericales
Lamiales
Solanales
Solanales
Malpighiales
Alismatales
Caryophyllales
Poales
Alismatales
Asteraceae
Cucurbitaceae
Hymenophyllaceae
Fabaceae
Asteraceae
Goodeniaceae
Boraginaceae
Loasaceae
Lamiaceae
Cyperaceae
Cyperaceae
Asteraceae
Brassicaceae
Apiaceae
Solanaceae
Schizaeceae
Anacardiaceae
Saxifragaceae
Saxifragaceae
Saxifragaceae
Podocarpaceae
Lamiaceae
Gesneriaceae
Amaranthaceae
Santalaceae
Apiaceae
Caryophyllaceae
Samolaceae
Lamiaceae
Solanaceae
Solanaceae
Salicaceae
Alismataceae
Caryophyllaceae
Poaceae
Ruppiaceae
3
50
1
300
1250
2
1
2
360
20
100
6
10
13
12
30
27
1
1
350
1
38
1
16
25
40
1
12
900
2
15
450
25
25
28
10
subtro & trop regs
Aus, Tasm, NZ, Hawaii, N & S Am, ID
S S Am, JF, Falkland
pantrops
Cosmop
Aus, NZ, Chile
JF
Chile
cosmop
North temperate, austral, JF, IP
subcosmop
trop & subtrop Am
C N Chile, Arg
11 NZ, 1 Aus, 1 S S Am
Chile, Arg
trop & austral, N Am, Madag
Mexico to Arg
Tfuego
Antarctic S Am
N temp, Eurasia, W N Am, Andes S Am
Chile, Arg
temp & subtrop, Medit
Chile
Eurasia, Afr, Aus, Tasm, NZ, N & S Am, JF
Indomalasia to Aus, Hawaii, JF (ex)
subcosmop, exc Aus & NZ
Cosmop, IP
cosmop
Chile, Arg
S Am: Ecuad, Peru, Bol, Bra, Parag, Uru, Chile, Arg
Arctic temp, few in trops, temp S Am, 1 S Afr
Cosmop
temp N Hemis, E Afr, Asia, N Guinea, Aus, NZ, Andes
Australia, NZ, Arg, Chile, IP
cosmop
C-PAC
AUS
W-NT
AND
AUS
AND
W-NT
W-ANT
SPT
AND
W-ANT
AUS
1
4
5
1
7
2
6
6
7
7
7
3
3
2
3
1
3
5
5
4
5
7
6
7
2
7
6
7
7
5
3
4
7
7
2
7
1
1
1
9
218
1
1
2
3
20
4
2
10
1
12
1
9
1
1
1
1
4
1
3
1
2
1
3
4
2
2
1
1
1
6
2
-32,83
-20,17
-39,63
-18,00
-17,58
-28,45
-31,00
-35,33
-17,67
-35,83
-17,58
-26,43
-38,58
-21,03
-39,88
-18,00
-51,00
-50,72
-36,00
-36,00
-18,50
-30,60
-20,63
-30,67
-33,42
-22,00
-27,00
-18,25
-18,58
-36,50
-31,50
-30,67
-21,00
-42,72
-33,40
-56,00
-40,00
-56,00
-42,72
-38,23
-53,00
-56,00
-56,00
-31,83
-35,33
-54,48
-41,00
-49,15
-46,55
-54,83
-56,00
-55,00
-45,75
-54,00
-43,83
-54,00
-44,50
-55,00
-25,33
-40,00
-19,87
-38,78
-40,00
-42,50
-51,00
-55,00
-37,78
-26,78
-47,82
-29,00
-36,79
-35,58
-34,62
-44,17
-36,83
-45,92
-24,71
-30,88
-46,53
-31,02
-44,52
-32,28
-52,92
-53,36
-45,50
-40,88
-36,25
-37,22
-37,32
-37,58
-44,21
-23,67
-33,50
-19,06
-28,68
-38,25
-37,00
-40,83
-38,00
9,88
13,23
16,37
22,00
38,42
14,27
0,00
7,23
17,67
38,33
20,17
14,25
8,90
15,90
19,97
9,27
28,55
3,83
5,28
19,00
9,75
35,50
13,23
33,37
0,00
13,83
0,00
21,58
3,33
13,00
1,62
20,20
3,50
11,00
20,33
34,00
242
Eudicots
Rosids
Monocots
Asterids
Asterids
Monocots
Monocots
Asterids
Asterids
Rosids
Monocots
Monocots
Eudicots
Rosids
Asterids
Monocots
Asterids
Asterids
Eudicots
Asterids
Asterids
Monocots
Asterids
Monocots
Monocots
Eudicots
Asterids
Monocots
Rosids
Rosids
Asterids
Asterids
Monocots
Rosids
Asterids
Rosids
Silene
Sisymbrium
Sisyrinchium
Skytanthus
Solanum
Solaria
Solenomelus
Solidago
Soliva
Sophora
Spartina
Speea
Spergularia
Sphaeralcea
Spilanthes
Sporobolus
Stachys
Stangea
Stellaria
Stemodia
Stenandrium
Stenomesson
Stevia
Stipa
Stuckenia
Suaeda
Tagetes
Tapeinia
Tara
Tarasa
Taraxacum
Tecoma
Tecophilaea
Tepualia
Tessaria
Tetilla
Geraniales
Asterales
Myrtales
Asparagales
Lamiales
Asterales
Malvales
Fabales
Asparagales
Asterales
Caryophyllales
Alismatales
Poales
Asterales
Asparagales
Lamiales
Lamiales
Caryophyllales
Dipsacales
Lamiales
Poales
Asterales
Malvales
Caryophyllales
Asparagales
Poales
Fabales
Asterales
Asterales
Asparagales
Asparagales
Solanales
Gentianales
Asparagales
Brassicales
Caryophyllales
Francoaceae
Asteraceae
Myrtaceae
Tecophilaeaceae
Bignoniaceae
Asteraceae
Malvaceae
Fabaceae
Iridaceae
Asteraceae
Amaranthaceae
Potamogetonaceae
Poaceae
Asteraceae
Amaryllidaceae
Acanthaceae
Gratiolaceae
Caryophyllaceae
Valerianaceae
Lamiaceae
Poaceae
Asteraceae
Malvaceae
Caryophyllaceae
Alliaceae
Poaceae
Fabaceae
Asteraceae
Asteraceae
Iridaceae
Alliaceae
Solanaceae
Apocynaceae
Iridaceae
Brassicaceae
Caryophyllaceae
1
1
1
2
12
60
30
3
1
50
100
6
300
230
40
40
45
150
5
300
160
6
60
30
1
20
47
8
150
2
2
1700
3
60
90
600
Chile
S Am, Brazil
Chile, Arg
Chile
Am tropical Arizona to Chile, 1 S Afr, Aus?
temp N Hemis & temp S Am, NZ, JF
2 sp Mexico, disjunct to Peru, Chile, Arg
Mexico, C Am to S Am
Chile, Arg
N & S Am
cosmop, ID
Subcosmop
temp N Hemis, montane trops, IP
S US to Chile
Andes
S US to C Chile
Trops and Subtrops, Madag
cosmop
Arg,Chil,Perú
trop, subtrop, temp regions exc. Australasia
Am, Asia, Afr, 1 Euro
trops
W N Am, Mex, temperate S Am
cosmop, JF, ID
Chile
Am, W Euro, NW Afr
trop & most N temp, JF, IP
S Am, N Am, Aus
Temperate America, Macaronesia, Eurasia
Chile, Arg
Chile, Arg
subcosmop, JF, ID, IP
Brasil, Chile
N, C & S Am
temp Eurasia, Medit, S Afr, Andes, N Am
temp N hemis, Medit, S Afr
S-AMZ
W-ANT
ANT-P
W-NT
W-NT
W-NT
AND
W-NT
W-ANT
AND
ANT-P
W-ANT
C-PAC
W-ANT
AND
S-AMZ
W-NT
W-ANT
6
3
5
6
1
4
4
3
5
3
7
7
7
3
3
3
4
7
3
7
7
1
4
7
6
4
7
4
4
5
3
7
3
3
4
7
1
1
1
2
1
2
10
1
1
3
4
3
1
2
1
1
1
5
1
11
1
1
2
13
2
1
2
4
2
2
1
51
1
11
23
8
-31,50
-18,42
-35,40
-30,00
-18,47
-33,45
-18,17
-18,47
-40,50
-17,58
-20,25
-18,00
-17,58
-18,00
-30,00
-29,00
-18,00
-17,67
-22,00
-23,62
-18,47
-29,45
-18,00
-32,92
-36,67
-32,00
-29,95
-30,95
-31,00
-33,00
-18,20
-25,20
-24,50
-18,20
-18,00
-35,42
-36,50
-53,00
-33,00
-18,50
-56,00
-38,78
-18,47
-56,00
-36,60
-55,00
-53,00
-25,43
-18,17
-37,80
-38,00
-55,83
-18,08
-44,00
-23,62
-18,47
-37,78
-40,00
-34,27
-37,50
-45,42
-41,83
-44,00
-50,00
-36,83
-46,50
-28,93
-55,25
-55,00
-55,00
-33,46
-27,46
-44,20
-31,50
-18,48
-44,73
-28,48
-18,47
-48,25
-27,09
-37,63
-35,50
-21,51
-18,08
-33,90
-33,50
-36,92
-17,88
-33,00
-23,62
-18,47
-33,62
-29,00
-33,59
-37,08
-38,71
-35,89
-37,48
-40,50
-34,92
-32,35
-27,07
-39,88
-36,60
-36,50
3,92
18,08
17,60
3,00
0,03
22,55
20,62
0,00
15,50
19,02
34,75
35,00
0,00
7,85
0,17
7,80
9,00
37,83
0,42
22,00
0,00
0,00
8,33
22,00
1,35
0,83
13,42
11,88
13,05
19,00
3,83
28,30
3,73
30,75
36,80
37,00
243
Asterids
Rosids
Eudicots
Monocots
Asterids
Asterids
Filicopsida
Filicopsida
Monocots
Monocots
Asterids
Monocots
Rosids
Eudicots
Asterids
Filicopsida
Monocots
Monocots
Rosids
Monocots
Asterids
Monocots
Asterids
Monocots
Monocots
Eudicots
Rosids
Asterids
Rosids
Rosids
Eudicots
Asterids
Monocots
Rosids
Monocots
Asterids
Tetrachondra
Tetraglochin
Tetragonia
Tetroncium
Teucrium
Thamnoseris
Thelypteris
Thyrsopteris
Tigridia
Tillandsia
Tiquilia
Traubia
Trevoa
Tribeles
Trichocline
Trichomanes
Trichoneura
Trichopetalum
Trifolium
Triglochin
Triodanis
Tripogon
Triptilion
Trisetum
Tristagma
Tristerix
Triumfetta
Trixis
Tropaeolum
Tropidocarpum
Tunilla
Tweedia
Typha
Ugni
Uncinia
Urbania
Lamiales
Poales
Myrtales
Poales
Gentianales
Caryophyllales
Brassicales
Brassicales
Asterales
