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Biogeographic Map of South America. A


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International Journal of Geobotanical Research, Vol. nº 1, December 2011, pp. 21-40 + Map

Biogeographic Map of South America. A preliminary survey.

Salvador RIVAS-MARTÍNEZ a, Gonzalo NAVARRO b, Angel PENAS c and Manuel COSTAd

a
Phytosociological Research Center, Collado-Villalba, Madrid, Spain. E-mail: rivas-martinez.cif@tsai.es
b
Catholic University "San Pablo". Cochabamba. Bolivia. E-mail: gonzalonavarrosanchez@gmail.com
c
Department of. Biodiversity & Environmental Management (Botany).Faculty of. Biology & Environmental Sciences. Mountain
Livestock Institute (CSIC-University of Leon). University of Leon. Spain. E-mail: a@unileon.es.
d
Botanical Garden of Valencia. Spain. E-mail: manuel.costa@uv.es

With the collaboration of: Javier Amigo (Chile and Argentina), Alindo Butzke (Brazil), Sara del Río (Argentina, Brazil and Para-
guay), Antonio Galán (Perú), José Guevara (Venezuela), Jesús Izco (Ecuador), Eduardo Martínez Carretero (Argentina), Orlando
Rangel (Colombia), Salvador Rivas Sáenz (South America bioclimate expert), Fidel Roig (†) (Argentina), Daniel Sánchez-Mata
(South Chile), Leopoldo G. Sancho (South Chile and Antarctica), Pilar Soriano (Venezuela) and Oscar Tovar (†) (Perú).

Abstract

The biogeographic map of South America showing the biogeographic units up to the provincial level is published in this paper as a
preliminary survey. From Honduras (Central America) to the Antarctic Peninsula we recognize: 1 kingdom, 3 subkingdoms, 4 supe-
regions, 13 regions and 53 biogeographic provinces. A brief description is given of each one of the 13 regions, indicatng its limits
and main bioclimatic factors. The biogeographic map is accompanied on a smaller scale by a bioclimate map, and another subking-
dom map and a physical geography map.

Keywords: Biogeography, Map, South Americ

Introduction one single type of potential vegetation (climax) and as


a result, a single sequence of substitute communities.
Biogeography is the science which studies the dis-
The tesela and the permatesela (conceived within
tribution of species and biocoenosis on earth, as well as
the framework of dynamic-catenal phytosociology lo-
the relationships between them and their causes. It ta-
cated in exceptional sites: polar, fluvial, lake and mari-
kes into account the areas of taxa and syntaxa (choro-
ne lands-capes, deserts, high-mountain summits, dunes
logy), in addition to information from other nature
and rock formations, coastal cliffs, etc. in which the
sciences (geography, soil science, bioclimatology, geo-
permanent vegetation growing in these elementary spa-
logy, etc.), and attempts to establish a hierarchical ty-
ces lacks perennial substitution units) are the only bio-
pology of the territories on the planet, whose main
geographical units which can be repeated in a disjunc-
units in decreasing rank are as follows: kingdom, re-
tive manner.
gion, province, sector, district, area, landscape cell and
tesela (RIVAS-MARTÍNEZ & AL., 2007, 2011). Biogeo- The landscape cells, such as peni-plains in horst,
graphy has been twinned with phytogeography due to river valleys, lake systems, marshes, mountain sum-
the value of plant species and communities in its defi- mits, etc., are constituted in a broad geographic space
nition and de-limittation. characterized by a series of teselas or permateselas and
their corresponding complexes, assembled by networks
of geosygmeta and geopermasygmeta based on the re-
Concepts on biogeographical units
lief or on the soils in the territory.
The elementary biogeographic terrestrial unit or the The biogeographical country must be an extensive
unit of the lowest rank is the tesela, which we define as and clearly delimited territory which possesses an a-
a geographic space with a greater or lesser extension, bundant group of landscape cells, species, associations,
ecologically homogeneous –that is to say, it has only and above all, peculiar topographical geosygmeta.

_______________________________________________________________________________
Correspondence: Ängel Penas. Department of. Biodiversity & Environmental Management (Botany).Faculty of. Biology &
Environmental Sciences. Mountain Livestock Institute (CSIC-University of Leon). University of Leon. Spain. E-mail:
apenm@unileon.es.
ISSN: 2253-6302 print/ISSN: 2253-6515 on line ©Editaefa
DOI: 10.5616/ijgr110002
22 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

The district is a series of biogeographical countries, entity is the distinguishing and mapping of taxa (fami-
characterized by the existence of numerous differential lies, genera, species and subspecies) whose territorial
species and even endemic taxa in the coastal, oreadic distribution is closely restricted to one particular area.
and interior halophilous zones, which permit their se- These taxa are termed endemisms. Endemisms have
paration from the adjacent taxa; it also comprises an in- been successfully used to define and delimit the choro-
dependent unit through its associations, series, geose- logical or biogeographical units (provinces, sectors), as
ries and geoclinoseries which are absent in nearby dis- they form part of the phytogeographical subelement
tricts. which characterises them. Moreover, the endemisms
The sector is a grouping of districts with a large- (taxa or syntaxa) which have a greater area or are re-
scale geographic entity, which possesses its own ende- gional, and those which for migratory reasons are dis-
mic taxa, associations and vegetation series, as well as persed across various biogeographical regions, cons-
original topographical and geoclinosequential geoseries titute the phytogeographical element or geoelement.
which are generally due to the existence of exclusive
climatophilous, permanent and subserial communities, Concepts on vegetation series and lanscape phyto-
as well as to paleoclimates or former migratory routes. sociology
The province is a vast geographic territory which,
Nowadays, the development of dynamic-catenal
as well as possessing a large number of endemisms and
phytosociology and the syntaxonomic knowledge of
differential species (its own subelement) has particular
broad territories on the Earth, as well as the cartogra-
macroseries and geomacroseries; a particular altitudinal
phic delimitation of vegetation series, geoseries and
zonation in the vegetation is also characteristic of each
geopermaseries, when available, have become the es-
biogeographical province (exclusive geoclinoseries).
sential criteria for defining biogeographical units, in
The region is a very extensive territory, formed by a
addition to suitably compiled and mapped bioclimatic
group of biogeographical provinces which has a flora
and soil factors.
or regional floristic element with species, genera or
The vegetation series, also termed sygmetum, ex-
even endemic families; in addition it has its own parti-
presses the whole set of plant communities or stages
cular megaseries, geomegaseries and geomegapermase-
which can be found within similar teselar spaces as a
ries and in consequence, its own bioclimatic belts (RI-
result of the succession process, and includes both the
VAS-MARTÍNEZ, 2005).
representative association of the mature stage or series
Finally, the kingdom is the supreme unit of biogeo-
head, which is used as a nomenclatural reference, and
graphy, generally pluricontinental and multinsular,
the initial or subserial associations that may replace it.
which in addition to taxonomic and ecosystematic con-
Based on this concept, the vegetation series or sygme-
siderations, addresses the origins of the flora and fauna,
tum represents the basic unit or essential model of dy-
the formation of the great continents, orogenies and cli-
namic phytosociology. Distinctions can also be made
mate in the present and past.
between climatophilic, edaphoxerophilic, temporihy-
As is by now traditional in this science, the deno-
grophilic and edaphohygrophilic series. Climatophilic
minations of the biogeographic units –both the primary
or zonal series are located on mature soils according to
and the auxiliary units (from the subregion to the
the mesoclimate, and only receive rainwater: mesophy-
area)– are given as names and place names based on
tic, submesophytic and subxerophytic; the temporihy-
known geographical, orographic and historical designa-
grophilic series, included among the climatophilic, are
tions which are considered to conicide with, inform or
those which have additional water contribution due to
represent the area they are intended to represent.
their topographical circumstances, and they thus deve-
Orthographically, all the units are considered to be pro-
lop on flooded or very wet soils throughout part of the
per names identifying the place; the name is formed by
year, and –at least during the summer or dry period–
juxtaposing two geographical nouns, joined by means
the soil horizons are well-drained and aerated. Finally
of a hyphen, maintaining the initial capital in each due
the edaphoxerophilic series are found in particularly
to their condition of proper noun.
dry or xerophytic soils or biotopes such as lithosols,
It must be emphasized that the biogeographical
arenosols, very windy sites, steep slopes, crests, ledges,
units can only be accurately delimited through their
etc.; and the edaphohygrophilic series grow on particu-
diagnosis and through the corresponding maps. All the
larly wet soils and biotopes such as fluvisols, halosols,
territories –except for the tesela and permatesela– must
histosols, etc., and are found on river beds, marsh are-
be contiguous by land, lake or sea routes, and include
as, salt flats, peat bogs, etc.
all the orographic accidents and lithological diversity
The vegetation geoseries or geosygmetum is the ba-
which may exist in the area. Sometimes in the biogeo-
sic unit of dynamic-catenal phytosociology. It corres-
graphical territories as a whole there are introgressions
ponds to a catena of vegetation series which is found
by other adjacent territories, and these islands frequen-
around a given bioclimatic belt and biogeographic te-
tly occur in regions with a varied lithology or in areas
rritory in the heart of the universal crest-slope-valley
near regional or provincial boudaries. Their possible
model. This topographic framework makes it possible
typological independence, always of a lower rank than
to distinguish the three geomorphological aspects of
the area into which they introgress, depends on their
any complete catena where the vegetation series consti-
originality, floristic richness and phytocoenotics, as
tuting the geosygmetum are located in zones; the eda-
well as on their surface area.
phoxerophilic series and geoseries (hyperxerophilic
One of the criteria traditionally used for recognising
and xerophilic) are located in the driest sites (crests, es-
and delimiting biogeographic units with their own
Biogeographic Map of South America. A preliminary survey 23

