Biogeographic Map of South America. A preliminary

Transcripción

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: [email protected]
Catholic University "San Pablo". Cochabamba. Bolivia. E-mail: [email protected]
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: [email protected].
d
Botanical Garden of Valencia. Spain. E-mail: [email protected]
b
With the collaboration of: Javier Amigo (Chile and Argentina), Alindo Butzke (Brazil), Sara del Río (Argentina, Brazil and Paraguay), 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 superegions, 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 subkingdom map and a physical geography map.
Keywords: Biogeography, Map, South Americ
Introduction
Biogeography is the science which studies the distribution of species and biocoenosis on earth, as well as
the relationships between them and their causes. It takes into account the areas of taxa and syntaxa (chorology), in addition to information from other nature
sciences (geography, soil science, bioclimatology, geology, etc.), and attempts to establish a hierarchical typology of the territories on the planet, whose main
units in decreasing rank are as follows: kingdom, region, province, sector, district, area, landscape cell and
tesela (RIVAS-MARTÍNEZ & AL., 2007, 2011). Biogeography has been twinned with phytogeography due to
the value of plant species and communities in its definition and de-limittation.
Concepts on biogeographical units
The elementary biogeographic terrestrial unit or the
unit of the lowest rank is the tesela, which we define as
a geographic space with a greater or lesser extension,
ecologically homogeneous –that is to say, it has only
one single type of potential vegetation (climax) and as
a result, a single sequence of substitute communities.
The tesela and the permatesela (conceived within
the framework of dynamic-catenal phytosociology located in exceptional sites: polar, fluvial, lake and marine lands-capes, deserts, high-mountain summits, dunes
and rock formations, coastal cliffs, etc. in which the
permanent vegetation growing in these elementary spaces lacks perennial substitution units) are the only biogeographical units which can be repeated in a disjunctive manner.
The landscape cells, such as peni-plains in horst,
river valleys, lake systems, marshes, mountain summits, etc., are constituted in a broad geographic space
characterized by a series of teselas or permateselas and
their corresponding complexes, assembled by networks
of geosygmeta and geopermasygmeta based on the relief or on the soils in the territory.
The biogeographical country must be an extensive
and clearly delimited territory which possesses an abundant group of landscape cells, species, associations,
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:
[email protected].
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,
characterized by the existence of numerous differential
species and even endemic taxa in the coastal, oreadic
and interior halophilous zones, which permit their separation from the adjacent taxa; it also comprises an independent unit through its associations, series, geoseries and geoclinoseries which are absent in nearby districts.
The sector is a grouping of districts with a largescale geographic entity, which possesses its own endemic taxa, associations and vegetation series, as well as
original topographical and geoclinosequential geoseries
which are generally due to the existence of exclusive
climatophilous, permanent and subserial communities,
as well as to paleoclimates or former migratory routes.
The province is a vast geographic territory which,
as well as possessing a large number of endemisms and
differential species (its own subelement) has particular
macroseries and geomacroseries; a particular altitudinal
zonation in the vegetation is also characteristic of each
biogeographical province (exclusive geoclinoseries).
The region is a very extensive territory, formed by a
group of biogeographical provinces which has a flora
or regional floristic element with species, genera or
even endemic families; in addition it has its own particular megaseries, geomegaseries and geomegapermaseries and in consequence, its own bioclimatic belts (RIVAS-MARTÍNEZ, 2005).
Finally, the kingdom is the supreme unit of biogeography, generally pluricontinental and multinsular,
which in addition to taxonomic and ecosystematic considerations, addresses the origins of the flora and fauna,
the formation of the great continents, orogenies and climate in the present and past.
As is by now traditional in this science, the denominations of the biogeographic units –both the primary
and the auxiliary units (from the subregion to the
area)– are given as names and place names based on
known geographical, orographic and historical designations which are considered to conicide with, inform or
represent the area they are intended to represent.
Orthographically, all the units are considered to be proper names identifying the place; the name is formed by
juxtaposing two geographical nouns, joined by means
of a hyphen, maintaining the initial capital in each due
to their condition of proper noun.
It must be emphasized that the biogeographical
units can only be accurately delimited through their
diagnosis and through the corresponding maps. All the
territories –except for the tesela and permatesela– must
be contiguous by land, lake or sea routes, and include
all the orographic accidents and lithological diversity
which may exist in the area. Sometimes in the biogeographical territories as a whole there are introgressions
by other adjacent territories, and these islands frequently occur in regions with a varied lithology or in areas
near regional or provincial boudaries. Their possible
typological independence, always of a lower rank than
the area into which they introgress, depends on their
originality, floristic richness and phytocoenotics, as
well as on their surface area.