Malvales
Santalales
Asparagales
Poales
Asterales
Poales
Asterales
Alismatales
Fabales
Asparagales
Poales
Hymenophyllales
Asterales
Saxifragales
Rosales
Asparagales
Unplaced
Poales
Asparagales
Dennstaedtiales
Blechnales
Asterales
Lamiales
Alismatales
Caryophyllales
Rosales
Lamiales
Verbenaceae
Cyperaceae
Myrtaceae
Typhaceae
Apocynaceae
Cactaceae
Brassicaceae
Tropaeolaceae
Asteraceae
Malvaceae
Loranthaceae
Alliaceae
Poaceae
Asteraceae
Poaceae
Campanulaceae
Juncaginaceae
Fabaceae
Laxmanniaceae
Poaceae
Hymenophyllaceae
Asteraceae
Saxifragaceae
Rhamnaceae
Amaryllidaceae
Boraginaceae
Bromeliaceae
Iridaceae
Dicksoniaceae
Thelypteridaceae
Asteraceae
Lamiaceae
Juncaginaceae
Aizoaceae
Rosaceae
Tetrachondraceae
1
60
10
13
6
9
4
50
50
70
11
6
70
12
30
8
12
250
2
7
80
22
1
1
1
27
540
35
1
800
1
250
1
70
8
2
Chile, Arg
Malesia, Pacific, Aus, SE Asia, C & S Am, JF
Am from Mex to Chile, JF
Cosmop
Bol, Uru, Chile, Arg
California, Chile
Mexico to Chile
SW N Am to Chile
Pantrop
Andes Colombia to Chile
Chile, Arg, Uru
cold & temp, montane tropics, JF
C Chile, 1 Patagonia
trops & subtrops
N Am, Mex, Guatem, Medit
N temperate, circumboreal, austral
temp & subtrop exc Aus
Chile, Arg
Arabia, trop Afr, S US, Peru
patrop, Madag, JF
S Am, 1 W Aus
Chile & TFuego
Chile
Chile
arid N & S Am
trop & subtrop Am, WI
Mex, Guatemala disjunct to Peru, Chile, Bol, Brazil
JF
trops & subtrops, Madag
Cosmop
temp S Am, Magallanes
N & S Afr, SE Asia, Aus, Tasm, NZ, Polinesia, Peru,
Chile
Arg, Chil, Perú
1 NZ, 1 Patagonia, Tfuego
AND
SPT
W-NT
S-AMZ
AND
ANT-P
W-NT
W-NT
AND
W-ANT
W-ANT
W-ANT
AUS
ANT-P
W-NT
ANT-P
W-ANT
AND
AUS
3
2
3
7
3
3
4
3
3
1
3
5
7
5
1
4
4
4
5
1
1
2
5
6
6
4
3
4
6
1
6
7
5
4
3
2
1
10
2
2
2
2
1
18
1
1
3
12
14
12
1
1
3
8
1
1
1
5
1
1
1
5
4
1
1
1
1
2
1
10
3
1
-18,00
-30,67
-34,17
-19,17
-26,75
-18,00
-30,67
-22,08
-18,55
-28,50
-29,00
-25,00
-23,00
-18,38
-25,00
-20,33
-25,00
-25,43
-18,47
-39,87
-18,43
-40,50
-30,42
-31,50
-18,18
-18,00
-22,00
-18,27
-25,00
-43,00
-21,03
-28,72
-49,00
-24,73
-56,00
-45,38
-39,00
-37,67
-22,00
-33,38
-44,00
-20,93
-42,50
-54,83
-56,00
-42,50
-23,58
-40,00
-55,00
-40,00
-38,23
-18,47
-43,10
-37,82
-55,50
-35,83
-34,00
-29,00
-38,00
-25,40
-37,33
-40,00
-56,00
-31,92
-46,17
-54,83
-21,37
-43,33
-39,78
-29,08
-32,21
-20,00
-32,03
-33,04
-19,74
-35,50
-41,92
-40,50
-32,75
-20,98
-32,50
-37,67
-32,50
-31,83
-18,47
-41,48
-28,13
-48,00
-33,13
-32,75
-23,59
-28,00
-23,70
-27,80
-32,50
-49,50
-26,48
-37,44
-51,92
6,73
25,33
11,22
19,83
10,92
4,00
2,72
21,92
2,38
0,00
14,00
25,83
31,00
19,50
5,20
15,00
34,67
15,00
12,80
0,00
3,23
19,38
15,00
5,42
2,50
10,82
20,00
3,40
0,00
19,07
0,00
15,00
13,00
10,88
17,45
5,83
244
Asterids
Rosids
Asterids
Asterids
Asterids
Rosids
Asterids
Asterids
Asterids
Rosids
Asterids
Asterids
Rosids
Filicopsida
Rosids
Monocots
Asterids
Rosids
Asterids
Rosids
Rosids
Rosids
Asterids
Monocots
Monocots
Filicopsida
Asterids
Rosids
Rosids
Asterids
Monocots
Monocots
Rosids
Monocots
Monocots
Urmenetea
Urtica
Utricularia
Valdivia
Valeriana
Vasconcellea
Verbena
Verbesina
Vestia
Vicia
Viguiera
Villanova
Viola
Vittaria
Viviania
Vulpia
Wahlenbergia
Weberbauera
Wedelia
Weinmannia
Wendtia
Werdermannia
Werneria
Wolffia
Wolffiella
Woodsia
Xenophyllum
Xerodraba
xMargyracaena
Yunquea
Zameioscirpus
Zannichellia
Zanthoxylum
Zephyra
Zoellnerallium
Asparagales
Asparagales
Sapindales
Alismatales
Poales
Asterales
Rosales
Brassicales
Asterales
Dryopteridales
Alismatales
Alismatales
Asterales
Brassicales
Geraniales
Oxalidales
Asterales
Brassicales
Asterales
Poales
Geraniales
Pteridales
Malpighiales
Asterales
Asterales
Fabales
Solanales
Asterales
Lamiales
Brassicales
Dipsacales
Unplaced
Lamiales
Rosales
Asterales
Alliaceae
Tecophilaeaceae
Rutaceae
Potamogetonaceae
Cyperaceae
Asteraceae
Rosaceae
Brassicaceae
Asteraceae
Dryopteridaceae
Lemnaceae
Lemnaceae
Asteraceae
Brassicaceae
Ledocarpaceae
Cunoniaceae
Asteraceae
Brassicaceae
Campanulaceae
Poaceae
Vivianiaceae
Vittariaceae
Violaceae
Asteraceae
Asteraceae
Fabaceae
Solanaceae
Asteraceae
Verbenaceae
Caricaceae
Valerianaceae
Escalloniaceae
Lentibulariaceae
Urticaceae
Asteraceae
2
1
250
5
3
1
1
6
21
25
10
11
40
3
3
160
100
18
150
25
6
70
430
10
180
140
1
300
225
20
230
1
200
75
1
Chile, Arg
Chile
Am, Afr, Asia, JF
cosmop
Arg, Chile, Bol
JF
JF
S Arg, Chile
Andes Colombia to N Arg, Chile
temp & cool temp Eurasia, Afr, Am, exc Aus
trop & warm Am, 1 S Afr
cosmop, warm temperate and tropical regions
Andes Ecuador to Chile
N Chile, Arg
Chile, Arg
Madag, Australasia, Mex, C & S Am
pantrops, W Australia
Andes Peru, Chile, Arg, Bol
Southern Hemisphere, Europe, SE Asia
N & S Am, Euro, N Afr, M Asia
S Brasil, Chile, Arg
pantrop
cosmop
Mex to Chile
temp N Hemis, Medit, few trop Afr, S Am, Hawaii
Chile
temp & trop Am, few OW
Mexico to N Arg & S Brasil
N temp Eurasia, S Afr, Andes
Chile
cosmop
subcosmop, temperate
N Chile, NW Arg
AND
AND
AND
W-ANT
AND
AND
C-AUS
AND
W-ANT
S-AMZ
W-NT
W-NT
W-ANT
W-NT
W-ANT
W-NT
W-ANT
AND
3
6
1
7
3
6
6
5
3
4
1
7
3
3
5
2
1
3
4
7
3
1
7
3
3
4
6
3
4
3
4
6
7
7
3
2
1
2
1
3
1
1
2
5
1
1
1
9
2
1
1
1
7
1
2
3
1
70
2
6
28
1
2
33
1
43
1
1
10
1
-30,00
-20,82
-22,00
-17,58
-51,00
-18,00
-18,22
-33,33
-36,75
-17,58
-21,27
-30,50
-35,33
-23,58
-30,67
-30,00
-23,33
-28,72
-18,00
-18,20
-18,18
-25,00
-33,03
-22,00
-18,00
-29,67
-18,00
-39,88
-28,72
-17,72
-21,50
-33,83
-29,00
-54,00
-33,67
-51,00
-23,58
-18,35
-36,75
-36,75
-33,82
-26,33
-41,73
-49,42
-27,37
-36,90
-40,00
-37,33
-36,83
-56,00
-25,00
-37,80
-55,33
-40,00
-29,90
-40,00
-33,00
-55,00
-40,25
-43,00
-55,50
-23,18
-31,92
-24,91
-38,00
-25,63
-51,00
-20,79
-18,28
-35,04
-36,75
-25,70
-23,80
-36,12
-42,38
-25,48
-33,78
-35,00
-30,33
-32,78
-37,00
-21,60
-27,99
-40,17
-36,52
-25,95
-29,00
-31,33
-36,50
-40,07
-35,86
-36,61
-22,34
3,83
8,18
0,00
32,00
16,08
0,00
0,00
0,00
5,58
0,13
3,42
0,00
16,23
5,07
11,23
14,08
3,78
6,23
10,00
14,00
8,12
0,00
38,00
6,80
19,62
30,33
6,97
7,90
22,00
3,33
37,00
0,37
14,28
37,78
1,68
Appendix A (continuation): Legend and summary for tracks and elements
TRACKS (follow subsections in the text)
N° of genera
AUS
31
SPT
25
C-AUS
3
W-NT
64
3.3.2 Andean track
AND
113
3.3.3 South Amazonian track
S-AMZ
39
3.4.1 Wide antitropical track
W-ANT
84
ANT-P
56
C-PAC
9
ELEMENTS
N° of genera
1. Pantropical
88
2. Australasiatic
59
3. Neotropical
216
4. Antitropical
152
5. South-temperate
81
6. Endemic
83
7. Cosmopolitan
134
TOTAL
813
245
246
Appendix B: Genera shared by several Chilean regions
247
GEN
Summe
ANT
Aa
1
1
BIO
COQ
Abrotanella
2
Acacia
2
Acaena
5
1
GEN
Summe
Laretia
1
Larrea
2
Lastarriaea
1
Lasthenia
3
Acantholippia
2
1
1
Lathyrus
4
Acrisione
2
1
1
Laurelia
1
1
Adenocaulon
1
Adenopeltis
2
Laureliopsis