carpments, lithosols, etc.); the climatophilic and tempo- and particularly what was then the Institute of Hispanic
rigrophilic series and geoseries are located on slopes Culture, sparingly but reliably financed the project in
and foothills where greater humidity is contributed by the early years (1977-1982), thus providing a stimulus
rainfall; and the edaphohygrophilic series and geoseries and a springboard for many. Rivas-Martínez has since
are found in the valleys and watercourses (fluvial, lake worked to share what he colloquially termed the
and watercourses), among which the river fractogeo- “American adventure” by promoting and organizing,
sygmetum is of great important to plant landscape sci- together with his good friend Manuel Costa, countless
ence due to its extrazonality, and also, in combination trips and studies in North America, Africa, China, Aus-
with the edaphoxerophilic and climatophilous sygmeta tralia and Tasmania. Moreover Manuel Costa has tra-
and geosygmeta, to the definition and structuring of re- veled independently and has an in-depth knowledge of
gional and global biogeography. Central America, Mexico, the east of North America
The vegetation geopermaseries, also known as geo- and Tibet. His last major project, still incomplete after
permasygmeta, is the catenal expression of a set of ne- a dozen years, is currently the geobotanical study of
ighboring permaseries or permasygmeta, delimited by Venezuela, where, with the help of a large group of
changing topographic or soil situations. These are con- first-rate young professional Venezuelan botanists and
ditioned by conditions of extreme climate (high moun- soil scientists, he is directing a series of doctoral theses
tains and polar areas) and exceptional microtopogra- on the ground, covering large parts of the territory, four
phic and soil conditions (walls, rock formations, mari- of which have already been judged “cum laude” at the
ne cliffs, salt flats, etc.) which give rise to a large num- University of Valencia. On several occasions accompa-
ber of neighboring ecological residences populated by nying Manuel Costa and his team, Rivas-Martínez has
diverse permanent perennial plant communities (con- had the opportunity to increase his knowledge of the
tinuous vegetation permaseries) with an absence of vegetation and the bioclimate of Venezuela and, above
non-nitrophilous serial perennial communities which all, to study the regions of Alto Orinoco, La Guayana
appear to have reached their equilibrium. The most fa- and the Gran Sabana.
vorable sites for the existence of geopermaseries or Gonzalo Navarro, Ph.D., a native of Madrid, now
geopermasygmeta, as well as sites corresponding to lives in Cochabamba, after completing his education
permanent types of vegetation in extreme high-moun- and writing an extensive doctoral thesis entitled “Flora
tain and polar region bioclimates, are ledges, rock cre- and vegetation of the Sistema Ibérico in Soria”, which
vasses, cliffs and coastal rock formations bathed by sea was directed by Rivas-Martínez. Shortly after, as a re-
waters, peat bogs, wind drifts, mobile sand dunes, lake sult of an exploratory trip through the Andes, Gonzalo
shores, streams etc. Navarro took the irrevocable decision to live and re-
search in Bolivia for most of the year. He thus applied
Studies performed and geobotanical sources for a leave of absence from his position as permanent
teacher in the institute of secondary education in Gua-
Salvador Rivas-Martínez and Manuel Costa, Pro-
dalajara (Spain), which he had won through public exa-
fessor of Botany at the University of Valencia, worked
mination. A little later he continued his connections as
together at the Faculty of Biology at the Complutense
a part-time associate professor in the Department of
University and at the Royal Botanical Gardens in Ma-
Botany at the Faculty of Pharmacy of the Complutense
drid in the 1970s, where they designed a long-term glo-
University, at that time headed by Rivas-Martínez, with
bal geobotanical program to further the syntaxonomical
the undertaking that during his winter stay in Madrid
study of the Earth following the European phytosocio-
he would teach the optional subject “Biogeography of
logical methodology of BRAUN-BLANQUET (1964). To
South America”, conduct research into Andean, Cha-
this method they incorporated the new synphytosocio-
coan and Amazonian ecosystems, and that at the Uni-
logical analyses proposed by RIVAS-MARTÍNEZ (1976),
versity of Cochabamba, with which there was an inter-
and shortly afterwards by GEHÚ & RIVAS-MARTÍNEZ
university agreement, he would train young students
(1982), as well as recent bioclimatic and biogeographi-
and graduates in geobotany so that under his direction
cal approaches which were being successfully tested in
they could begin studying their doctoral thesis on Boli-
Europe and North Africa, and in whose scientific and
vian subjects, and that thanks to the Mutis grant sys-
methodological debates with Tüxen, Gehú, Bolós, Qué-
tem, they could spend a long period of training in Ma-
zel and Tahktajan, they played an active role.
drid studying the subjects for their doctorate. Gonzalo
Primo Yúfera, at that time general secretary of the
Navarro amply and generously fulfilled this commit-
Higher Council for Scientific Investigation (CSIC) in
ment, which continued successfully maintained for al-
the mid-1970s, convened various directors of the Insti-
most a decade. Of the six Mutis grant-holders who we-
tutes in the CSIC, including Salvador Rivas-Martínez,
re in Madrid, two are already doctors. Unfortunately in
to present research proposals which could be used in
2004, after the requisite yearly democratic and secret
due course for the scientific commemoration of the
departmental vote for the renewal of his post as asso-
fifth centenary of the discovery of America in 1992.
ciate professor, he found himself to be one vote short,
They wasted no time in adapting part of their global
and was rejected. Fortunately we his friends still enjoy
geobotanical program to the wildest possible subject
the privilege of continuing to work and collaborate
they could imagine, in keeping with their dual facets of
with him.
botanists and climbers at that time: the “Phytosociolo-
In the last 20 years, Gonzalo Navarro has conduc-
gical and bioclimatological study of the Andes in the
ted a continuous, profound and methodical investiga-
Viceroyalties of Peru and Nueva Granada”. The CSIC,
tion into the Bolivian Andes, and the areas of Beni,
24 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

Chaco, Cerrado, Yungas and Amazonia, as well as in a expeditions and various botanical and professional ex-
more general manner in the United States, Mexico and peditions. However, it was between 1977 and 1990,
over almost all of South America from Patagonia to the owing to several well-organized projects financed by
Venezuelan Caribbean. On many expeditions and jour- Spain, when he was able uninterruptedly to develop the
neys he has used his own resources, and on others, re- “Phytosociological and bioclimatic study of the Andes
sources obtained in projects and competitions proposed of the Viceroyalties of Perú and Nueva Granada”,
by organizations such as TNC, WCS, WWF, etc. As a which from 1985 became known as “Bioclimatic and
result of this research, as well as multiple reports and cultivation belts of Perú” thanks to another better fi-
publications, we should also mention the unrivalled nanced project. During those 13 years he made more
“Vegetation of Bolivia” published in 2002, in the book than 20 botanical expeditions around a large part of the
“Ecological geography of Bolivia” [ISBN 99905-0- Andes, Amazonia, Chaco, the Pacific coasts and de-
225-0] and the astounding “Vegetation map of Bolivia serts, the Pampa, Patagonia and Valdivia, where he
at a scale of 1:250.000” published by Nature Conser- took 3,000 phytosociological inventories, compiled cli-
vancy [ISBN 978-99954-0-168-9]. For all these rea- mate data for more than 2,000 South American weather
sons he has achieved the distinction on his own merits stations, and mapped the vegetation and bioclimate of
of becoming the maximum geobotanical authority on most of the territories accessible by road. Manuel Cos-
Bolivia and a key reference in the neotropics of South ta, as well as many of his disciples, colleagues from the
America. Complutense University such as Carlos Arnaiz, Paloma
Ángel Penas, professor at the University of León, Cantó, Ana Crespo, Antonio Galán, Javier Loidi, Gon-
began his partnership with Rivas-Martínez 30 years zalo Navarro and Conchita Sáenz, also took part in so-
ago in an unforgettable geobotanical research project in me of these journeys. From the outset he was lucky
the Cordillera Cantábrica mountains, conducted with enough on almost all these campaigns to enlist the co-
Tomás E. Díaz, José Antonio F. Prieto and Javier Loi- llaboration and valuable help of Oscar Tovar, the emi-
di, and which culminated with the publication of a nent Peruvian agrostologist and botanist, Professor of
book on the vegetation in the Picos de Europa moun- Botany at the University of San Marcos in Lima, as
tains. Since then their professional relationship and fri- well as the support, teachings and opinions of Stephan
endship has continued to grow, particularly through the Beck, Ángel Cabrera, José Cuatrecasas, Ramón Ferrei-
publications in Itinera Geobotánica and the Habitats ra, Raúl Lara, Máximo Lieberman, Eduardo Martínez
projects. However their main point in common is their Carretero, Eric Oberdorfer, Salvador Rivas Goday, Fi-
mutual interest in the study of the vegetation of Ameri- del Roig and Paul Seibert. Unfortunately, two years
ca and Africa. In January 1996 they successfully inves- before the official conclusion of this last project, high-
tigated, in the company of Leonard Llorens, the south- level political differences between Spain and Peru, un-
eastern tropical and oreadic temperate vegetation of related to any personal considerations, led to the abrupt
South Africa; in America they undertook two major termination of the agreement and to the project’s finan-
geobotanical expeditions, one in September 1994, with cing. As it was not possible to redirect the South Ame-
Gonzalo Navarro and Francisco Amich, in which they rican research program to Spain, it had to be post-
toured the Pampa, Monte Argentino and Patagonia ma- poned. In view of these difficulties he took the decision
king inventories, concluding in the Mediterranean An- to spend two sabbatical years in the United States, in
dean forests of Austrocedrus chilensis and in the Valdi- the Missouri Botanical Garden, in order to devote him-
vian temperate forests of Nothofagus; the second was a self completely to the study of the vegetation of North
fruitful and extensive geobotanical transect in April America and finally to formulate the bioclimatology at
1995, in the company of Gonzalo Navarro, Federico the global level. Luck would have it that Gonzalo Na-
Fernández and Daniel Sánchez Mata, where they suc- varro, a young doctor with considerable training in ta-
cessfully studied the tropical south of the United Sta- xonomy and geobotany, had moved to Bolivia and with
tes: Texas, Tamaulipas, Chihuahua and Sonora, as well great enthusiasm and effort began to study in great
as the continental Mediterranean territories of the Great detail and with marked success the oreadic and flatlan-
Basin and the oceanic Mediterranean territories of Cali- dic vegetation of Bolivia. This circumstances led Rivas
fornia. However Ángel Penas most important research Martínez in 1994 to return to South America to work
activity in South America, apart from his geobotanical with Navarro, who had by now become a considerable
excursions to Colombia, Venezuela, Peru, Chile and expert and specialist, and together they designed the
Argentina, was in Brazil, where only between 1992 and biogeographical synthesis of South America (1994),
2006 he personally directed six doctoral theses on the which has today, after many years, been improved and
region of Parana, judged cum laude in the University of updated with the help and mutual responsibility of Án-
León, and still found time to travel extensively and stu- gel Penas and Manuel Costa, as well as with the colla-
dy numerous aspects of Amazonia, La Catinga, Cerra- boration and advice of other botanists with extensive
do Oriental, Mata Atlántica and the mesophytic Pampa. experience in the territory: J. Amigo, A. Butzke, S. del
Salvador Rivas-Martínez began his study of tropical Río, A. Galán, J. Guevara, J. Izco, E. Martínez Carrete-
and Andean flora and vegetation in 1961 on the occa- ro, O. Rangel, F. Roig (†), D. Sánchez-Mata, L.G. San-
sion of a major mountaineering expedition to the An- cho, P. Soriano and O. Tovar (†); as well as with the
des in Peru (Apolobamba massif and Cordillera Blan- experience of S. Rivas Sáenz, expert and co-author of
ca). Between 1965 and 1976 he continued these studies the program of computerized bioclimatic cartography.
intermittently in Colombia, Ecuador, Bolivia, Chile and At the same time a detailed review was made of the
Argentina, taking advantage of other mountaineering bibliography which has enabled us to compare our ob-
Biogeographic Map of South America. A preliminary survey 25