1
1
1
1
Lebetanthus
1
Adesmia
4
1
1
1
1
Legrandia
1
Adiantum
5
1
1
1
1
Lemna
3
1
Aextoxicon
2
Agalinis
1
1
1
Lenzia
2
1
1
Lepidium
4
1
Agallis
1
1
Lepidoceras
1
Ageratina
3
1
1
Agoseris
3
1
1
Agrostis
4
1
1
Allionia
1
1
Alona
2
1
Alonsoa
2
1
Alopecurus
3
1
1
Aloysia
1
Alstroemeria
4
1
Alternanthera
1
1
Amaranthus
1
Amblyopappus
2
1
Ambrosia
3
1
Amomyrtus
1
Amphibromus
1
Amsimkia
1
Amsinckia
3
1
1
1
Anagallis
4
1
1
1
Anarthrophyllum
2
1
Anatherostipa
1
Androsace
1
Anemone
3
Anisomeria
2
Antennaria
3
Anthochloa
1
Anthoxanthum
2
1
Antidaphne
1
Aphanes
1
Apium
5
Apodasmia
2
1
Arachnitis
2
1
Araeoandra
1
Araucaria
1
Arenaria
4
1
Argemone
2
1
Argylia
3
1
Aristeguietia
2
Aristida
3
Aristolochia
1
Aristotelia
2
1
1
1
1
JF
MAG
1
1
1
1
1
1
1
COQ
JF
MAG
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Lepidophyllum
1
1
Lepidothamnus
1
1
1
Leptinella
1
Leptocarpha
1
1
1
Leptophyllochloa
2
1
1
Leptostigma
1
1
Lepuropetalon
1
Leucheria
4
1
1
1
Leucocoryne
3
1
1
1
Leunisia
1
Libertia
3
1
Ligaria
2
1
Lilaeopsis
4
1
Limonium
1
1
Limosella
4
Linanthus
1
Lindernia
1
1
Linum
3
1
1
Lippia
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
1
Loasa
4
1
Lobelia
3
locenes
1
Lomatia
3
1
Lophosoria
2
1
Lotus
3
1
Ludwigia
1
Luma
1
Lupinus
3
Luzula
4
Luzuriaga
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
2
1
1
2
1
1
1
1
1
1
1
2
1
Lycopersicon
1
1
1
1
Lycopodium
3
1
1
1
Lysimachia
1
1
1
1
Machaerina
1
1
Macrachaenium
1
1
Madia
3
1
1
1
Lycium
248
1
1
1
1
1
1
1
Llagunoa
1
1
1
1
1
1
1
1
1
1
1
2
1
1
1
Littorella
1
1
1
1
Lithrea
1
1
1
1
1
1
BIO
1
1
1
ANT
1
1
1
1
1
1
1
1
1
1
1
Arjona
2
Armeria
3
Artemisia
2
Arthropteris
1
Asplenium
5
Astelia
1
Aster
3
1
Asteranthera
2
1
Asteriscium
3
1
1
1
Astragalus
4
1
1
1
Atriplex
4
1
1
1
Austrocedrus
1
1
Azara
3
1
1
Azolla
4
1
1
1
Azorella
4
1
1
1
Baccharis
4
1
1
1
Bacopa
2
1
Bahia
3
1
Bakerolimon
2
1
Balbisia
2
1
Balsamocarpon
Barneoudia
1
1
1
Maihuenia
1
1
1
Maihueniopsis
2
1
Malacothamnus
1
Malacothrix
2
1
1
1
Malesherbia
2
1
1
1
Malvella
2
1
1
1
Mancoa
1
1
1
Margyricarpus
4
1
Marsilea
1
1
Marsippospermum
2
1
Mastigostyla
1
1
Mathewsia
2
1
Maytenus
3
1
Megalachne
1
1
Megalastrum
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3
1
1
Melica
2
1
1
1
Melosperma
2
1
1
1
Menodora
1
1
Menonvillea
4
1
1
Mentzelia
2
1
1
1
Microphyes
3
1
1
1
Micropsis
2
Bartsia
1
1
Microseris
3
Beilschmiedia
1
Microsteris
3
Belloa
2
Miersia
1
Benthamiella
1
Mikania
1
Berberidopsis
1
Mimulus
5
Berberis
5
1
1
1
Minuartia
1
Bidens
3
1
1
1
Miqueliopuntia
1
Bipinnula
3
1
1
1
Mirabilis
2
Blechnum
4
1
1
Misodendrum
2
1
Blennosperma
1
1
Mitraria
3
1
1
1
Blepharocalyx
1
1
Montia
3
1
1
1
Boehmeria
1
Montiopsis
3
1
1
1
Boerhavia
2
1
1
Monttea
2
1
Boisduvalia
2
1
1
Moscharia
2
Bolax
1
Moschopsis
1
Bomarea
1
1
Moschopsìs
1
Boopis
2
1
Muehlenbeckia
2
Boquila
1
1
Muhlenbergia
2
1
Botrychium
1
Mulinum
4
1
Bouteloua
1
1
Munroa
1
1
Bowlesia
4
1
1
Brachyclados
1
Brachystele
2
Bridgesia
1
Bromidium
2
Bromus
5
Buddleja
Bulbostylis
Bulnesia
1
Caesalpinia
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
Myosotis
1
1
Myosurus
3
1
1
Myrceugenia
3
1
1
Myrcianthes
1
Myriophyllum
4
Myrteola
3
1
Nama
1
1
Nanodea
1
2
1
1
1
1
249
1
1
1
1
1
1
1
3
1
1
1
3
1
1
1
Myoschilos
1
1
1
1
Mutisia
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Caiophora
3
1
1
1
Nardophyllum
3
Calamagrostis
4
1
1
1
1
Nassauvia
3
Calandrinia
4
1
1
1
1
Nassella
5
Calceolaria
4
1
1
1
1
Nastanthus
3
Caldcluvia
1
Nasturtium
1
Calliandra
1
Navarretia
2
1
Callitriche
4
Nertera
4
1
Calopappus
1
Nicandra
1
Calotheca
1
Nicotiana
4
Caltha
3
Calycera
3
Calydorea
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Nierembergia
1
1
Nitrophila
2
1
2
1
1
Nolana
3
1
Calystegia
2
1
Notanthera
2
Camissonia
2
1
Campsidium
2
1
Cardamine
4
1
1
Cardionema
3
1
1
1
Carex
5
1
1
1
Carica
1
Carpha
1
Carpobrotus
3
Castilleja
2
Catabrosa
3
1
Centaurea
3
1
1
1
Centaurium
3
1
1
1
Centaurodendron
1
Centella
2
1
Centipeda
2
1
Cerastium
3
Ceratophyllum
1
Cestrum
3
1
1
Chaetanthera
3
1
1
Chaetotropis
2
1
1
Chaptalia
1
1
Chascolytrum
2
1
1
Cheilanthes
3
1
1
1
Chenopodium
5
1
1
1
Chersodoma
1
1
Chevreulia
1
Chiliophyllum
1
Chiliotrichum
2
Chiropetalum
3
Chloraea
3
Chorizanthe
2
Chrysosplenium
1
Chuquiraga
2
Chusquea
3
1
1
Cicendia
2
1
1
Ciclospermum
1
Cissarobryon
1
Cissus
2
Cistanthe
3
1
1
1
2
2
1
Nothoscordum
2
1
Noticastrum
1
1
Nototriche
2
Ochagavia
3
Oenothera
4
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Olsynium
4
1
Ombrophytum
1
1
Onuris
1
Ophioglossum
4
1
1
1
1
1
Ophryosporus
2
Oreobolus
3
1
Oreocereus
1
Oreomyrrhis
1
Oreopolus
2
1
Orites
1
1
1
Ornithopus
1
1
1
Orobanche
1
Ortachne
1
Osmorhiza
3
1
1
Otholobium
2
1
1
Ourisia
3
1
1
1
Ovidia
1
1
Oxalis
4
1
1
1
Oxychloe
2
1
1
Oxyphyllum
1
1
1
Oxytheca
2
1
Oziroe
3
1
Pachylaena
1
Palaua
1
Panicum
1
Parastrephia
1
1
Parietaria
3
1
Paronychia
3
1
1
1
Pasithea
3
1
1
1
Paspalum
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Passiflora
1
1
1
Patosia
3
250
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Notholaena
1
1
1
Nothofagus
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Citronella
2
1
1
Pectocarya
3
1
1
1
Clarkia
2
1
1
Pellaea
3
1
1
1
Cleome
1
Pelletiera
1
Cliococca
1
1
Pennisetum
2
1
Codonorchis
2
1
Peperomia
3
1
Colletia
3
1
1
Perezia
4
1
Colliguaja
3
1
1
1
Collomia
2
Colobanthus
4
Combera
1
Conanthera
3
1
1
1
Convolvulus
3
1
1
1
Conyza
4
1
1
1
Copiapoa
2
1
Coprosma
1
Cordia
1
Coriaria
1
1
Coronopus
2
1
1
Corrigiola
2
1
1
Cortaderia
4
1
1
Corynabutilon
2
1
1
Cotula
1
1
Crassula
4
1
Cressa
2
1
Crinodendron
1
Cristaria
2
1
Croton
1
1
Cruckshanksia
2
1
Cryptantha
3
1
Cryptocarya
1
1
1
1
1
1
1
1
2
1
1
1
Peumus
2
1
1
Phacelia
4
1
1
Phalaris
2
1
1
Philesia
1
Philibertia
1
Philippiella
1
Phleum
2
Phragmites
3
Phrodus
1
Phycella
2
Phyllachne
1
Phylloscirpus
1
Phytolacca
1
1
Picrosia
1
1
Pilea
1
1
Pilgerodendron
1
Pilostyles
3
Pilularia
2
Pinguicola
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Pinguicula
1
1
Piptochaetium
3
1
2
1
1
Pitavia
1
1
Cryptogramma
2
1
1
Pitraea
2
Cuatrecasasiella
1
Placea
1
Cuminia
1
Plagiobothrys
3
Cumulopuntia
2
1
1
Plantago
5
Cuscuta
3
1
1
1
Plazia
1
Cyclospermum
2
1
1
Plectritis
1
Cynodon
2
1
1
Pleocarphus
1
Cynoglossum
1
1
Pleopeltis
3
1
Cyperus
4
1
Pleurophora
2
1
Cyphocarpus
1
Pleurosorus
2
Cystopteris
4
Pluchea
1
1
Dalea
1
Plumbago
2
1
Danthonia
3
1
Poa
4
1
Dasyphyllum
1
1
Podagrostis
1
1
Daucus
3
Podanthus
2
1
Dendroseris
1
Podocarpus
2
1
Dennstaedtia
1
Deschampsia
3
Descurainia
3
Desfontainia
2
1
Desmaria
1
1
Deuterocohnia
1
1
Dichondra
4
1
Dicksonia
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Podophorus
1
1
Polemonium
2
1
1
Polyachyrus
2
1
Polycarpon
1
Polygala
3
1
1
Polygonum
2
1
1
1
Polylepis
1
1
1
Polypodium
5
1
1
1
251
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Persea
1
1
1
1
Perityle
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Dicllptera
1
1
Polypogon
4
Dielsiochloa
1
1
Polystichum
4
Dinemagonum
1
Porlieria
1
Dinemandra
1
1
Portulaca
1
1
Dioscorea
3
1
1
Diostea
2
Diplachne
1
1
Diplolepis
3
1
Diplostephium
1
1
Diposis
1
Discaria
3
Distichia
1
1
Distichlis
3
1
Dodonaea
1
Domeykoa
1
Donatia
1
Doniophyton
2
Downingia
2
1
Draba
3
1
1
Drapetes
1
Drimys
4
Drosera
1
Drymaria
1
Dysopsis
4
Eccremocarpus
1
Echinopsis
2
Elaphoglossum
1
Elatine
1
Eleocharis
5
Elodea
1
Elymus
4
Elytropus
1
1
Embothrium
2
1
Empetrum
3
Encelia
2
1
Enneapogon
1
1
Ephedra
4
1
1
1
1
Epilobium
4
1
1
1
1
Epipetrum
3
1
1
Equisetum
2
1
Eragrostis
3
1
Ercilla
1
Erechtites
1
Eremocharis
1
Eriachaenium
1
Erigeron
4
Eriosyce
3
1
Errazurizia
2
1
Eryngium
4
1
Escallonia
Eucryphia
Eudema
2
1
Eulychnia
2
1
Euphorbia
4
1
1
1
1
1
1
1
1
Potamogeton
4
1
Pouteria
1
Pozoa
2
1
1
1
Pratia
2
1
Primula
1
Prosopis
2
1
Proustia
3
1
Prumnopitys
1
1
1
Pseudopanax
2
1
1
Psilocarphus
2
1
1
Pteris
3
1
1
Pteromonnina
2
1
1
1
Puccinellia
3
1
1
Puya
3
1
1
Pycnophyllum
1
1
1
Quillaja
2
1
Quinchamalium
3
1
1
Raimundochloa
2
1
Ranunculus
5
1
Reicheella
1
1
Relchela
1
Retanilla
2
Reyesia
2
Rhamnus
1
Rhaphithamnus
3
Rhodophiala
3
Ribes
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
Rostkovia
1
Rubus
3
1
Rumex
3
1
1
Rumohra
4
1
1
Ruppia
3
1
Rytidosperma
3
1
1
Sagittaria
1
1
1
Salix
3
Salpiglossis
2
1
Salvia
1
1
Samolus
2
1
1
Sanicula
2
1
1
Santalum
1
1
1
1
1
1
1
1
1
1
1
1
1
Rorippa
1
1
1
1
Robinsonia
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Sarcocornia
5
1
Sarmienta
2
Satureja
4
Saxegothaea
1
1
Saxifraga
2
1
Saxifragella
1
1
Saxifragodes
1
1
Schinus
2
1
1
1
4
1
1
1
1
1
1
1
1
1
1
1
252
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Euphrasia
4
1
Evolvulus
1
1
1
1
Exodeconus
1
1
Fabiana
3
1
1
1
Facelis
3
1
1
Fagonia
2
1
Fascicularia
1
Festuca
4
1
1
1
Flaveria
3
1
1
1
Flourensia
2
1
1
Fragaria
1
1
Francoa
1
Frankenia
4
Fuchsia
3
Fuertesimalva
2
Gaimardia
1
Galinsoga
3
1
1
1
Galium
5
1
1
1
1
Gamocarpha
2
Gamochaeta
5
1
1
Gamochaetopsis
1
Gaultheria
5
Gavilea
Gayophytum
Gentiana
4
Gentianella
Geoffroea
1
Schizaea
1
Schizanthus
3
Schizeilema
1
Schizopetalon
1
1
Schkuhria
2
1
1
Schlnus
1
1
Schoenus
2
Scirpus
5
Scutellaria
2
1
Scyphanthus
2
1
Selkirkia
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Selliera
2
1
1
1
Senecio
4
1
1
1
1
1
1
Senna
3
1
1
1
Serpyllopsis
2
Sicyos
2
Sigesbeckia
1
1
Silene
4
1
1
1
1
1
Sisymbrium
4
1
1
1
1
1
Sisyrinchium
4
1
1
1
1
Skytanthus
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Solanum
4
4
1
1
1
1
Solaria
1
1
2
1
1
Solenomelus
3
1
1
1
1
1
1
Solidago
2
1
1
4
1
1
1
1
Soliva
2
1
1
2
1
Sophora
3
1
1
1
Geranium
4
1
Spartina
1
1
Gethyum
1
Spergularia
4
1
1
1
Geum
2
1
Sphacele
2
1
1
Gevuina
1
1
Sphaeralcea
2
1
1
Gilia
3
Gilliesia
2
Glandularia
3
Gleichenia
3
1
Glinus
2
1
Glyceria
1
Glycyrrhiza
1
Gnaphalium
4
Gochnatia
2
Gomortega
1
Gomphrena
1
1
Grabowskia
1
1
Grammitis
3
Grammosperma
1
Gratiola
1
1
Greigia
2
1
Griselinia
4
Guindilia
1
Gunnera
4
Gutierrezia
3
Guynesomia
1
Gymnachne
1
Gymnophyton
2
1
1
1
1
1
1
1
1
1
Spirodela
1
1
1
Stachys
3
1
1
1
1
1
Stellaria
4
1
1
1
Stemodia
2
1
1
Stenandrium
2
1
1
Stevia
1
1
Stipa
4
1
1
1
1
Suaeda
3
1
1
1
Tagetes
3
1
1
Tapeinia
1
Tarasa
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
Tecophilaea
1
1
Tepualia
2
1
Tessaria
2
1
Tetilla
1
Tetrachondra
1
Tetraglochin
2
1
Tetragonia
2
1
Tetroncium
1
Teucrium
3
1
1
1
Thelypteris
3
1
1
1
Thlaspi
1
1
1
1
1
253
1
1
Taraxacum
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Gypothamnium
1
Habenaria
2
1
Thyrsopteris
1
Tigridia
1
1
Haloragis
1
Hamadryas
1
Tillandsia
3
1
Tiquilia
1
1
Haplopappus
3
Hebe
1
Traubia
1
Trevoa
2
Hedyotis
2
Tribeles
1
Helenium
3
Helictotrichon
1
Trichocline
3
Trichomanes
1
Heliotropium
4
1
Trichopetalum
2
Helogyne
1
1
Trifolium
3
Herbertia
1
Herreria
1
1
Triglochin
1
Triodanis
Heterosperma
1
Heterozostera
1
Hieracium
2
1
Hierochloe
2
1
Hippuris
2
1
Histiopteris
2
Hoffmannseggia
2
1
1
Homalocarpus
2
1
1
Hordeum
4
1
Hornungia
1
Huanaca
2
Huidobria
2
Huperzia
1
Hybanthus
1
1
Hydrangea
1
1
Hydrocotyle
3
1
Hymenoglossum
3
1
Hymenophyllum
4
1
Hypericum
2
1
1
Hypochaeris
4
1
1
1
Hypolepis
4
1
1
Hypsela
2
Imperata
2
Ipomopsis
1
Isoetes
3
Jaborosa
4
Jarava
3
Jovellana
1
Juania
1
Jubaea
1
Juncus
5
1
Junellia
3
1
Kageneckia
2
Koeleria
1
Krameria
2
Kurzamra
1
Lactoris
1
Lagenophora
3
Lampaya
1
Lapageria
2
1
Lardizabala
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
4
1
1
1
1
1
Triptilion
3
1
1
1
Trisetum
5
1
1
1
Tristagma
3
1
1
Tristerix
2
1
1
1
Tropaeolum
3
1
1
1
1
Tunilla
1
1
Tweedia
3
1
1
1
Typha
3
1
1
1
Ugni
2
Uncinia
4
Urbania
1
1
Urmenetea
1
1
Urtica
5
1
Utricularia
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3
1
1
1
1
1
Verbesina
2
1
1
1
Vestia
1
1
Vicia
4
1
1
1
Viguiera
3
1
1
1
1
Villanova
1
1
1
Viola
4
1
1
1
Viviania
2
1
1
1
1
Vulpia
3
1
1
1
1
1
Wahlenbergia
3
1
1
1
1
1
1
Weberbauera
3
1
1
1
1
1
Weinmannia
2
1
Wendtia
2
Werdermannia
1
1
1
1
1
1
Werneria
2
1
Wolffia
1
1
Wolffiella
1
Xenophyllum
1
Xerodraba
1
1
xMargyracaena
1
1
Yunquea
1
Zannichellia
3
Zanthoxylum
1
Zephyra
2
Zoellnerallium
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
254
1
1
1
1
1
1
4
1
1
1
Verbena
1
1
1
Valeriana
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Appendix C: Matrix for PAE: Distribution of Endemic Genera
255
N°
GENERO
A an
B co
B an
C co
C an
D co
D an
E co
E an
F co
F an
G co
G in
G an
H co
1
Acrisione
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
Adenopeltis
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
Alona
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
4
Anisomeria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
5
Araeoandra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
Avellanita
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
Bakerolimon
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
8
Balsamocarpon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9
Bridgesia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
10
Calopappus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
11
Cissarobryon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
12
Conanthera
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
13
Copiapoa
0
0
0
0
0
0
0
1
0
1
0
1
0
0
1
14
Cyphocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
15
Desmaria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
16
Dinemagonum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
17
Dinemandra
0
0
0
0
0
0
0
1
0
1
1
1