servations with those of other authors, of both general MERA, A. (2007), GALÁN DE MERA, A., GONZÁLEZ, A.,
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MACO, J. & REÁTEGUI, F. (2007), LABUNTSOVA, M.A. CLEEF, A.M. & SALAMANCA, S. (2005), RANGEL-CH.,
(1969), LAVRENKO, E. M. (1964), LLORENTE BOUS- J.O., GARAY-P., H. & AVELLA, A. (2010), RANGEL-
QUETS, J. & MORRONE, J. J. (EDS.) (2001), LUTEYN, J. CH., J.O., SÁNCHEZ, D. & ARIZA, N.C. (2005), RIVERA
(1999), MORELLO, J. (1958), MORRONE, J. J. (2000, DÍAZ, O. & FERNÁNDEZ-ALONSO, J. L. (2003), SALA-
2001A, 2001B), OZENDA, P. (1964), POSADAS, P. E.; ES- MANCA S., CLEEF, A.M. & RANGEL, J.O. (2003), SÁN-
TÉVEZ, J. M. & MORRONE, J. J. (1997), PRADO, D.E. &. CHEZ, R. & RANGEL, J.O. (1990) and WITTE, H.J.J.
GIBBS, P. E. (1993), RAVEN, P.H. (1963), RAVEN, P.H. (1995).
& AXELROD, D.J. (1974), RICARDI, M., GAVIRIA, J. & In Peru and Bolivia we should mention the works
ESTRADA, J. (2001), RIKLI, M. (1913, 1934), RIVAS- of DE LA BARRA, N. (2003), GALÁN DE MERA, A.
MARTÍNEZ, S. (1976, 1994, 1997, 2001, 2004, 2005), (1995, 1999), GALÁN DE MERA, A., BALDEÓN, S., BEL-
RIVAS-MARTÍNEZ, S & RIVAS-SAÉNZ (2009), RIVAS- TRÁN, H., BENAVENTE, M. & GÓMEZ, J. (2004), GALÁN
MARTÍNEZ, S. & COLS. (2007, 2011), RIVAS-MARTÍNEZ, DE MERA, A., CÁCERES, C. & GONZÁLEZ, A. (2003),
S. & NAVARRO, G. (1998), RIVAS-MARTÍNEZ, S., RI- GALÁN DE MERA, A., ROSA, M.V. & CÁCERES, C.
VAS-SÁENZ, S., PENAS, A., NAVARRO, G. & COSTA, M. (2002), GALÁN DE MERA, A. & VICENTE ORELLANA,
& COLS. (2011), RIVAS-MARTÍNEZ, S., SÁNCHEZ-MA- J.A (1996, 2006), GUTTE, P. (1980, 1986, 1988), MO-
TA, D. & COSTA, M. (1999), RIVAS-MARTÍNEZ, S. & LINA, J.A., NAVARRO, G., DE LA BARRA, N. &
TOVAR O. (1983), SCHMITHÜSEN, J. (1961), SCHWABE, LUMBRERAS, A. (2007).NAVARRO, G. (1993, 1997,
G. H. (1968), SMITH, A.C. & JOHNSTON, I.M. (1945), 2003), NAVARRO, G. & FERREIRA, W. (2007, 2009),
TAKHTAJAN, A. (1988), THORNE, R.F. (1963), DEIL, U., NAVARRO, G. & MALDONADO, M. (2002), NAVARRO,
ÁLVAREZ, M., BAUER, E.V. & RAMÍREZ, C. (2011), G., MOLINA, J. A. & DE LA BARRA, N. (2005), RA-
UDVARDY, M.D.F. (1975), WALTER, H. (1898) and MÍREZ, C. & BECK, S. (1981), RIVAS-MARTÍNEZ, S. &
WALTER, H. & STRAKA, H. (1970). TOVAR, O. (1982), SEIBERT, P. & MENHOFER, X. (1991,
We have also based this work on the studies carried 1992, 1993), WEBERBAUER, A. (1945) and WILLIAMS,
out in territories which –although extensive– are more L. (1945).
specific than the previous ones, by different authors In Ecuador the works by ACOSTA-SOLIS, M. (1966,
which we mention below, for their work done in vari- 1984), AGUIRRE, Z., COOMBES, L. & RAMSAY, P. M.
ous South American countries. From Venezuela and (2001), MADSEN, J.E., COTTON, E. & BALSLEV, H.
the Guayanas we should highlight the works of (EDS.) (2002), CAÑADAS, L. (1983), GREHAN, J. R.
ATAROFF, M. & SARMIENTO, L. (2003), AYMARD, G. 2001, IZCO, J., PULGAR, I., AGUIRRE, Z. & SANTÍN, F.
(2003), CASTROVIEJO, S. & LÓPEZ, G. (1985), COSTA, (2004), JOHNSTON, M.P. & P.H. RAVEN (1973), MOS-
M.; CEGARRA, J.; LUGO, L.; GUEVARA J.R.; CARRERO, COL OLIVERA, MARCELA C. & CLEEF, A. M. (2009),
O.; LOZADA, J. & SORIANO, P. (2008), COSTA, M.; CE- PÚLGAR, I., IZCO, J. & JADÁN, O. (2010), QUINTANILLA,
GARRA, A.; LUGO, L.; LOZADA, J.; GUEVARA, J. & SO- V.G. (1983), SIERRA, R. (ED.) (1999), SIERRA, R., CICE-
RIANO, P. (2007), CUELLO, N. & CLEEF. A.M. (2009 a, RÓN, C., PALACIOS, W. & VALENCIA, R. (1999), TERÁN,
b), DUNO DE ESTEFANO, R., AYMARD, G. & HUBER, O. E. (1979), TERNEUS, E. (2002) and ULLOA, C. & JØR-
(2007), ESTRADA SÁNCHEZ, J.C. (2003), GALÁN DE GENSEN, P.M. (1993).
26 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

In Chile the works carried out by AMIGO J. (2009), Results


AMIGO, J., IZCO, J. &.RODRÍGUEZ-GUITIÁN, M.A.
(2007), AMIGO J. & RAMÍREZ, C. (1998), AMIGO, J., A. Biogeogeographical typology of South America
RAMÍREZ, C. & GARCÍA QUINTANILLA, L. (2004, 2007), up to provincial level
AMIGO, J., SAN MARTÍN, J. & GARCÍA QUINTANILLA, L.
(2000), ETAYO, J. & SANCHO, L.G. (2008), HILDE- In South America and from Honduras to the Ant-
arctic Peninsula we recognise: 1 kingdom, 3 subking-
BRAND-VOGEL, R. (1984), HILDEBRAND-VOGEL, R.,
doms, 4 superegions, 13 regions and 53 biogeographic
GODOY, R. & VOGEL, A. (1990), LUEBERT, F. & GA-
provinces (see Anex Map).
JARDO, R. (2005), LUEBERT, F. & P PLISCOFF. (2006),
The biogeographical denominations, as is tradi-
MÉNDEZ, E. (2007), MÉNDEZ, E. & AMBROSETTI, A.J. tional, are place names and adjectives which express
(1985), NAVARRO, G. & RIVAS-MARTÍNEZ, S. (2005), territories, history or toponyms. Generally these deno-
OBERDORFER, E. (1960), PINTO, R. & LUEBERT, F. minations are given in Spanish and translated into
(2009), RAMÍREZ, C., SAN MARTÍN, C., CONTRERAS, D. English.
& SAN MARTÍN, J. (1994) and RUTHSATZ, B. (1995).
In Argentina, Paraguay, Uruguay and Brasil, the B. NEOTROPICAL-AUSTROAMERICAN Kingdom
works published by BOLÓS, O., CERVI, A.C. & HAT- [Reino NEOTROPICAL-AUSTROAMERICANO]
SCHBACH, G. (1991), BUTZKE, A. (1997), CABIDO, M.
Ba. NEOTROPICAL Subkingdom
(1985), CABIDO, M. & ACOSTA, A. (1986), CABIDO, M., [Subreino NEOTROPICAL]
Bab. CARIBBEAN-NEOGRANADIAN Superegion
ACOSTA, A. & DÍAZ, S. (1990), CABRERA, Á. L. (1971),
[Superregión CARIBEÑA-NEOGRANADINA]
CABRERA, Á. L. (1976), DIESEL, S. (2005), EITEN, G.
9. CARIBBEAN-MESOAMERICAN Region]
(1972, 1983), ESKUCHE, U. (1984, 2005), FAGGI, A.M.
[Región CARIBEÑA-MESOAMERICANA]
(1985), FIASCHI, P & J.R. PIRANI. (2009), FONTANA, 9.3. Lesser Antillean Province
S.L. (2005), GALÁN DE MERA, A. & NAVARRO G. [Provincia Antillana Menor]
(1992), GANDULLO, R. & FAGGI, A. M. (2003, 2005), 9.5. Chiapan-Honduran Province]
GANDULLO, R. & SCHMID, P. (2001), GIBBS, P. E., [Provincia Chiapaneca-Hondureña]
LEITAO FILHO, H. & SHEPHERD, G. (1983), KEGLER, A. 9.6. Panamanian-Costa Rican Province
(2005), KEGLER, A., DIESEL, S., WASUM, R.A., HERRE- [Provincia Panameña-Costarricense]
RO, L., DEL RÍO, S. & PENAS, A. (2010), LEÓN, R.J.C. & 10. NEOGRANADIAN Region
BURKART, S.E. (1988), LEWIS, J:P:, COLLANTES, M:B:, [Región NEOGRANADINA]
PIRE, E.F., CARNEVALE, N.J., BOCCANELLI, S.I., STOFE- 10.1. Guajiran-Caribbean Province
LLA, S.L. & PRADO, D.E. (1985), MARTÍNEZ CARRETE- [Provincia Guajireña-Caribeña]
RO, E. (1993, 1995, 2000, 2001, 2006), MÉNDEZ, E., E. 10.2. Llaneran Province
MARTÍNEZ CARRETERO & I. PERALTA, I. (2006), NAVA- [Provincia Llanera]
RRO, G., MOLINA, J. A. & PÉREZ DE MOLAS, L. (2006), 10.3. Colombian Andean Province
PRANCE, G. T. (1977, 1978, 1979), RATTER, J. A., LEI- [Provincia Andina Colombiana]
TAO-FILHO, H. DE F., ARGENT, G., GIBBS, P. E., SEMIR, 10.4. Cordobesa-Lower Magdalena Province
J., SHEPHERD, G. & TAMASHIRO, J. (1988), REITZ, P. R. [Provincia Cordobesa-Bajomagdalena]
(1961), RIZZINI, C. T. (1979), ROIG, F.A. (1972, 1998), 10.5. Colombian Pacific Province
ROIG, F.A., ANCHORENA, J., DOLLENZ, O., FAGGI, A.M. [Provincia Pacífico Colombiana]
& MÉNDEZ, E. (1985), ROIG, F.A., DOLLENZ, O., & 10.6. Guayaquilian-Ecuadorean Province
MÉNDEZ, E. (1985), ROIG, F.A.& FAGGI, A. (1985), [Provincia Guayaquileña-Ecuatoriana]
10.7. Insular Galapagos Province
RUTHSATZ, B. (1977), SAMPAIO, A. J. DE (1934), SCUR
[Provincia Islas Galápagos]
L. (2005), STUTZ DE ORTEGA, L. C. (1983, 1984, 1986,
Bac. AMAZONIAN-GUYANAN Superegion
1987, 1990), VELOSO H. P. (1946, 1948), VELOSO H. P.
[Superregión AMAZÓNICA-GUAYANENSE]
& KLEIN, R. M. (1957, 1959) and WASUM, R. A. 11. GUYANAN-ORINOQUIAN Region
(2005). [Región GUAYANA-ORINOQUENSE]
And finally we should indicate that we have ana- 11.1. Guyanan Province
lysed proposals for territories other than South America [Provincia Guayanense]
but which bear a relation with South America for flo- 11.2. Deltaic Orinoquian Province
ristic or other reasons, among which we should men- [Provincia Orinoquense Deltaica]
tion: CANO CARMONA , E., VELOZ, A. & CANO ORTÍZ, 11.3. Guaviarean-Orinoquian Province
A. (2010), CONTRERAS MEDINA., R.; LUNA VEGA, I. & [Provincia Guaviareña-Orinoquense]
MORRONE, J. J. (1999), EMMERICH, K. H. (1988), GA- 11.4. Tepuyan Province
LÁN DE MERA, A. (2005), LAUER, W. (1968), LUNA VE- [Provincia Tepuyana]
GA, I.; MORRONE, J. J.; ALCÁNTARA AYALA, O. & ESPI- 11.5. Guyanese Brazilian Province
NOSA ORGANISTA, D. (2001), MATTICK, F. (1964), [Provincia Brasileña Guayanense]
DEIL, U. (1994, 1999). We have also particularly taken 12. AMAZONIAN Region
into account the bioclimatic typology of RIVAS-MAR- [Región AMAZÓNICA]
TÍNEZ, S. (2004), RIVAS-MARTÍNEZ, S. & RIVAS SÁENZ, 12.1. West Amazonian Province
S. (2009) and RIVAS-MARTÍNEZ, S., RIVAS SÁENZ, S. & [Provincia Amazónica Occidental]
PENAS, A. (2011). 12.2. North Amazonian Province
Biogeographic Map of South America. A preliminary survey 27

[Provincia Amazónica Septentrional] 18. MIDDLE CHILEAN-PATAGONIAN Region


12.3. Deltaic Amazonian Province [Región MESOCHILENA-PATAGÓNICA]
[Provincia Amazónica Deltaica] 18.1. Desertic Mediterranean Chilean Province
12.4. Southwest Amazonian Province [Provincia Chilena Mediterránea Desértica]
[Provincia Amazónica Suroccidental] 18.2. Central Chilean Province
12.5. Central Amazonian Province [Provincia Chilena Central]
[Provincia Amazónica Central] 18.3. Mediterranean Andean Province
Bad. CHACOAN-BRAZILIAN Superegion [Provincia Andina Mediterránea]
[Superregión CHAQUEÑA-BRASILEÑA] 18.4. Argentine Monte Province
13. BRAZILIAN-PARANENSE Region [Provincia Monte Argentino]
[Región BRASILEÑA-PARANAENSE] 18.5. North Patagonian Province
13.1. Brazilian Atlantic Province [Provincia Patagónica Septentrional]
[Provincia Atlántica Brasileña] 18.6. South Patagonian Province
13.2. Paranense Province [Provincia Patagónica Meridional]
[Provincia Paranaense] 19. VALDIVEAN-MAGELLANIAN Region
13.3. Catingan Province [Región VALDIVIANA-MAGALLÁNICA]
[Provincia Catinguense] 19.1. Valdivean Province
13.4. Tocantins Province [Provincia Valdiviana]
[Provincia Tocantinense] 19.2. Temperate Magellanian Province
13.5. East Cerrado Province [Provincia Magallánica Templada]
[Provincia Cerradense Oriental] 19.3. Boreal Austromagellanian Province
13.6. West Cerrado Province [Provincia Austromagallánica Boreal]
[Provincia Cerradense Occidental] 19.4. Insular Falkland Province
13.7. Pantanalian Province [Provincia Islas Malvinas]
[Provincia Pantanalense] 19.5. Insular Juan Fernandez Province
13.8. Benian Province [Provincia Islas Juan Fernández]
[Provincia Beniana] Bc. CIRCUMANTARCTIC Subkingdom
14. CHACOAN Region [Subreino CIRCUNANTÁRTICO]
[Región CHAQUEÑA] 20. INSULAR ANTARCTIC Region
14.1. North Chacoan Province [Región ANTÁRTICA INSULAR]
[Provincia Chaqueña Septentrional] 20.1. Insular Atlantical Antarctic Province
14.2. South Chacoan Province [Provincia Islas Antárticas Atlánticas]
[Provincia Chaqueña Meridional] 21. CONTINENTAL ANTARCTIC Region
Bae. TROPICAL SOUTH ANDEAN Superegion [Región ANTÁRTICA CONTINENTAL]
[Superregión SURANDINA TROPICAL] 21.1. West Antarctic Province
15. TROPICAL SOUTH ANDEAN Region [Provincia Antártica Occidental]
21.2. East Antarctic Province
[Región SURANDINA TROPICAL]
[Provincia Antártica Oriental]
15.1. Desertic Peruvian-Ecuadorean Province
[Provincia Peruana-Ecuatoriana Desértica]
B. Descriptions of biogeographic units
15.2. Mesophytic Punenian Province
[Provincia Puneña Mesofítica] Amazonian: South-American equatorial and Atlantic
15.3. Xerophytic Punenian Province southern eutropical biogeographic region (12), plu-
[Provincia Puneña Xerofítica] vial and tropical mesophytic infra-lower thermotropi-
15.4. Bolivian-Tucumanan Province cal bioclimates and with few rare exceptions subme-
[Provincia Boliviana-Tucumana] sophytic, with varzeal compensation, on soils and
15.5. Yungenian Province waters enriched with ions and nutrients. To the north
[Provincia Yungueña] it borders on the hyperoligotrophic soils and waters
16. HYPERDESERTIC TROPICAL PACIFIC Region with Guyanan-Orinoquian Region; to the south on
[Región PACÍFICA TROPICAL HIPERDESÉRTICA] the infra-thermotropical pluviseasonal lower meso-
16.1. Hyperdesertic North Peruvian Province phytic and submesophytic border with the Brazilian-
[Provincia Norperuana Hiperdesértica] Paranense Region and, at the headwaters of the An-
16.2. Hyperdesertic Tropical Chilean-Arequipan dean mountains with the yungas of the Tropical
Province South Andean Region which have a height of be-
[Provincia Chilena-Arequipeña Tropical tween 500-1000 m. The Amazonian Region compri-
Hiperdesértica] ses five biogeographic provinces: 12.1. West Amazo-
Bb. AUSTROAMERICAN Subkingdom nian, 12.2. North Amazonian, 12.3. Deltaic Amazo-
nian, 12.4. Southwest Amazonian, 12.5 and Central
[Subreino AUSTROAMERICANO]
Amazonian. [Región Amazónica]
17. PAMPEAN Region
[Región PAMPEANA] Amazonian-Guyanan: South-American biogeogra-
17.1. Mesophytic Pampean Province phical superegion (Bac), with tropical macrobiocli-
[Provincia Pampeana Mesofítica] mate formed by the Guyanan-Orinoquian and Ama-
17.2. Xerophytic Pampean Province zonian Regions (11+12). [Superregión Amazónica-
[Provincia Pampeana Xerofítica] Guayanense].
28 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

Argentinian Montean: Biogeographic province of the pical coastal Braziliantropical. South America south
Middle Chilean-Patagonian Region (18.4). Non-tro- of San Francisco river. [Provincia Atlántica Brasile-
pical, desertic and xeric Mediterranean mostly sub- ña].
tropical South America. [Provincia Monte Argenti-
no]. Brazilian-Paranense: Atlantic South-American bio-
geographic region (13), located in amazonic and rio-
Austroamerican: Non-tropical South-American sub- platensean basin. Meridional equatorial (Ceará, Ma-
kingdom (Bb), covering a wide territory usually ranhao), meridional eutropical (Bahía, Planalto, Mato
south of parallel 30ºS, both Mediterranean (Middle Grosso, Rondonia, Pantanal and Beni), and meridio-
Chilean-Patagonian Region) and temperate or boreal nal subtropical to the east of the Paraguay-Paraná ri-
macrobioclimates (Pampean and Valdivean-Magel- ver (parallel 30ºS), whose bioclimate is mostly tropi-
lanian Region). To the north it borders on the tropical
cal pluviseasonal submesophytic, as well as tropical
macrobioclimate with Brazilian-Paranense, Chacoan,
pluvial, at the south of the tropic of Capricorn, and
Tropical South Andean and Hyperdesertic Tropical
Pacific regions. It has gondwanic paleorelationships tropical xeric in the Catinga-Sertao (basin of the San
and later ones with the Neozelandic-Australian King- Francisco river). The Brazilian-Paranense region in-
dom and the Circumantarctic Subkingdom. Three re- cludes eight biogeographic provinces: 13.1. Brazilian
gions are recognized: Pampean (17), Middle Chilean- Atlantic, 13.2. Paranense, 13.3. Catingan, 13.4. To-
Patagonian (18) and Valdivean-Magellanian (19) cantins, 13.5. East Cerrado, 13.6. West Cerrado,
[Subreino Austroamericano]. 13.7. Pantanalian and 13.8 Benian [Región Brasile-
ña-Paranaense].
Austrocircummediterranean: Meridional pluriconti-
nental geographic area with a gondwanic origin, and Caribbean-Mesoamerican:Mesoamerican biogeogra-
Mediterranean macrobioclimate, potentially made up phic region (9), with tropical bioclimate, chiefly plu-
of the Austro-American, South-African and Austra- viseasonal, to a lesser extent pluvial or xeric and ra-
lian evergreen climatophilous forests, semideserts rely desertic in some islands or Antilleancoasts. It in-
and deserts. Territories with Mediterranean macro- cludes all the Caribbean islands, the Florida and Yu-
bioclimate of the southern hemisphere [Área Austro- catan peninsulas, the rainy territories of Veracruz and
circunmediterránea] all over Central America to the north of Darien. The
Caribbean-Mesoamerican Region comprises six bio-
Austrocircumtemperate: Meridional pluricontinental geographic provinces: 9.1. Floridian, 9.2. Cuban, 9.3.
geographic area with a gondwanic origin, with tem-
Lesser Antillean, 9.4. Veracruzenian-Yucatecan, 9.5.
perate bioclimate, potentially formed by the Austro-
Chiapan-Honduran and 9.6. Panamanian-Costa Rican
American, Australian-New-Zealander and South-Af-
rican warm mostly evergreen and gymnospermic [Región Caribeña-Mesoamericana]
climatophilous forests. Territories with temperate Caribbean-Neogranadian: Caribbean and North So-
macrobioclimate of the southern hemisphere [Área
uth-American biogeographic superegion (Bab), with
Austrocircuntemplada].
tropical macrobioclimate formed by the Caribbean-
Austrocoldtemperate: Meridional pluricontinental Mesoamerican and Neogranadian Regions (9+10).
wide geographic area with a gondwanic origin, mos- Caribbean and New Granada. [Superregión Caribe-
tly located south of parallel 35ºS, formed by cold and ña-Neogranadina].
temperate latitudinal and altitudinal zones, to the
Catingan: Biogeographic province of the Brazilian-
south of the subtropical boundary. Territories with
Paranense Region (13.3) mostly tropical xeric. Tro-
temperate macrobioclimate of the southern hemi-
pical South America. Catinga. [Provincia Catinguen-
sphere, often with deciduous forest, with a thermicity
se].
index lower than It<200. [Área Austrocriotemplada].
Central Amazonian: Biogeographic province of the
Benian: Biogeographic province of the Brazilian-Pa-
Amazonian Region (12.5); it has been also called
ranense Region (13.8). Tropical South America; low
Madeira-Tapajoz. Tropical South America. Central
and middle basin of Mamore and Beni rivers, mostly
Amazon. [Provincia Amazónica Central].
infra-thermotropical humid. [Provincia Beniana].
Central Chilean: Biogeographic province of the
Bolivian-Tucumanan: Biogeographic province of the
Middle Chilean-Patagonian Region (18.2). Mediter-
Tropical South Andean Region (15.4). Tropical So-
ranean non-tropical South America. [Provincia Chi-
uth America; eastern Andes of Bolivia and Argentina
lena Central].
as far as Tucuman above the Chacoan Region. [Pro-
vincia Boliviana-Tucumana]. Chacoan: Atlantic tropical South-American biogeo-
graphic region (14), located to the northwest of Para-
Boreal Austromagellanian: Austro-American biogeo- guay river (Paranense biogeographic province) with
graphic province (19.3), with boreal hyperoceanic dry and semiarid tropical xeric bioclimate. Its central
bioclimate, in the south of the Valdivean-Magella- axis is the Pilcomayo river when it reaches the plain
nian Region [Provincia Austromagallánica Boreal]. after flowing across the tropical Andes in the Xero-
phytic Punenian and Bolivian-Tucumanan Provinces.
Brazilian Atlantic: Biogeographic province of the Towards the north it borders on Santa Cruz de la Sie-
Brazilian-Paranense Region (13.1). Infra-thermotro- rra with Chiquitanía, West Cerrado Province, and
Biogeographic Map of South America. A preliminary survey 29