1
0
1
18
Epipetrum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
19
Ercilla
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
20
Fascicularia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
21
Francoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
22
Gethyum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
23
Gomortega
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
24
Guynesomia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
25
Gymnachne
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
26
Gypothamnium
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
Hollermayera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
28
Homalocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
29
Huidobria
0
0
0
1
1
0
1
1
0
1
0
1
0
1
1
30
Hymenoglossum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
31
Ivania
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
32
Jubaea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
33
Lapageria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
34
Latua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
35
Legrandia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
36
Leontochir
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
37
Leptocarpha
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
38
Leucocoryne
0
0
0
0
0
1
0
1
0
0
0
1
0
0
0
39
Leunisia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
40
Marticorenia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
41
Metharme
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
42
Microphyes
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
43
Miersia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
44
Miqueliopuntia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
45
Moscharia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
46
Neuontobothrys
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
47
Notanthera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
48
Ochagavia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
49
Oxyphyllum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
50
Peumus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
51
Phrodus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
52
Pintoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
53
Pitavia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
54
Placea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
55
Pleocarphus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
56
Podanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
57
Reicheella
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
58
Sarmienta
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
59
Scyphanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
60
Speea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
61
Tecophilaea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
62
Tetilla
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
63
Traubia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
64
Trevoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
65
Valdivia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
66
Vestia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
67
Zephyra
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
TOT
0
0
0
1
1
4
2
4
1
6
2
5
1
1
9
256
N°
GENERO
H an
I co
I an
J co
J an
K co
K an
L co
L an
M co
M an
N co
N an
1
Acrisione
0
0
0
0
0
0
0
0
0
0
0
0
0
2
Adenopeltis
0
0
0
0
0
0
0
0
0
0
0
1
0
3
Alona
0
1
0
1
0
1
0
1
0
1
0
1
0
4
Anisomeria
0
1
0
0
0
0
0
1
0
1
0
1
0
5
Araeoandra
0
0
0
0
0
0
0
0
0
1
0
1
0
6
Avellanita
0
0
0
0
0
0
0
0
0
0
0
0
0
7
Bakerolimon
0
1
0
1
0
1
0
0
0
1
0
0
0
8
Balsamocarpon
0
0
0
0
0
1
0
1
1
1
0
0
0
9
Bridgesia
0
0
0
0
0
0
0
1
0
1
0
1
1
10
Calopappus
0
0
0
0
0
0
0
0
0
0
0
0
0
11
Cissarobryon
0
0
0
0
0
0
0
0
0
0
0
0
0
12
Conanthera
0
1
0
0
0
1
0
1
0
1
0
1
0
13
Copiapoa
0
1
0
1
0
1
0
1
0
1
0
1
0
14
Cyphocarpus
0
0
0
0
0
1
0
1
0
1
0
1
1
15
Desmaria
0
0
0
0
0
0
0
0
0
0
0
0
0
16
Dinemagonum
0
0
0
0
0
1
0
1
0
1
1
1
1
17
Dinemandra
0
1
0
1
0
1
0
0
0
0
0
0
0
18
Epipetrum
0
1
0
0
0
0
0
0
0
0
0
1
0
19
Ercilla
0
0
0
0
0
0
0
0
0
0
0
0
0
20
Fascicularia
0
0
0
0
0
0
0
0
0
0
0
0
0
21
Francoa
0
0
0
0
0
0
0
0
0
0
0
0
0
22
Gethyum
0
0
0
0
0
0
0
0
0
1
0
1
0
23
Gomortega
0
0
0
0
0
0
0
0
0
0
0
0
0
24
Guynesomia
0
0
0
0
0
0
0
0
0
0
1
0
1
25
Gymnachne
0
0
0
0
0
0
0
0
0
0
0
0
0
26
Gypothamnium
0
1
0
1
0
0
0
0
0
0
0
0
0
27
Hollermayera
0
0
0
0
0
0
0
0
0
0
0
0
0
28
Homalocarpus
0
1
0
0
0
1
0
1
1
1
0
1
1
29
Huidobria
0
1
0
1
1
1
0
0
1
1
0
0
0
30
Hymenoglossum
0
0
0
0
0
0
0
0
0
0
0
0
0
31
Ivania
0
0
0
0
0
1
0
0
0
0
0
0
0
32
Jubaea
0
0
0
0
0
0
0
0
0
0
0
0
0
33
Lapageria
0
0
0
0
0
0
0
0
0
0
0
1
0
34
Latua
0
0
0
0
0
0
0
0
0
0
0
0
0
35
Legrandia
0
0
0
0
0
0
0
0
0
0
0
0
0
36
Leontochir
0
0
0
0
0
1
0
1
0
0
0
0
0
37
Leptocarpha
0
0
0
0
0
0
0
0
0
0
0
0
0
38
Leucocoryne
0
1
0
1
0
1
0
1
0
1
1
1
0
39
Leunisia
0
0
0
0
0
0
0
0
0
0
0
1
0
40
Marticorenia
0
0
0
0
0
0
0
0
0
0
0
0
0
41
Metharme
0
0
0
0
0
0
0
0
0
0
0
0
0
42
Microphyes
0
1
0
1
0
1
0
1
0
1
1
1
1
43
Miersia
0
0
0
0
0
0
0
0
0
0
0
0
0
44
Miqueliopuntia
0
0
0
0
0
1
0
1
0
1
0
0
0
45
Moscharia
0
0
0
0
0
0
0
0
0
1
0
1
0
46
Neuontobothrys
0
0
0
0
0
0
0
0
0
0
0
0
0
47
Notanthera
0
0
0
0
0
0
0
0
0
0
0
0
0
48
Ochagavia
0
0
0
0
0
0
0
0
0
0
0
0
0
49
Oxyphyllum
0
1
0
1
0
0
0
0
0
0
0
0
0
50
Peumus
0
0
0
0
0
0
0
0
0
0
0
1
0
51
Phrodus
0
0
0
0
1
1
0
1
1
1
1
1
1
52
Pintoa
0
0
0
0
0
1
0
0
1
0
0
0
0
53
Pitavia
0
0
0
0
0
0
0
0
0
0
0
0
0
54
Placea
0
0
0
0
0
0
0
0
0
0
0
0
0
55
Pleocarphus
0
0
0
0
0
0
0
0
0
1
0
1
0
56
Podanthus
0
0
0
0
0
0
0
0
0
0
0
1
0
57
Reicheella
0
0
0
0
0
0
0
0
0
0
0
0
0
58
Sarmienta
0
0
0
0
0
0
0
0
0
0
0
1
0
59
Scyphanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
60
Speea
0
0
0
0
0
0
0
0
0
0
0
0
0
61
Tecophilaea
0
0
0
0
0
0
0
0
0
0
0
1
0
62
Tetilla
0
0
0
0
0
0
0
0
0
0
0
0
0
63
Traubia
0
0
0
0
0
0
0
0
0
0
0
0
0
64
Trevoa
0
0
0
0
0
0
0
0
0
0
0
1
0
65
Valdivia
0
0
0
0
0
0
0
0
0
0
0
0
0
66
Vestia
0
0
0
0
0
0
0
0
0
0
0
0
0
67
Zephyra
0
1
0
0
0
1
0
1
0
1
0
0
0
TOT
0
14
0
9
2
18
0
15
5
20
5
24
7
257
N°
GENERO
O co
O an
P co
P an
Q co
Q an
R co
R an
S co
S an
T co
T an
U co
U an
V co
1
Acrisione
1
0
1
0
1
1
1
1
1
1
1
1
1
1
1
2
Adenopeltis
1
0
1
0
1
0
1
0
1
0
1
0
0
0
0
3
Alona
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
4
Anisomeria
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
5
Araeoandra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
Avellanita
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
7
Bakerolimon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
8
Balsamocarpon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9
Bridgesia