towards the northeast with the Pantanal. Towards the Shelf (20ºW-160º E: West Antarctic or Lesser Ant-
south, when the tropical macrobioclimate disappears arctic), which includes the Ellsworth Mountains with
and evolves to temperate, it come into contact with the Vinson Peak (5140 m) culminating at the top of
the Pampean Region (Xerophytic Pampean Province) Antarctica (West Antarctic Province). 21.2 East Ant-
and when it becomes Mediterranean xeric or desertic arctic Province: this extends along the coast towards
it is replaced by the subdesertic vegetation of the Ar- the east from the Usarp mountains on Oates Coast
gentinean Montean, the start of the North Patagonian (160º E) to the Ekström ice field on Princess Martha
Province of the Middle Chilean-Patagonian Region. Coast (20º W: East Antarctic or Greater Antarctic),
The Chacoan Region comprises two biogeographic which includes as its culminating part the Dome Ar-
provinces: 14.1. North Chacoan and 14.2 South Cha- gus (4030 m); to the west of the French station of
coan. [Region Chaqueña]. Charcot (2435 m) an ice glacier thickness of 4776
has been measured; the record for the coldest tempe-
Chacoan-Brazilian: South-American biogeographic rature on the surface of the Earth (-89.4º C) is held
superegion (Bad) with tropical macrobioclimate ma- by the Russian station of Vostok (3488 m). The Ant-
de up of Brazilian-Paranense and Chacoan Regions arctic Continent, with more than 13.5 million square
(13 + 14). [Superregión Chaqueña-Brasileña]. kilometres, was the center of the Gondwana super-
continent 180 million years ago, after which a set of
Chiapan-Honduran : Biogeographic province of the subcontinents: South America, Africa, India, Austra-
Caribbean-Mesoamerican Region (9.5). Central A- lia and New Zealand separat and slid toward the
merica. [Provincia Chiapaneca-Hondureña]. north, leaving the Antarctica more or less fixed
around the South Pole, isolated and perhaps partly or
Circumantarctic: Biogeographic subkingdom (Bc), wholly under ice cover for 40 million years. Today,
with polar macrobioclimate, exceptionally boreal hy- 98 percent of the surface of Antarctica is covered by
peroceanic in some subtemperate isles, hypergelid to a thick ice cover of about 2.5 km of thickness on av-
a large extent made up of the Antarctic Continent and erage. There are also wide flat glacial floating planes
the islands and archipelagos peripheral to the Ant- (ice shelf), some of 1000 km in length in protected
arctic Peninsula and the Antarctic Continent (Conti- coastal areas (Ross & Ronne Ice Shelves). The an-
nental Antarctic Region), as well as the islands far nual precipitation exceeds 600 mm in some hypero-
from the continent, immersed in the westerly winds ceanic windward coastal or insular adjacent locali-
and the icy waters of the Antarctic Convergence (In- ties, while many extreme continental areas inside the
sular Antarctic Region: South Orkney, South Geor- center-east have less than 30 mm. The dominant bio-
gia, South Sandwich, Bouvet, Prince Edward, Crozet, climate is polar pergelid and only upper suprapolar
Kerguélen, McDonald, Macquarie, Balleny, Scott, (Tp<20) in the marked hyperoceanic coasts and at-
etc.). The Circumantarctic Subkingdom comprises tached islands (Graham Land at the end of the Ant-
two biogeographic regions: 20. Insular Antarctic and arctic Peninsula and surrounding islands, especially
21. Continental Antarctic. [Subreino Circunantárti- in sunny places with environmental humidity). Only
two vascular plants are known, with distribution
co].
Austro-American linked to coastal habitats with
Colombian Andean: Biogeographic province of the freshwater or very little brackish and seasonal hu-
Neogranadian Region (10.3). Tropical South Ameri- midity available, nevertheless there are several hun-
ca of the Andes from Cotopaxi volcano in Ecuador dred lichen species, bryophytes, fungi and proto-
phytes, especially on well-exposed rock habitats
next to Quito to the Andes of Mérida, Eastern and
where melt water or cryptoprecipitations are avail-
Central Range and Cauca Valley, extending from in-
able at least some time during the summer. Among
fratropical xeric in the inner Magdalena deep valleys birds and mammals that breed on the continent or
to cryorotropical pluvial in the snowy high moun- adjacent islands, always in low continental sea-land
tains. It has been also named Andina Paramuna due stations, it is worth mentioning five species of pen-
to its cold and humid high plains enriched with espe- guins, more than a dozen bird and six seal species.
letineans (frailejones). [Provincia Andina Colombia- Despite having been able to detect warmer and
na]. colder periods in Antarctica over the past 150,000
years and correlate them with the concentrations of
Colombian Pacific: Biogeographic province of the carbon dioxide –an increased greenhouse-gas effect
Neogranadian Region (10.5). Tropical South Ameri- in the Earth– in the last 50 years the increase of CO2
ca. [Provincia Pacífico Colombiana]. from 280 ppm to 370 ppm (global warming) does not
seem to have had a significant impact on the current
Continental Antarctic: Biogeographic region belon- increase in temperature in East Antarctica. [Región
ging to the Circumantarctic Subkingdom (21) with Antártica Continental].
pergelid polar bioclimate and scarce suprapolar bio-
climate Tp<20. Because of its orography, bioclimate, Cordobesa-Lower Magdalena: Biogeographic prov-
geography and biota, two great territories can be ince of the Neogranadian Region (10.4). Xeric to
identified on the Antarctic Continent, to the west and pluvial infra-thermotropical Caribbean tropical South
east of the Transantarctic Mountains, to which we America of the plains, hills and dams of the Sinu,
confer the provincial biogeographic level. 21.1. West Magdalena and Cauca basins from Barrancabermeja
Antarctic Province: this extends along the coast to-
and Cáceres to the swamps close to Plato. [Provincia
ward the west and south from the Filchner and Ronne
Cordobesa-Bajomagdalena].
ice shelves to follow the coasts close to the Ross Ice
30 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

Deltaic Amazonian: Biogeographic province of the Guajiran-Caribbean: Biogeographic province of the


Amazonian Region (12.3), infratropical pluvial and Neogranadian Region (10.1). Tropical South Ameri-
pluviseasonal mesophytic. Tropical South America. ca. Caribbean from Cartagena to the Paria Peninsula,
Amazonian Delta. [Provincia Amazónica Deltaica]. with the Guajira, Maracaibo, Falcón, Barquisimeto,
Valencia, Caracas and coastal Venezuelan moun-
Deltaic Orinoquian: Biogeographic province of the tains. To the south it borders with the Llaneran
Guyanan-Orinoquian Region (11.2), infratropical (10.2), Colombian Andean (10.3) and Cordobesa-Lo-
pluvial. Tropical South America. Orinoquian Delta. wer Magdalena (10.4) biogeographical provinces.
[Provincia Orinoquense Deltaica]. [Provincia Guajireña-Caribeña].
Desertic Mediterranean Chilean: Biogeographic Guaviarean-Orinoquian : Biogeographic province of
province of the Middle Chilean-Patagonian Region the Guyanan-Orinoquian Region (11.3). Tropical
(18.1). Nontropical South America. Toward the south South America. [Provincia Guaviareña-Orinoquen-
of Antofagasta (24º C) the coastal and interior hyper- se].
deserts, deserts and semideserts are already Mediter-
ranean (winter rainfall) until Region V (32ºS), and Guayaquilian-Ecuadorean: Biogeographic province
belong to the Desertic Mediterranean Chilean Prov- belonging to the Neogranadian Region (10.6). Pacific
ince which, from the extreme hyperdeserts (Io 0.0- equatorial tropical South America. It extends along
0.1) lacking in vascular climatophilous vegetal cover the coast from the Guayaquil Gulf (3ºS) up the An-
to the III Region (26ºS), continues northwards with con Bay on the Colombian border (1º 30’N), ex-
the acute and moderate deserts with arid cacti such as cluding a desertic narrow coastal territory with man-
Eulychnia tenuis and Copiapoa marginata. North of groves going from the Puná Island to the Manta Bay.
Vallenar in the IV Region (29ºS) the arid deserts (Io In contrast, the coast is always hyperarid and ultra-
0.4-0.9) with large cacti of Eulychnia breviflora ap- hyperarid with cold sea waters and without man-
pear; and reaching up until north of Coquimbo (30º groves extending from latitude 4º 10’ S up to 24ºS to
S), the semideserts (Io 1.0-1.9) with large Cactaceae, the Hyperdesertic Tropical Pacific Region (16)]. The
rosulate puya and shrubs of Eulychnia acida, Echi- Andean ranges of the Colombian Andean Province
nopsis skottsbergii, Echinopsis litoralis, Puya chilen- (10.3), with the typical frailejones, extend as far as
sis and Lithraea caustica, preamble to the Mediterra- Ecuador up to the snow covered volcanoes of Coto-
nean pluviseasonal sclerophylous forests and chapar- paxi (5897 m) and Iliniza (5383 m) (1ºS). South of
rals of the class Lithraeo-Cryptocaryetea, typical of this latitude the mountain chain continues the
the Central Chilean Province (18.2). [Provincia Chi- Guayaquilian-Ecuadorean Province, whose highest
lena Mediterránea Desértica]. peak is the Chimborazo (6310 m). In the high humid
or even hyperhumid supra-cryorotropical high
Desertic Peruvian-Ecuadorean. Biogeographic prov- mountains there are short and tall pajonales and
ince of the Tropical South Andean Region (15.1). scrubland climatophilous communities, as well as the
Deserts and semideserts (Io 0.4-2.0) of the tropical perennifolius meso-micro-cloud-forests which be-
mountains and coasts of the Andean western slopes, come serial shrubby pajonales when damaged. In the
from the coast of Manta Bay in Ecuador (1º S), that upper semiarid-subhumid infra-mesotropical belts,
together with the Tumbes Peruvian coast up to Ma- mostly on the pacific slope, the natural matured
chala, with the short tree Loxopterigium huasango, vegetation, seriously damaged by agriculture, corre-
constitute two biogeographic districts: Desertic Coas- sponds to deciduous and semideciduous micro-meso-
tal Ecuadorean and Tumbesian, in Peru from El Alto forests that in the hottest areas can be thorny or doli-
in Piura (4º 10’ S) towards the south leaves the lit- form. The xeric mesotropical deciduous forests with
toral to cover the interior deserts and semideserts and small doliform trees and cacti are also visible at the
the western Andean foothills up until Tarata in Tacna south of the province (6ºS) next to the Abra Porculla
and the borders of Chile (18º S). In contrast, the ul- (2145 m) and Bagua low hot valley, in the Marañón
trahyperarid and hyperarid hyperdeserts (Io 0.0-0.4) basin. Eastwards from the Napo to the Pastaza rivers
belong to the Hyperdesertic Tropical Pacific Region it borders the humid lower thermotropical pluvial
(16) extending along the coasts, ridges and moun- belt with the West Amazonian Province (12.1) and at
tains ranges from Talara (4º 30’ S) in Peru to Antofa- the Condor Mountain Range and the Chinchipe ba-
gasta (24º S) in Chile. Tropical South America. [Pro- sin, the humid-hyperhumid meso-lower supratropical
vincia Peruana-Ecuatoriana Desértica]. pluvial belt with the lauroid and Podocarpus humid
cloud forests of the Yungenian Province (15.5)
East Antarctic: Biogeographic province of the Conti- coming from the Eastern Peruvian Andes. [Provincia
nental Antarctic Region (21.2), polar pergelid and Guayaquileña-Ecuatoriana].
scarcely upper suprapolar (Tp<10) in sun exposed-
coasts. East Antarctic. This could be also named Gre- Guyanan: Biogeographic province of the Guyanan-
ater Antarctic. [Provincia Antártica Oriental]. Orinoquian Region (11.1). Tropical South America.
[Provincia Guayanense].
East Cerrado: Biogeographic province of the Brazil-
ian-Paranense Region (13.5). Tropical South Ameri- Guyanan-Orinoquian: Equatorial and Atlantic north-
ca. Eastern Cerrado. [Provincia Cerradense Orien- ern eutropical south American biogeographic region
tal]. (11), with pluvial and pluviseasonal bioclimate, ex-
Biogeographic Map of South America. A preliminary survey 31

tending over the whole of the hard Guyanese Shield Insular Antarctic: Biogeographic region belonging to
and the surrounding sandy fluvial and windy sand the Circumantarctic Subkingdom (20) with polar hy-
deposits which give rise to soils and waters which are peroceanic, oceanic and pergelid bioclimate, and ex-
extraordinarily poor in nutrients and bases such as ceptionally boreal hyperoceanic only formed by isles
Old Guyana, Vichada, Guaviare, Vaupés and Ro- located in the austral glaciated sea around the Ant-
raima territories. To the north and south it borders on arctic Continental (Continental Antarctic Region),
the Neogranadian and Amazonian Regions respecti- linked by the permanent or temporary iced sea all the
vely, both of which have rich soils and waters. The year (banquisa) and the meeting between the ice wa-
Guyanan-Orinoquian Region comprises five biogeo- ters (-2ºC) and temperate ones (0º to 3ºC) of the Ant-
graphic provinces: 11.1. Guyanan, 11.2. Deltaic Ori- arctic Convergence (about 1600 km of the coast),
noquian, 11.3. Guaviarean-Orinoquian , 11.4. Tepu- adjacent to the circumpolar current, driven by the
yan and 11.5. Guyanese Brazilian . [Región Guaya- west wind drift. Inside the region we accept three
na-Orinoquense]. oceanic groups of islands or archipelagos at the bio-
geographic province level: 20.1 Insular Atlantic Ant-
Guyanese Brazilian : Biogeographic province of the arctic Province (20ºE-80ºW): Bouvet Island, South
Guyanan-Orinoquian Region (11.5). Tropical South Georgia Islands, Diego Ramírez Islands, South
America. [Provincia Brasileña Guayanense]. Sandwich Islands and South Shetland Islands; 20.2
Insular Indian Antarctic Province (20ºE-140ºE): Cro-
Hyperdesertic North Peruvian : Biogeographic pro-
zet Island, Prince Edward Island, Heard Island, Ker-
vince of the Hyperdesertic Tropical Pacific Region
guélen Islands and McDonald Islands and 20.3. In-
(16.1). Tropical South America. Hyperdesertic north-
sular Pacific Antarctic Province (140ºE-80ºW):
ern Peru. [Provincia Norperuana Hiperdesértica].
Balleny Islands, Macquarie Islands, Peter Island and
Hyperdesertic Tropical Chilean-Arequipan: Biogeo- Scott Islands. The terrestrial and oceanic-terrestrial
graphic province of the Hyperdesertic Tropical Paci- biota is quite poor and with gondwanic lineage, but
fic Region (16.2). Tropical South America. Hyperde- with influences of the nearby large continents and
sertic Tropical Chile and Arequipa, ultrahyperarid islands: South America, Africa, Australia, New Zea-
and hyperarid coasts, ridges and mountain ranges of land and Tasmania. [Región Antártica Insular].
Northern Arequipa, Moquegua and Tacna Peruvian Insular Atlantical Antarctic: Biogeographic insular
departments. [Provincia Chilena-Arequipeña Tropi- province of the Insular Antarctic Region (20.1). At-
cal Hiperdesértica]. lantic Antarctic archipelagos and islands: South
Hyperdesertic Tropical Pacific: South American tro- Georgia, Diego Ramirez, Bouvet, South Sandwich
pical biogeographic region (16), extending from 4º and South Shetland (20º E-80º W) with thermo-su-
10’ S in the hyperarid coastal of Talara (Peru) to the prapolar and pergelid bioclimates. [Provincia Islas
extreme ultrahyperarid tropical deserts of Antofa- Antárticas Atlánticas].
gasta (24ºS). To this region belongs the extreme
marked and moderate hyperdeserts, thermo-mesotro- Insular Falkland: Biogeographic insular province of
pical hyperarid and ultrahyperarid (It > 320; Io < the Valdivean-Magellanian Region (19.4). Atlantic
0.4), which in the Hyperdesertic Tropical Chilean- boreal (antiboreal) South America. [Provincia Islas
Arequipan Province can reach almost 3000 meters, Malvinas].
with the moderate hyperdeserts of columnar cactus
of Browningia candelaris. The rocky or clayey hy- Insular Galapagos: Biogeographic insular province
perdeserts, without occasional hydric contribution or of the Neogranadian Region (10.7). Xeric-desertic
fog, lack vascular vegetation cover, but the misty de- pacific equatorial tropical South America. [Provincia
serts with unmeasurable drizzled moisture during Islas Galápagos].
some weeks in year, have different types of vegeta-
tion depending on the topography and substrate Insular Juan Fernandez: Biogeographic insular pro-
(coastal ridges), such as the aerohygrophile commu- vince of the Valdivean-Magellanian Region (19.5).
nities of Tillandsia sp. pl. which grow on several Mediterranean and temperate subtropical pacific non
substrates and dune communities, or those than grow tropical South America. [Provincia Islas Juan
on deep sandy soils formed by numerous radicant, Fernández].
geophytic or terophytic plants. In the hyperdeserts
the natural matured vegetation corresponds to very Lesser Antillean: Biogeographic province of the Ca-
open formations of different cactus, some of them ribbean-Mesoamerican Region (9.3). Antilles Is-
crassiarborescents columnar microphanerophytics lands. [Provincia Antillana Menor].
with very low growth such as Neoraimondia arequ-
ipensis. The Hyperdesertic Tropical Pacific Region Llaneran: Biogeographic province of the Neograna-
comprises two biogeographic provinces: 16.1. Hy- dian Region (10.2). Xeric and pluviseasonal infratro-
perdesertic North Peruvian and 16.2. Hyperdesertic pical Atlantic tropical South America of the plains,
Tropical Chilean-Arequipan. However it has few flo- hills, rivers and dams of water in rich soils from the
ristic and vegetational relationships with the deserts river Meta to Guajira river, with hydrophytic herba-
and semideserts of the Desertic Peruvian-Ecuadorean ceous and humid wooded natural permanent savanna
biogeographic province (15.1). [Región Pacífica Tro- to pluviseasonal evergreen or deciduous rich soil for-
pical Hiperdesértica]. est vegetation climax. [Provincia Llanera].
32 S. Rivas-Martínez, G. Navarro, A. Penas & M. Costa

Mediterranean Andean: Biogeographic province of Neotropical-Austroamerican: American and Antarc-


the Middle Chilean-Patagonian Region (18.3). Non- tic Biogeographic Kingdom (B) established by three
tropical South America. Mediterranean Andes. [Pro- broad subkingdoms having distinct bioclimates: Ba.
vincia Andina Mediterránea]. Neotropical (tropical bioclimates: North, Central and
South America); Bb. Austro-American (temperate,
Mesophytic Pampean: Biogeographic province of the mediterranean and boreal bioclimates): Bc. Circum-
Pampean Region (17.1). Subhumid and humid tem-
antarctic (polar bioclimates: Antarctic and related ar-
perate Atlantic (30º-40º S) South America. [Provin-
cia Pampeana Mesofítica]. chipelagos). [Reino Neotropical-Austroamericano]

Mesophytic Punenian: Biogeographic province of the North Amazonian: Biogeographic province of the
Tropical South Andean Region (15.2). Tropical Amazonian Region (12.2); it has been also named
mesophytic and hygrophytic subhumid to hyperhu- Roraima. Tropical South America. Western Amazon.
mid puna of South America. [Provincia Puneña Me- [Provincia Amazónica Septentrional].
sofítica].
North Chacoan: Biogeographic province of the Cha-
Middle Chilean-Patagonian: Pacific and Atlantic coan Region (14.1). Tropical South America. North-
non-tropical South American biogeographic region ern Chaco. [Provincia Chaqueña Septentrional].
(18), all with Mediterranean macrobioclimate. The
region obliquely crosses the subcontinent from one North Patagonian: Biogeographic province of the
ocean to another (Pacific coast: 24º-38º S, Atlantic Middle Chilean-Patagonian Region (18.5). Non-tro-
coast 40º-52º S); across a broad Mediterranean pical Mediterranean South America. [Provincia Pa-
bridge in the high Andes (approx. 30º-35ºS). Its tagónica Septentrional].
northern Andean mountain zone corresponds to the
puna floristic district Cuyano (MARTÍNEZ CARRE- Neogranadian: Neotropical biogeographic region
TERO 1995). From the extreme ultrahyperarid ther- (10) cover a wide northern territory of South Ameri-
mo-mesomediterranean hyperdeserts of Atacama ca from the inner Guayaquil Bay and the Andean Pe-
(24ºS) and the central Chilean deciduous and scle- ruvian-Ecuadorean mountains south of Loja (6ºS) to
rophyllous dry to subhumid thermo-mesomedite- the Gulf of Panama in the Pacific and the Gulf of Da-
rranean micro-mesoforests (38ºS), it reaches the oro- rien, the Guajira (12º 30’ N) and Paria (11ºN) penin-
mediterranean belt of the cordillera with the clima- sulas in Caribbean Sea (10ºN). To the east and south
tophilous scrubland and “pajonales” of the Medite- it borders on the Guyanan-Orinoquian (11), Amazo-
rranean Andean Province and, in the high Andes, the nian (14) and Tropical South Andean (15) regions.
climatophilous cryoromediterranean dry-humid pul- The Neogranadian Region (see provinces text) com-
vinate vegetation. Beyond the cordillera appears the prises seven biogeographic provinces: 10.1. Guaji
meso-megascrubland, more or less thermic semiarid- ran-Caribbean, 10.2. Llaneran, 10.3. Colombian An-
arid of the Argentine Monte biogeographic Province, dean, 10.4. Cordobesa-Lower Magdalena, 10.5.
and continues towards the south, on the eastern side Colombian Pacific, 10.6. Guayaquilian-Ecuadorean
of the cordillera, with northern or southern Pata- and 10.7. Insular Galapagos. [Región Neogranadina]
gonian supra-oromediterranean semiarid-dry dwarf
shrubby vegetation. As far as the coast in Rio Galle- Pampean: Non-tropical Atlantic Temperate South-
gos (51º 50’ S), appears the temperate xeric macro- American biogeographic region (17) belonging to the
bioclimate with the productive grasslands in summer Austro-American Subkingdom. All the region has a
and, more toward the southwest the remains of the temperate macrobioclimate (temperate oceanic and
old micro-mesoforests of Nothofagus, particularly xeric), located between 30ºS and 40ºS parallels and
when the ombrotype is subhumid or humid with the chiefly in the low basins of the Paraná and Uruguay
Valdivean-Magellanian mesoforest of Wintero-No-
rivers which come from the tropical Brazilian-Para-
thofagetea class. The Middle Chilean-Patagonian Re-
nense Region (approx. 30ºS). A considerable part of
gion comprises six biogeographic provinces: 18.1.
Desertic Mediterranean Chilean, 18.2. Central Chi- the Pampean Region territory, except for its internal
lean, 18.3. Mediterranean Andean, 18.4. Argentine and surrounding mountains ranges: Ventana, Carape,
Monte, 18.5. North Patagonian and 18.6. South Pata- San Luis and Córdoba (2884 m), has a very recent
gonian. [Región Mesochilena-Patagónica]. origin (Holocene). For this reason the native flora
and particularly the arborescent one is very poor.
Neotropical: Biogeographic subkingdom of America Towards the south and west of the rioplatensean te-
(Ba), with tropical macrobioclimate; that is to say, all rritorial region the bioclimate changes to Mediterra-
territories of the equatorial and eutropical latitudinal nean xeric or desertic and consequently to the wide
belts (0º-23º N & S), as well as the subtropical belts Middle Chilean-Patagonian Region, always across
(23º a 35º N & S) having tropical macrobioclimate. the Argentine Monte Province. The Pampean Region
Five biogeographic superegions are recognized in comprises two biogeographic provinces: 17.1. Meso-
this subkingdom: Baa. Mexican (Mexican Xerophy- phytic Pampean and 17.2. Xerophytic Pampean. [Re-
tic and Madrean (regions: 7+8), Bab. Caribbean- gión Pampeana]
Neogranadian (regions: 9+10), Bac. Amazonian-Gu-
yanan: (regions: 11+12), Bad. Chacoan-Brazilian (re- Panamanian-Costa Rican: Biogeographic province
gions: 13+14) and Bae. Tropical South Andean (re- of the Caribbean-Mesoamerican Region (9.6). Cen-
gions: 15+16). [Subreino Neotropical]. tral America. [Provincia Panameña-Costarricense].
Biogeographic Map of South America. A preliminary survey 33

Pantanalian: Biogeographic province of the Brazil- and connect with the Tropical South Andean along a
ian-Paranense Region (13.7). Tropical South Ameri- narrow band coming from the Chiquitanía. The con-
ca. The Pantanal of South America, mostly covered tact with the tropical xeric Chacoan Region is esta-
with hydrophytic herbaceous and permanent or tem- blished slightly towards the south, to finish at the
porary flooded wooded natural permanent savanna. south of the 30ºS parallel in the Mediterranean
[Provincia Pantanalense]. mountains and plains of the Middle Chilean-Patago-
nian Region of the provinces: Mediterranean An-
Paranense: Biogeographic province of the Brazilian- dean, Argentine Monte and Central Chilean. The
Paranense Region (13.2). Tropical South America. Tropical South Andean Region comprises five bio-
[Provincia Paranaense]. geographic provinces: 15.1. Desertic Peruvian-Ecua-
dorean, 15.5. Yungenian, 15.2. Mesophytic Pune-
South Chacoan: Biogeographic province of the Cha- nian, 15.3. Xerophytic Punenian and 15.4. Bolivian-
coan Region (14.2). Tropical South America. South- Tucumanan. [Región Surandina Tropical].
ern Chaco. [Provincia Chaqueña Meridional].
Valdivean: Biogeographic province of the Valdivean-
South Patagonian: Biogeographic province of the Magellanian Region (19.1), from mesotemperate to
Middle Chilean-Patagonian Region (18.6). Non.tro- cryorotemperate subhumid to ultrahyperhumid. Non-
pical Mediterranean South America. [Provincia Pa- tropical temperate and boreal South America. [Pro-
tagónica Meridional]. vincia Valdiviana].