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
10
Calopappus
0
0
0
1
0
1
0
1
0
1
0
0
0
0
0
11
Cissarobryon
0
0
0
0
1
1
0
1
0
1
0
1
1
1
0
12
Conanthera
1
0
1
1
1
1
1
1
1
1
1
1
1
0
0
13
Copiapoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
14
Cyphocarpus
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
15
Desmaria
0
0
0
0
0
0
0
0
0
1
0
1
1
1
1
16
Dinemagonum
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
17
Dinemandra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
18
Epipetrum
1
0
1
0
1
0
1
1
1
0
0
0
0
1
0
19
Ercilla
0
0
0
0
1
1
1
1
1
1
1
0
1
0
1
20
Fascicularia
0
0
0
0
1
0
0
0
1
0
1
0
1
0
1
21
Francoa
0
0
1
0
1
0
1
0
1
1
1
1
1
1
1
22
Gethyum
1
0
0
0
0
1
0
1
0
0
0
0
0
0
0
23
Gomortega
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
24
Guynesomia
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
25
Gymnachne
0
0
0
0
1
0
1
0
1
0
1
0
1
0
1
26
Gypothamnium
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
Hollermayera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
28
Homalocarpus
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
29
Huidobria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
30
Hymenoglossum
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
31
Ivania
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
32
Jubaea
1
0
1
0
1
0
1
0
1
0
0
0
0
0
0
33
Lapageria
0
0
0
0
1
0
0
0
1
0
1
1
1
0
1
34
Latua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
35
Legrandia
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
36
Leontochir
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
37
Leptocarpha
0
0
0
0
0
0
0
0
1
0
1
1
1
1
1
38
Leucocoryne
1
1
1
1
1
1
1
0
1
1
1
0
1
0
0
39
Leunisia
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
40
Marticorenia
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
41
Metharme
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
42
Microphyes
1
1
1
0
1
1
0
0
0
1
1
0
1
1
0
43
Miersia
1
0
1
0
1
0
1
0
1
0
0
0
0
0
0
44
Miqueliopuntia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
45
Moscharia
1
0
1
0
1
1
1
0
1
0
1
0
0
0
0
46
Neuontobothrys
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
47
Notanthera
0
0
1
0
1
0
1
1
1
1
1
1
1
1
1
48
Ochagavia
1
0
1
0
1
0
1
1
1
1
1
1
1
1
0
49
Oxyphyllum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
50
Peumus
1
0
1
0
1
1
1
1
1
1
1
1
1
1
1
51
Phrodus
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
52
Pintoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
53
Pitavia
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
54
Placea
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
55
Pleocarphus
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
56
Podanthus
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
57
Reicheella
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
58
Sarmienta
0
0
0
0
0
0
0
0
1
0
1
0
1
0
1
59
Scyphanthus
1
1
1
1
0
1
0
1
1
1
1
1
0
1
0
60
Speea
0
0
1
0
1
1
0
1
0
0
0
0
0
0
0
61
Tecophilaea
1
0
1
0
1
1
0
0
0
0
0
0
0
0
0
62
Tetilla
1
0
1
0
1
1
1
1
1
0
0
0
0
0
0
63
Traubia
1
0
1
0
1
0
1
0
0
0
0
0
0
0
0
64
Trevoa
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
65
Valdivia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
66
Vestia
0
0
1
0
1
0
0
1
1
0
1
1
1
1
1
67
Zephyra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
TOT
26
12
28
12
31
20
20
18
26
17
22
14
21
13
14
258
N°
GENERO
V an
W co
W an
X co
X an
Y co
Y an
Z co
Z an
AA co
AA an
AB co
AB an
1
Acrisione
0
1
0
0
0
0
0
0
0
0
0
0
0
2
Adenopeltis
0
0
0
0
0
0
0
0
0
0
0
0
0
3
Alona
0
0
0
0
0
0
0
0
0
0
0
0
0
4
Anisomeria
0
0
0
0
0
0
0
0
0
0
0
0
0
5
Araeoandra
0
0
0
0
0
0
0
0
0
0
0
0
0
6
Avellanita
0
0
0
0
0
0
0
0
0
0
0
0
0
7
Bakerolimon
0
0
0
0
0
0
0
0
0
0
0
0
0
8
Balsamocarpon
0
0
0
0
0
0
0
0
0
0
0
0
0
9
Bridgesia
0
0
0
0
0
0
0
0
0
0
0
0
0
10
Calopappus
0
0
0
0
0
0
0
0
0
0
0
0
0
11
Cissarobryon
0
0
0
0
0
0
0
0
0
0
0
0
0
12
Conanthera
0
0
0
0
0
0
0
0
0
0
0
0
0
13
Copiapoa
0
0
0
0
0
0
0
0
0
0
0
0
0
14
Cyphocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
15
Desmaria
1
1
0
1
0
0
0
0
0
0
0
0
0
16
Dinemagonum
0
0
0
0
0
0
0
0
0
0
0
0
0
17
Dinemandra
0
0
0
0
0
0
0
0
0
0
0
0
0
18
Epipetrum
0
0
0
0
0
0
0
0
0
0
0
0
0
19
Ercilla
1
1
0
1
1
1
1
1
1
0
0
0
1
20
Fascicularia
0
1
0
1
1
0
1
1
0
0
0
0
0
21
Francoa
1
1
1
1
1
1
1
1
0
1
0
0
0
22
Gethyum
0
0
0
0
0
0
0
0
0
0
0
0
0
23
Gomortega
0
0
0
0
0
0
0
0
0
0
0
0
0
24
Guynesomia
0
0
0
0
0
0
0
0
0
0
0
0
0
25
Gymnachne
0
1
0
0
0
0
0
0
0
0
0
0
0
26
Gypothamnium
0
0
0
0
0
0
0
0
0
0
0
0
0
27
Hollermayera
1
0
0
0
1
0
0
0
0
0
0
0
0
28
Homalocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
29
Huidobria
0
0
0
0
0
0
0
0
0
0
0
0
0
30
Hymenoglossum
0
1
1
1
1
1
1
1
1
1
1
1
0
31
Ivania
0
0
0
0
0
0
0
0
0
0
0
0
0
32
Jubaea
0
0
0
0
0
0
0
0
0
0
0
0
0
33
Lapageria
1
1
0
1
1
0
0
0
0
0
0
0
0
34
Latua
0
0
0
1
0
1
0
1
0
0
0
0
0
35
Legrandia
0
0
0
0
0
0
0
0
0
0
0
0
0
36
Leontochir
0
0
0
0
0
0
0
0
0
0
0
0
0
37
Leptocarpha
0
1
1
1
1
0
0
0
0
0
0
0
0
38
Leucocoryne
0
1
0
0
0
0
0
0
0
0
0
0
0
39
Leunisia
0
0
0
0
0
0
0
0
0
0
0
0
0
40
Marticorenia
0
0
0
0
0
0
0
0
0
0
0
0
0
41
Metharme
0
0
0
0
0
0
0
0
0
0
0
0
0
42
Microphyes
0
0
0
0
0
0
0
0
0
0
0
0
0
43
Miersia
0
0
1
0
0
0
0
0
0
0
0
0
0
44
Miqueliopuntia
0
0
0
0
0
0
0
0
0
0
0
0
0
45
Moscharia
0
0
0
0
0
0
0
0
0
0
0
0
0
46
Neuontobothrys
0
0
0
0
0
0
0
0
0
0
0
0
0
47
Notanthera
1
1
0
0
1
0
0
1
0
0
0
0
0
48
Ochagavia
1
0
0
0
0
0
0
0
0
0
0
0
0
49
Oxyphyllum
0
0
0
0
0
0
0
0
0
0
0
0
0
50
Peumus
0
1
0
1
1
0
0
0
0
0
0
0
0
51
Phrodus
0
0
0
0
0
0
0
0
0
0
0
0
0
52
Pintoa
0
0
0
0
0
0
0
0
0
0
0
0
0
53
Pitavia
0
0
0
0
0
0
0
0
0
0
0
0
0
54
Placea
0
0
0
0
0
0
0
0
0
0
0
0
0
55
Pleocarphus
0
0
0
0
0
0
0
0
0
0
0
0
0
56
Podanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
57
Reicheella
0
0
0
0
0
0
0
0
0
0
0
0
0
58
Sarmienta
0
1
0
1
1
1
0
1
0
1
0
0
0
59
Scyphanthus
1
0
0
0
0
0
0
0
0
0
0
0
0
60
Speea
0
0
0
0
0
0
0
0
0
0
0
0
0
61
Tecophilaea
0
0
0
0
0
0
0
0
0
0
0
0
0
62
Tetilla
0
0
0
0
0
0
0
0
0
0
0
0
0
63
Traubia
0
0
0
0
0
0
0
0
0
0
0
0
0
64
Trevoa
0
0
0
0
0
0
0
0
0
0
0
0
0
65
Valdivia
0
1
0
1
0
0
0
0
0
0
0
0
0
66
Vestia
1
1
0
0
0
0
0
1
0
0
0
0
0
67
Zephyra
0
0
0
0
0
0
0
0
0
0
0
0
0
TOT
9
15
4
11
10
5
4
8
2
3
1
1
1
259
N°
GENERO
AC co
AC an
AD co
AD in
AD an
AE co
AE an
AF co
AF an
AG co
AG an
AH co
AH an
AI co
AI an
1
Acrisione
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
Adenopeltis
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
Alona
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
Anisomeria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5
Araeoandra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
Avellanita
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