Southwest Amazonian: Biogeographic province of Valdivean-Magellanian: Pacific and Austro-Atlantic


the Amazonian Region (12.4); it has been also named South-American biogeographic region (19), extend-
Acre and Madre de Dios. Tropical South America. ing southwards from the Araucanía Region IX in
South-western Amazon. [Provincia Amazónica Sur- Chile (38ºS) and Rio Gallegos (51º50’S) in Argenti-
occidental]. na, with woodland and forest of Wintero-Nothofage-
tea vegetation class. Its macrobioclimate is temperate
Temperate Magellanian: Biogeographic province of and boreal. The boundary between the temperate and
the Valdivean-Magellanian Region (19.2). Humid-ul- boreal macrobioclimates is located in the middle of
trahyperhumid temperate non-tropical Pacific South Isla Grande in Tierra de Fuego and the adjacent con-
America.[Provincia Magallánica Templada]. tinental territory of the Strait of Magellan at parallel
53º30’, as well as in Queen Adelaide Archipelago in
Tepuyan: Biogeographic province of the Guyanan- the Pacific (52ºS). The Falkland (Malvinas) islands
Orinoquian Region (11.4), mostly meso-supratropi- with boreal hyperoceanic bioclimate constitute a bio-
cal pluvial. Tropical South America. Tepuys. [Pro- geographic province of this region. The extreme hy-
vincia Tepuyana]. peroceanic, temperate and Mediterranean subhumid-
humid Juan Fernandez Islands with a high number of
Tocantins: Biogeographic province of the Brazilian-
endemisms with gondwanic and magellanic origin
Paranense Region (13.4), mostly infratropical pluvi-
are also included in this region. The Valdivean-Ma-
seasonal. Tropical South America. Tocantins River.
gellanian Region comprises five biogeographic pro-
[Provincia Tocantinense].
vinces: 19.1. Valdivean, 19.2. Temperate Magella-
Tropical South Andean: South American biogeogra- nian, 19.3. Antiboreal Magellanian, 19.4. Insular Fal-
phical superegion (Bae), with tropical macrobiocli- kland and 19.5. Insular Juan Fernandez. [Región Val-
mate formed by the Tropical South Andean and Hy- diviana-Magallánica].
perdesertic Tropical Pacific Regions (15+16). Tropi-
West Amazonian: Biogeographic province of the
cal western coast and Andes. [Superregión Surandi-
Amazonian Region (12.1); also named Loreto Pro-
na Tropical]
vince. Tropical South America. Western Amazon.
Tropical South Andean: South American biogeogra- [Provincia Amazónica Occidental].
phic region (15) belonging to the tropical Andes of
the southern hemisphere, extending from the yungas West Antarctic: Biogeographic province of the Con-
and humid punas of the Amazon and Cajamarca de- tinental Antarctic Region (21.1), polar pergelid and
partments and South Piura semideserts, as well as the scarcely upper suprapolar (Tp <20) in the coast and
arid Ecuadorean coasts at the south of parallel (1ºS), adjacent islands to Palmer Land in the Antarctic
to the cardonales and Chilean-Argentinian tropical Peninsula. West Antarctic. It is also known as Lesser
desertic and semidesertic punas of the Cerro las Antarctic. [Provincia Antártica Occidental].
Tórtolas (6120 m) near the Agua Negra Pass (4735
m) (30ºS). To the north and west it borders on the West Cerrado: Biogeographic province of the Brazi
Neogranadian Region, inside Guayaquil Bay. To the lian-Paranense Region (13.6). Tropical South Ame-
east, between the 4ºS and 17ºS parallels, below the rica. Western Cerrado. [Provincia Cerradense Occi-
Andean yungas (600-900 m) the pluvial and meso- dental].
phytic pluviseasonal humid-hyperhumid warm infra-
thermotropical (It>640) rainforests belong to the Xerophytic Pampean: Biogeographic province of the
Amazonian Region. In Santa Cruz de la Sierra (Boli- Pampean Region (17.2). Non-tropical temperate At-
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Steyermark, JA. 1979. Plant refuge and dispersal centres in gical systems of the geo-biosphere/ Siegmar-Walter Breckle;
Venezuela: their relict and endemic element. In Tropical translated from 7th, completely revised and enlarged German
Botany, eds. K. Larsen and LB. Holm-Nielsen, 185-221. edition by Gudrun & D. Lawlor.-4th, completely rev. and enl.
London. ed. p. cm.
Stutz de Ortega, LC. 1983. Études floristiques de divers stades Walter, H. & Straka, H. 1970. Arealkunde. Floristisch-histo-
secondaires des formations forestières du Haut Parana (Pa- rische Geobotanik. Eugen Ulmer Verlag. 2 Aufl. Stuttgart.
raguay oriental). Inventaire floristique d’une reserve fores- Wasum, R. A. 2005. Estudio fitosociológico de la vegetación
tiére. Candollea 38: 543-573. de los municipios de São Francisco de Paula, Bom Jesus y
Stutz de Ortega, LC. 1984. Études floristiques de divers stades Jaquirana, RS. (Brasil). Tesis Doctoral. Universidad de
secondaires des formations forestières du Haut Parana (Pa- León. España. 680 pp.
raguay oriental). dynamisme et reconstitution d’une forêt se- Weberbauer, A. 1945. El mundo vegetal de los Andes Perua-
condaire peu degradée. Candollea 39(2): 385-394. nos. Estación Experimental Agrícola de la Molina.Ministerio
Stutz de Ortega, LC. 1986. Études floristiques de divers stades de Agricultura. 776 pp. Lima.
secondaires des formations ferestières du Haut Parana (Pa- Williams, L. 1945. The phytogeography of Peru. In Plants and
raguay oriental). Floraison, fructification et dispersion des Plant Science in Latin America, ed. F. Verdoorn, 308-312.
espèces forestiès. Candollea 41: 121- 144. Waltham, Mass.
Stutz de Ortega, LC. 1987. Études floristiques de divers stades Witte, H.J.J. 1995. Seasonal and altitudinal distribution of
secondaires des formations forestières du Haut Parana (Pa- precipitation, temperature and humidity in the Parque Los
raguay oriental). Structure, composition floristique et régé- Nevados transect (Central Cordillera, Colombia). In Ham-
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fôret sélectivement esploitée. Candollea 42: 205-262. Andean Ecosystems, 4: 279- 327.
International Journal of Geobotanical Research, 1

BIOGEOGRAPHIC MAPS OF
BIOGEOGRAPHICAL UNITS
-90° -80° -70° -60° -50° -40° THE WORLD: SOUTH AMERICA
9.3
B. NEOTROPICAL-AUSTROAMERICAN Kingdom [NEOTROPICAL-AUSTROAMERI- onduras Basse-Terre
Guadeloupe SALVADOR RIVAS-MARTÍNEZ, GONZALO NAVARRO, ÁNGEL
CANO] G . de H
Dominica PENAS & MANUEL COSTA
P
!

Ba. NEOTROPICAL Subkingdom [NEOTROPICAL]


Bab. CARIBBEAN-NEOGRANADIAN Superegion [CARIBEÑA-NEOGRANADINA] C. Gracias a Dios Fort-de-France !P Martinique

C a r i b b e a n S e a
9. CARIBBEAN-MESOAMERICAN Region [CARIBEÑA-MESOAMERICANA] Guatemala 9.5 Castries !P St. Lucía
with collaboration of:

o
Kingstown !P 9.3 Barbados
P
!

c
9.3. Lesser Antillean Province [Antillana Menor] San Salvador !P Javier Amigo (Chile and Argentina), Alindo Butzke (Brazil), Sara del Río (Argentina,

Co
9.5. Chiapan-Honduran Province [Chiapaneca-Hondureña] Tegucigalpa
Gua 10.1 St. Vincent Bridgetown Brazil and Paraguay), Antonio Galán (Peru), José Guevara (Venezuela), Jesús
P
! P
!

9.6. Panamanian-Costa Rican Province [Panameña-Costarricense] tem Izco (Ecuador), Eduardo Martínez Carretero (Argentina), Orlando Rangel
N O R T H
Managua St George's
10. NEOGRANADIAN Region [NEOGRANADINA] a la (Colombia), Salvador Rivas Sáenz (South America bioclimate expert), Fidel
I. Margarita Grenada
P
!

10.1. Guajiran-Caribbean Province [Guajireña-Caribeña] Tre L. Nicaragua C. de la Aguja Tobago


Roig (†) (Argentina), Daniel Sánchez-Mata (South Chile), Leopoldo G. Sancho
P
!

10.2. Llaneran Province [Llanera] nc Barranquilla


5800 !(Maracaibo 10.1 Caracas P Port of Spain
h (South Chile and Antarctica), Pilar Soriano (Venezuela) and Oscar Tovar (†) (Peru)
#
Sierra NevadaL. Maracaibo
(
!

10.3. Colombian Andean Province [Andina Colombiana] Trinidad


!
Cartagena
San José Panamá de Sta. Marta
P
!
(
! ( Valencia
!

10.4. Cordobesa-Lower Magdalena Province [Cordobesa-Bajomagdalena] G. of Darién


Barquisimeto
(
!

10°
P
!
Canal 10.1 10.1 11.2
10°
0 250 500 1.000 1.500 km
10.5. Colombian Pacific Province [Pacífico Colombiana] 9.6

A T L A N T I C
Ciudad
10.6. Guayaquilean-Ecuadorean Province [Guayaquileña-Ecuatoriana] 10.4 co
Panamá O rino !(Guayana
P
!

10.7. Insular Galapagos Province [Islas Galápagos] Cúcuta Lambert Equal-Area Azimuthal Projection

a le na
( San Cristóbal
Bac. AMAZONIAN-GUYANAN Superegion [AMAZÓNICA-GUAYANENSE] 11.3
(
!

10.2 Georgetown
!
Bucaramanga
BIOCLIMATES

a gd
11. GUYANAN-ORINOQUIAN Region [GUAYANA-ORINOQUENSE] 10.5

á
(
!
P
!

M e ta

n am
11.1. Guyanan Province [Guayanense] Paramaribo

M
O C E A N
Medellín
11.2. Deltaic Orinoquian Province [Orinoquense-Deltaica]
(
!

Authors: S. Riva s Sáenz; Rivas-Martínez, S.; Pe nas, Á; Navarro , G.; Costa, M.


P
!

2810 # Cayenne

Pa
11.3. Guaviarean-Orinoquian Province [Guaviareña-Orinoquense] 11.4 Mt. Roraima
11.4. Tepuyan Province [Tepuyana] Bogotá C. Orange -90° -80° -70° -60° -50° -40°

de
P
!

lf 11.3 11.1
Go
11.5. Guyanese Brazilian Province [Brasileña Guayanense]
P
!

10.3

o
12. AMAZONIAN Region [AMAZÓNICA] 15 13

Co
e q ui b o
12.1. West Amazonian Province [Amazónica Occidental]

ur an
12.2. North Amazonian Province [Amazónica Septentrional] Guaviare 13

ty n
Es s
12.3. Deltaic Amazonian Province [Amazónica Deltaica]

e
10° 10°

n co
12.4. Southwest Amazonian Province [Amazónica Suroccidental]
12.5. Central Amazonian Province [Amazónica Central] C. de San Francisco 14

Br a
11.5 Marajó I.
Bad. CHACOAN-BRAZILIAN Superegion [CHAQUEÑA-BRASILEÑA] Ca Ne g ro
q uet Equator
13. BRAZILIAN-PARANENSE Region [BRASILEÑA-PARANAENSE] Ga lápagos Is. Quito!P
Cotopaxi
á
0° 14
13.1. Brazilian Atlantic Province [Atlántica Brasileña] 0° 5897
#
12.2
13.2. Paranense Province [Paranaense] 10.7 15.1 Japurá zo n
10.6 # Chimborazo ma Belém 0°
A 15
(
!

13.3. Catingan Province [Catinguense] Guayaquil 6267 Putumayo São Luis 13
Na
13.4. Tocantins Province [Tocantinense] 10.7 p Santarém
(
!
(
!
Manaus
13.5. East Cerrado Province [Cerradense Oriental] 12.1 Am
(
!

12.3 12
(
!

G. of Guayaquil
a z on

Toca ntins
13.6. West Cerrado Province [Cerradense Occidental] Fortaleza
13.7. Pantanalian Province [Pantanalense]
Iquitos
13
(
!
(
!
Ma

Parnaib a
13.8. Benian Province [Beniana] Pta. Pariñas r añón C. São Roque 14

Xi n g u
14. CHACOAN Region [CHAQUEÑA] Teresina!( -10°

jós
-10°
ira
12.5

pa
14.1. North Chacoan Province [Chaqueña Septentrional] Ju r u á

Ta
Natal
14.2. South Chacoan Province [Chaqueña Meridional] Pta. Negra 16.1

de
s
(
!

Ma
Bae. TROPICAL SOUTH ANDEAN Superegion [SURANDINA TROPICAL] ru
Pu

Uc ay
Ar
(
!

15. TROPICAL SOUTH ANDEAN Region [SURANDINA TROPICAL]


Chiclayo
15

ip
a
15.1. Desertic Peruvian-Ecuadorean Province [Peruana-Ecuatoriana Desértica]

Tel e
li

uan
Roose
15.2. Mesophytic Punenian Province [Puneña Mesofítica] Trujillo Huascarán ir e 13.4
(
!
Pôrto Velho
11 13 -20°

s
á
13
(
!
s isc o -20°

P
(
!
Recife
15.3. Xerophytic Punenian Province [Puneña Xerofítica] #6768 anc
Fr 12

ia
velt
12.4
(
!

15.4. Bolivian-Tucumanan Province [Boliviana-Tucumana] Chimbote o 13.3

gua

15.5. Yungenian Province [Yungueña] 22 15

Ara
(
!
Maceió
15.5 os
16. HYPERDESERTIC TROPICAL PACIFIC Region [PACÍFICA TROPICAL HIPERDE- Di -10°
de
SÉRTICA]
-10°
dr
e 24

Ar
Aracaju

in
(
!

16.1. Hyperdesertic North Peruvian Province [Norperuana Hiperdesértica] 33

os
Callao

a
Gu

M
16.2. Hyperdesertic Tropical Chilean-Arequipan Province [Chilena-Arequipeña Tropical Lima ap 34 -30°
(!
!P

Hiperdesértica]
or é -30°
33
Bb. AUSTROAMERICAN Subkingdom [AUSTROAMERICANO] 13.8 Salvador 31
28
(
!
Cusco
(
!

15.2

M am
17. PAMPEAN Region [PAMPEANA]
17.1. Mesophytic Pampean Province [Pampeana Mesofítica] L. Titicaca 34

or é
17.2. Xerophytic Pampean Province [Pampeana Xerofítica]
Cuiabá
Brasìlia
26
Nevado de Ancohuma
(
!

18. MIDDLE CHILEAN-PATAGONIAN Region [MESOCHILENA-PATAGÓNICA] 15.1 13.6


Arequipa 6550 33 -40°
P
!
#
Goiânia
18.1. Desertic Mediterranean Chilean Province [Chilena Mediterránea Desértica] -40°
La Paz
(
! P
!

18.2. Central Chilean Province [Chilena Central]


(
!
Cochabamba

18.3. Mediterranean Andean Province [Andina Mediterránea] 13.5


(
!

Santa Cruz
(
!

18.4. Argentine Monte Province [Monte Argentino] 13.7 13.1


L. Poopó
18.5. North Patagonian Province [Patagónica Septentrional] Sucre
Abrolhos Bank 31
18.6. South Patagonian Province [Patagónica Meridional]
(
!

19. VALDIVEAN-MAGELLANIAN Region [VALDIVIANA-MAGALLÁNICA] 46 -50°


Iquique!(
19.1. Valdivean Province [Valdiviana] -50°
(
!
Belo Horizonte
19.2. Temperate Magellanian Province [Magallánica Templada]
I C
Vitória
15.4
(
!
-20° 55

P A C I F

Pa rag uay
19.3. Boreal Austromagellanian Province [Austromagallánica Boreal] -20° 14.1 ra
Juiz de Fora Campos


19.4. Insular Falkland Province [Islas Malvinas] 16.2 Pa
(
! (
!

19.5. Insular Juan Fernandez Province [Islas Juan Fernández] Campinas 55


Bc. CIRCUMANTARCTIC Subkingdom [CIRCUNANTÁRTICO]
(
!

Pilc C. Frío
(
!
Sao Paulo!( -110° -100° -90° -80° -70° -60° -50° -40° -30° -20° -10°
20. INSULAR ANTARCTIC Region [ANTÁRTICA INSULAR] om Rio de Janeiro
Tropic of C
Antofagasta
13.2
(
!
ayo
apricorn
20.1. Insular Atlantical Antarctic Province [Islas Antárticas Atlánticas] Salta

21. CONTINENTAL ANTARCTIC Region [ANTÁRTICA CONTINENTAL] 15.3


(
!
Asunción Cat. Iguazú
P
!
Curitiba!( Tropical [Tropical] Temperate [Templado]
21.1. West Antarctic Province [Antártica Occidental]
Cerro Ojos San Miguel 11. Hyperdesertic [Hiperdesértico] 31. Xeric [Xérico]
21.2. East Antarctic Province [Antártica Oriental] del Salado de Tucumán 12. Desertic [Desértico] 33. Oceanic [Oceánico]
# 6863
(
!
Resistencia Corrientes Urug
(!
! (
ua
y 13.1 13. Xeric [Xérico] 34. Hyperoceanic [Hiperoceánico]
14. Pluviseasonal [Pluviestacional] Boreal [Boreal]
14.2

Sa
region (region) province (provincia) 15. Pluvial [Pluvial] 46. Hyperoceanic [Hiperoceánico]

lad
N
O C E A
Mediterranean [Mediterráneo] Polar [Polar]

Paraná
o
18.1
Pôrto Alegre 22. Hyperdesertic-Oceanic [Hiperdesértico-Oceánico] 55. Hyperoceanic [Hiperoceánico]
24. Desertic-Oceanic [Desértico-Oceánico]
(
!

L. Mar 26. Xeric-Oceanic [Xérico-Oceánico]


Chiquita Santa Fé!( Paraná
L. de los Patos
Pelotas
18.3 (
!
-30° 28. Pluviseasonal-Oceanic [Pluviestacional-Oceánico]
Mt. Aconcagua
(
!

-30°
# 6960 17.1
Fern ández
Valparaíso
an
Arch. de Ju
(
!

Santiago
P
!

Buenos Aires Montevideo


19.5
P
!

R ío
(
! P
!
La Plata
17.2
-90° -80° -70° -60° -50° -40° 18.2 d e la Guatemala
-90° -80° -70° -60° -50° -40°
P lata G. d e
Hon du ras
Honduras
Guadeloupe

Concepción
Dominica
C. Gracias a Dios Martinique
(
! C a r i b b e a n S e a St. Lucía
Barbados
Gu St. Vincent
Mar del Plata
at em
a la Nicaragua
N O R T H

CARIBBEAN-NOVOGRANATENSEAN Superegion
I. Margarita Grenad
Tr a
en C. de la Aguja Tobago
Bahía Blanca
(
!
ch

Colora
18.4
Trinidad
10° 10°
Panamá
10° do 10°
G. o f Dar ién
id
a
s

S O U T H
(
! Canal
ér

Ba A T L A N T I C
Costa Rica M
de o Venezuela Guyana
hía
d. n
or
Valdivia Panama a n a H i g Suriname

á
u i
C

B
Neg h l

na m
G O C E A N
lan
ro a
(
!
n

Pa

a
al
ca
d
C. Orange

cen de n t
de
lf a

ta l
19.1
S. P m s French Guyana

l
Go a car a i

ien
C. al

L
C . . Oc ci
Colombia
G. San Matías

tr
Or
Puerto Montt S erra
18.5
AMAZONIAN-GUYANAN Superegion
uc umaq u
Tum

C ord
C. de San Francisc o e

C h
(
!
Marajó I.

Ch
0° Valdés Peninsula -40° 0°
Equator
Galápagos Is.
0° ub 0°
i l
Ecuador
Chiloé I.
u
-40° e R i s e
t

G. of Guayaquil

A T L A N T
Pta. Pariñas C. São Roque

I C
Pta. Negra
S a Plat. of
Chonos
e l v
NEOTROPICAL Subkingdom
Brazil

Arch ip ié la go
Borborema
TR

Comodoro Perú
San Valentín Rivadavia
OP

-10° Gulf of San Jorge -10°


(
!
-10° 4058 -10°

s
insula

nd
Argentine
IC up

Taitao Pen

la
#
Plateau of
A L er e

gh
18.6
S

Basin
Mato Grosso
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Gulf of Pen
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S O gi o

liv Bolivia
ia

an
CHACOAN-BRAZILIAN Sup
O C E A N
UT n

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Brazili
Pl
ate
Abrolh os Ba nk

I.
Wellington
HA

ra
au
u
-20° iq -20°

ei
-20° -20° I C nt
P A C I F Ma
Falkland Is.

aco
Paraguay
West Falkland da
ND

S.
os I.
Madre de Di

Ch
C. Frío

an
EAN

Gr
Chile
19.2 East Falkland

s
19.4
Str.
ellan´s

ío s
N
O C E A

Mag
-50°

eR
e
-30° -30°

E ntr
L. de los Patos
Argentina Uruguay
Santa Inés I. 19.3
-30° -30°

a s
Staten I. Ferná ndez
Arch. de Juan

d
bur
-50°
AUSTROAM ERICAN Subking

P a m p
20.1
ock
Río

gle
d

dom
e la P

C
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a l
Can

n
B ea C. Horn So uth G
eorgia
al
Can
Ba
hía S O U T H

A
Bla
n ca
G. San Matías

a
-40° C
h i
Chiloé I.
Valdés Peninsula
-40°

n i
l e
-40° -40° R i s
e
A T L A N T
w i c h I s.

ipiéla go Chon
os I C

p a t a g o
Arch
Gulf of San Jorge
ula A rg e nt in e
Taita o Penins

South Orkn
as Basin
Gul f of Pen

ey Is . 20.1 Wellin gton I.


O C E A
N

Sa
nd

Shetland Is.
West Falkland Falkland Is.
I.
Madre de Dios

20.1 20.1
East Falkland
´s St r.
e lla n
-50° M ag
Santa Inés I.
Tierra del Staten
Fuego I.
-50°
-50° -50°
rn
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Be C. Horn

ANTARCTICA
rgia
al
C an

TARC TIC Subkingd


CIRCUMAN om

w i c h I s.
21.1 Sou th Orkn
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Sa

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Shetland Is.

-60° ANTARCTICA

-110° -100° -90° -80° -70° -60° -50° -40° -30° -20° -10° -110° -100° -90° -80° -70° 19.6 -60° -50° -40° -30° -20° -10° -110° -100° -90° -80° -70° -60° -50° -40° -30° -20° -10°

Technical cartography: Miguel Álvarez, Ignacio Prieto


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