Bakerolimon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
8
Balsamocarpon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9
Bridgesia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
10
Calopappus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
11
Cissarobryon
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
12
Conanthera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
13
Copiapoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
14
Cyphocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
15
Desmaria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
16
Dinemagonum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
17
Dinemandra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
18
Epipetrum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
19
Ercilla
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
20
Fascicularia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
21
Francoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
22
Gethyum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
23
Gomortega
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
24
Guynesomia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
25
Gymnachne
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
26
Gypothamnium
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
Hollermayera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
28
Homalocarpus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
29
Huidobria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
30
Hymenoglossum
1
0
1
1
1
0
0
0
0
1
0
0
0
0
0
31
Ivania
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
32
Jubaea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
33
Lapageria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
34
Latua
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
35
Legrandia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
36
Leontochir
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
37
Leptocarpha
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
38
Leucocoryne
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
39
Leunisia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
40
Marticorenia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
41
Metharme
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
42
Microphyes
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
43
Miersia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
44
Miqueliopuntia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
45
Moscharia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
46
Neuontobothrys
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
47
Notanthera
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
48
Ochagavia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
49
Oxyphyllum
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
50
Peumus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
51
Phrodus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
52
Pintoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
53
Pitavia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
54
Placea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
55
Pleocarphus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
56
Podanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
57
Reicheella
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
58
Sarmienta
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
59
Scyphanthus
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
60
Speea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
61
Tecophilaea
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
62
Tetilla
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
63
Traubia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
64
Trevoa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
65
Valdivia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
66
Vestia
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
67
Zephyra
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
TOT
1
1
1
1
1
0
0
0
0
1
0
0
0
0
0
260
N°
GENERO
AJ co
AJ in
AJ an
AK co
AK in
AK an
AL co
AL an
AM co
AM an
SUM
1
Acrisione
0
0
0
0
0
0
0
0
0
0
14
2
Adenopeltis
0
0
0
0
0
0
0
0
0
0
7
3
Alona
0
0
0
0
0
0
0
0
0
0
10
4
Anisomeria
0
0
0
0
0
0
0
0
0
0
15
5
Araeoandra
0
0
0
0
0
0
0
0
0
0
2
6
Avellanita
0
0
0
0
0
0
0
0
0
0
2
7
Bakerolimon
0
0
0
0
0
0
0
0
0
0
6
8
Balsamocarpon
0
0
0
0
0
0
0
0
0
0
4
9
Bridgesia
0
0
0
0
0
0
0
0
0
0
9
10
Calopappus
0
0
0
0
0
0
0
0
0
0
4
11
Cissarobryon
0
0
0
0
0
0
0
0
0
0
7
12
Conanthera
0
0
0
0
0
0
0
0
0
0
19
13
Copiapoa
0
0
0
0
0
0
0
0
0
0
10
14
Cyphocarpus
0
0
0
0
0
0
0
0
0
0
7
15
Desmaria
0
0
0
0
0
0
0
0
0
0
8
16
Dinemagonum
0
0
0
0
0
0
0
0
0
0
7
17
Dinemandra
0
0
0
0
0
0
0
0
0
0
9
18
Epipetrum
0
0
0
0
0
0
0
0
0
0
9
19
Ercilla
0
0
0
0
0
0
0
0
0
0
19
20
Fascicularia
0
0
0
0
0
0
0
0
0
0
10
21
Francoa
0
0
0
0
0
0
0
0
0
0
19
22
Gethyum
0
0
0
0
0
0
0
0
0
0
5
23
Gomortega
0
0
0
0
0
0
0
0
0
0
3
24
Guynesomia
0
0
0
0
0
0
0
0
0
0
3
25
Gymnachne
0
0
0
0
0
0
0
0
0
0
7
26
Gypothamnium
0
0
0
0
0
0
0
0
0
0
2
27
Hollermayera
0
0
0
0
0
0
0
0
0
0
2
28
Homalocarpus
0
0
0
0
0
0
0
0
0
0
13
29
Huidobria
0
0
0
0
0
0
0
0
0
0
14
30
Hymenoglossum
0
0
0
0
0
0
0
0
0
0
18
31
Ivania
0
0
0
0
0
0
0
0
0
0
1
32
Jubaea
0
0
0
0
0
0
0
0
0
0
5
33
Lapageria
0
0
0
0
0
0
0
0
0
0
11
34
Latua
0
0
0
0
0
0
0
0
0
0
3
35
Legrandia
0
0
0
0
0
0
0
0
0
0
2
36
Leontochir
0
0
0
0
0
0
0
0
0
0
2
37
Leptocarpha
0
0
0
0
0
0
0
0
0
0
10
38
Leucocoryne
0
0
0
0
0
0
0
0
0
0
22
39
Leunisia
0
0
0
0
0
0
0
0
0
0
3
40
Marticorenia
0
0
0
0
0
0
0
0
0
0
2
41
Metharme
0
0
0
0
0
0
0
0
0
0
2
42
Microphyes
0
0
0
0
0
0
0
0
0
0
18
43
Miersia
0
0
0
0
0
0
0
0
0
0
6
44
Miqueliopuntia
0
0
0
0
0
0
0
0
0
0
3
45
Moscharia
0
0
0
0
0
0
0
0
0
0
9
46
Neuontobothrys
0
0
0
0
0
0
0
0
0
0
1
47
Notanthera
0
0
0
0
0
0
0
0
0
0
15
48
Ochagavia
0
0
0
0
0
0
0
0
0
0
12
49
Oxyphyllum
0
0
0
0
0
0
0
0
0
0
3
50
Peumus
0
0
0
0
0
0
0
0
0
0
17
51
Phrodus
0
0
0
0
0
0
0
0
0
0
10
52
Pintoa
0
0
0
0
0
0
0
0
0
0
2
53
Pitavia
0
0
0
0
0
0
0
0
0
0
3
54
Placea
0
0
0
0
0
0
0
0
0
0
7
55
Pleocarphus
0
0
0
0
0
0
0
0
0
0
5
56
Podanthus
0
0
0
0
0
0
0
0
0
0
15
57
Reicheella
0
0
0
0
0
0
0
0
0
0
2
58
Sarmienta
0
0
0
0
0
0
0
0
0
0
11
59
Scyphanthus
0
0
0
0
0
0
0
0
0
0
12
60
Speea
0
0
0
0
0
0
0
0
0
0
4
61
Tecophilaea
0
0
0
0
0
0
0
0
0
0
5
62
Tetilla
0
0
0
0
0
0
0
0
0
0
7
63
Traubia
0
0
0
0
0
0
0
0
0
0
4
64
Trevoa
0
0
0
0
0
0
0
0
0
0
12
65
Valdivia
0
0
0
0
0
0
0
0
0
0
2
66
Vestia
0
0
0
0
0
0
0
0
0
0
12
67
Zephyra
0
0
0
0
0
0
0
0
0
0
6
TOT
0
0
0
0
0
0
0
0
0
0
261
262
Appendix D: Native Genera in the Chilean Pacific Islands
263
Islas Desventuradas
Isla de Pascua
Juan Fernández
Juan Fernández (cont.)
Atriplex
Agrostis
Abrotanella
Leptophyllochloa
Chenopodium
Apium
Acaena
Libertia
Cristaria
Asplenium
Adiantum
Lobelia
Eragrostis
Axonopus
Agrostis
Lophosoria
Frankenia
Davallia
Apium
Luzula
Fuertesimalva
Diplazium
Arthropteris
Lycopodium
Lepidium
Doodia
Asplenium
Machaerina
Lycapsus
Dryopteris
Azara
Margyricarpus
Maireana
Elaphoglossum
Berberis
Megalachne
Nesocaryum
Ipomoea
Blechnum
Megalastrum
Parietaria
Juncus
Boehmeria
Mimulus
Plantago
Kyllinga
Bromus
Myrceugenia
Sanctambrosia
Lycium
Calystegia
Myrteola
Sicyos
Microlepia
Cardamine
Nassella
Solanum
Microsorum
Carex
Nertera
Spergularia
Ophioglossum
Centaurodendron
Nicotiana
Suaeda
Paspalum
Centella
Notanthera
Tetragonia
Polystichum
Chenopodium
Notholaena
Thamnoseris
Psilotum
Chusquea
Ochagavia
Pycreus
Colletia
Ophioglossum
Rytidosperma
Coprosma
Oreobolus
Samolus
Cuminia
Parietaria
Scirpus
Cyperus
Peperomia
Solanum
Cystopteris
Piptochaetium
Sophora
Danthonia
Plantago
Stipa
Dendroseris
Pleopeltis
Thelypteris
Dichondra
Podophorus
Triumfetta
Dicksonia
Polypodium
Vittaria
Drimys
Polypogon
Dysopsis
Polystichum
Elaphoglossum
Pteris
Eleocharis
Ranunculus
Empetrum
Rhaphithamnus
Erigeron
Robinsonia
Eryngium
Rubus
Escallonia
Rumohra
Euphrasia
Santalum
Galium
Sarcocornia
Gamochaeta
Scirpus
Gaultheria
Selkirkia
Gavilea
Serpyllopsis
Gleichenia
Solanum
Grammitis
Sophora
Greigia
Spergularia
Gunnera
Taraxacum
Haloragis
Thyrsopteris
Hedyotis
Trichomanes
Histiopteris
Trisetum
Hymenoglossum
Ugni
Hymenophyllum
Uncinia
Hypolepis
Urtica
Juania
Wahlenbergia
Juncus
xMargyracaena
Lactoris
Yunquea
Lagenophora
Zanthoxylum
264
Agradecimientos / Danksagung / Aknowledgements
Quiero, en primer lugar, dar las gracias a Paola Soto, mi mujer; sin ella ni este trabajo ni nada
hubiese sido posible. A ella se lo dedico. Y a mi pequeña flor, mi pequeño roble, y mi pequeño amor.
Agradezco también a Mélica Muñoz, no sólo por ser mi madre que ya es mucho, sino por compartir
siempre sus inagotables conocimientos botánicos y por haber sido desde que tengo memoria el
pilar fundamental de mi formación naturalista. A través de ella no puedo dejar de mencionar
a mi abuelo Carlos Muñoz y su esposa Ruth, quienes aún nos iluminan en los momentos de
desconcierto. A través de ellos he tenido la suerte de conocer a Calvin y Linda Heusser, quienes
me iniciaron en los intrincados caminos de la ciencia.
Vaya mi gratitud a mi padre Sergio, no sólo por ser mi padre sino por haberme impulsado siempre
a viajar en busca de nuevos horizontes. Mis hermanos han colaborado cada uno a su manera: con
Tiarella compartimos el nervio de una tesis, Iván ha enviando imprescindible música y videos;
Simón ha descuidando su propia tesis para conseguir literatura ‚indispensable‘.
Federico Luebert ha revisado la tesis en forma exhaustiva y crítica, y ha sido junto con Patricio
Pliscoff siempre partícipe de una discusión enriquecedora. Espero contar con su amistad por
muchos años (viva Berlín!) También Sergio Elórtegui ha sido un gran compañero en la distancia,
mágico creador de ilustraciones e ilusiones.
Agradezco también la amistad y la compañía musical de Carlos Ledermann, y el duende de los
Mártires del Compás; sin él hubiese perdío el rumbo hace tiempo.
Meinem Doktorvater, Herrn Prof. Dr. Michael Richter, danke ich für die engagierte Betreuung
der Arbeit am Anfang und dafür, mir am Ende die notwendigen Freiräume gewährt zu haben.
Frau Prof. Dr. Perdita Pohle und Herrn Prof. Dr. Achim Bräuning danke ich herzlich für die
Bereitschaft, bei meinem Rigorosum den Prüfungsvorsitz übernommen bzw. mich geprüft zu
haben.
Dank dafür gebührt natürlich auch Herrn Prof. Dr. Werner Nezadal, meinem Prüfer im Nebenfach;
ihm verdanke ich auch, viele Pflanzen am Cabo de Gata (Spanien) kennengelernt zu haben.
Walter Welß hieß mich in der Mitte seiner umfangreichen Bibliothek im Botanischen Garten
willkommen; langsam sind viele Bücher auch zu mir gewandert... Er ist in Teilen verantwortlich
für die Ergebnisse meiner Arbeit (zumindest die guten). Herrn Prof. Dr. Tod Stuessy (Wien) danke
ich für die Übernahme des Zweitgutachtens.
Lieben Dank an Sabine Donner für die stetige Hilfe in allen bürokratischen Fragen und danke allen
Mitarbeitern des Instituts für Geographie für die freundliche Atmosphäre und die Möglichkeit
zum fachlichen Austausch!
265
Besonders danke ich dabei meinen Mit-Doktoranden (viele sind mittlerweile schon promoviert):
Luisa Vogt, Hendrik Wagenseil, Natalie Schulz, Cristina Dall‘ Ozzo, Markus Pingold, Michaela
Ise, Paul Emck, Daniel Lingenhöhl, Henning Schröder, Tobias Bolch, Frieda Grüninger und Tom
Fickert. Klaus Geiselhart verdient einen speziellen Dank für seine grundlegende Erfindung: den
Geofanten.
Auch Alex Brenning muss ich hier erwähnen, obwohl ich ihn nicht leiden kann (…nur ein Scherz!).
Und von einer erlebnisreichen Exkursion nach Tunesien bleibt mir die gute Freundschaft mit
den Studierenden Viktor Kollmannsberger, Eva Neubauer, Matthias Patrzek und vor allem Ales
Macik erhalten. Es war ein Glück Euch kennenzulernen!
Die sehr hilfreiche Englisch-Korrektur der Dissertation übernahm Iris Burchardt. Nochmals
vielen Dank! Marian Jüngling gehört zwar nicht zur Geographie, doch war seine sprachliche
Hilfe entscheidend für den guten Verlauf des Rigorosums.
Speziellen Dank an Thomas Sokoliuk für seine langjährige Unterstützung und seine unerschöpfliche
Feststimmung (danke für den Fisch, Thomas!) Bei Rolf Kastner möchte ich mich unter anderem
für die Überlassung seines Computers bedanken und bei Stefan Adler für die große Hilfe beim
Einleben in Deutschland im ersten Jahr. Irma Richter, Anette Welß, Arietta Eberstadt sowie Coni
Augustin sorgten auch für das gute Eingewöhnen und stets für das seelische und leibliche Wohl.
Bei Frau Dr. Brigitte Perlick, Akademisches Auslandsamt der Universität, möchte ich mich für
ihre unendliche Energie bedanken und bei Frau Merker vom Promotionsbüro für den besten Rat
in einer schwierigen Situation.
Schließlich danke ich dem DAAD für die gewährte finanzielle Unterstützung. Besonders der
zuständigen Referentin Maria Hartmann gebührt für die schnelle Lösung unlösbarer Probleme
mein herzlicher Dank. Der Zantner-Busch Stiftung gilt auch mein Dank für die finanzielle
Unterstützung des Besuchs verschiedener Kongresse und Seminare.
Malte Ebach, Michael Heads, and Dennis McCarthy contributed with the review and edition of
several papers. I hope our friendship grows and never suffers vicariance.
S. Liede, S. Renner, P. Endress, V. Funck, M. Gandolfo, M. Griffith, and J.J. Morrone collaborated
providing important papers. Of course I’m alone responsible for the interpretation of these
papers.
Am Schluss möchte ich den Erlangern danken, die Stadt bot uns fast vier Jahre lang ein Zuhause.
Und einen Ratschlag möchte ich zu guter Letzt geben: Nehmt alles nicht so ernst! Das Leben ist
zu kurz dafür! Ich wünsche mir, Deutschland hätte die WM 2006 doch gewonnen…
Die gute Stimmung hätte vielleicht ein paar Jahre angedauert.
266
LEBENSLAUF
Persönliche Daten
Name: Andrés Moreira-Muñoz
Eltern:
Sergio Moreira / Mélica Muñoz-Schick
Geburtsdatum: 25. Feb. 1971 in Los Angeles, Chile
Familienstand: verheiratet mit Paola Soto aus Viña del Mar, Chile
Kinder:
Sayén (4), Silene (2), Coyán (1/4)
Schule
1976 - 1988 Abschluß: Deutsche Schule, Santiago de Chile
Prueba de Aptitud Académica (entspricht Abitur)
Studium
1989 - 1994
Studium der Geographie an der Universidad Católica de Chile
Abschluß:
Diplom-Geograph. Thema der Diplomarbeit: „Naturschutzgebiete an
der Küste Zentral Chiles“
1994 - 1995 Studium der Angewandte Geographie an der Universidad Católica de Chile
Abschluß:
Geógrafo Profesional. Thema der Abschlußarbeit:: „Umwelt-Erziehung
im Natur-Reservat Yerba Loca, Andes Zentral Chile“
Promotion
2003 - 2007
Institut für Geographie, Friedrich Alexander Universität Erlangen - Nürnberg
Thema: Plant Geography of Chile
Andrés Moreira-Muñoz
Erlangen, 14. Juni 2007
267

Plant Geography of Chile

Transcription

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