One of the criteria traditionally used for recognising
and delimiting biogeographic units with their own
entity is the distinguishing and mapping of taxa (families, genera, species and subspecies) whose territorial
distribution is closely restricted to one particular area.
These taxa are termed endemisms. Endemisms have
been successfully used to define and delimit the chorological or biogeographical units (provinces, sectors), as
they form part of the phytogeographical subelement
which characterises them. Moreover, the endemisms
(taxa or syntaxa) which have a greater area or are regional, and those which for migratory reasons are dispersed across various biogeographical regions, constitute the phytogeographical element or geoelement.
Concepts on vegetation series and lanscape phytosociology
Nowadays, the development of dynamic-catenal
phytosociology and the syntaxonomic knowledge of
broad territories on the Earth, as well as the cartographic delimitation of vegetation series, geoseries and
geopermaseries, when available, have become the essential criteria for defining biogeographical units, in
addition to suitably compiled and mapped bioclimatic
and soil factors.
The vegetation series, also termed sygmetum, expresses the whole set of plant communities or stages
which can be found within similar teselar spaces as a
result of the succession process, and includes both the
representative association of the mature stage or series
head, which is used as a nomenclatural reference, and
the initial or subserial associations that may replace it.
Based on this concept, the vegetation series or sygmetum represents the basic unit or essential model of dynamic phytosociology. Distinctions can also be made
between climatophilic, edaphoxerophilic, temporihygrophilic and edaphohygrophilic series. Climatophilic
or zonal series are located on mature soils according to
the mesoclimate, and only receive rainwater: mesophytic, submesophytic and subxerophytic; the temporihygrophilic series, included among the climatophilic, are
those which have additional water contribution due to
their topographical circumstances, and they thus develop on flooded or very wet soils throughout part of the
year, and –at least during the summer or dry period–
the soil horizons are well-drained and aerated. Finally
the edaphoxerophilic series are found in particularly
dry or xerophytic soils or biotopes such as lithosols,
arenosols, very windy sites, steep slopes, crests, ledges,
etc.; and the edaphohygrophilic series grow on particularly wet soils and biotopes such as fluvisols, halosols,
histosols, etc., and are found on river beds, marsh areas, salt flats, peat bogs, etc.
The vegetation geoseries or geosygmetum is the basic unit of dynamic-catenal phytosociology. It corresponds to a catena of vegetation series which is found
around a given bioclimatic belt and biogeographic territory in the heart of the universal crest-slope-valley
model. This topographic framework makes it possible
to distinguish the three geomorphological aspects of
any complete catena where the vegetation series constituting the geosygmetum are located in zones; the edaphoxerophilic series and geoseries (hyperxerophilic
and xerophilic) are located in the driest sites (crests, es-
Biogeographic Map of South America. A preliminary survey
carpments, lithosols, etc.); the climatophilic and temporigrophilic series and geoseries are located on slopes
and foothills where greater humidity is contributed by
rainfall; and the edaphohygrophilic series and geoseries
are found in the valleys and watercourses (fluvial, lake
and watercourses), among which the river fractogeosygmetum is of great important to plant landscape science due to its extrazonality, and also, in combination
with the edaphoxerophilic and climatophilous sygmeta
and geosygmeta, to the definition and structuring of regional and global biogeography.
The vegetation geopermaseries, also known as geopermasygmeta, is the catenal expression of a set of neighboring permaseries or permasygmeta, delimited by
changing topographic or soil situations. These are conditioned by conditions of extreme climate (high mountains and polar areas) and exceptional microtopographic and soil conditions (walls, rock formations, marine cliffs, salt flats, etc.) which give rise to a large number of neighboring ecological residences populated by
diverse permanent perennial plant communities (continuous vegetation permaseries) with an absence of
non-nitrophilous serial perennial communities which
appear to have reached their equilibrium. The most favorable sites for the existence of geopermaseries or
geopermasygmeta, as well as sites corresponding to
permanent types of vegetation in extreme high-mountain and polar region bioclimates, are ledges, rock crevasses, cliffs and coastal rock formations bathed by sea
waters, peat bogs, wind drifts, mobile sand dunes, lake
shores, streams etc.
Studies performed and geobotanical sources
Salvador Rivas-Martínez and Manuel Costa, Professor of Botany at the University of Valencia, worked
together at the Faculty of Biology at the Complutense
University and at the Royal Botanical Gardens in Madrid in the 1970s, where they designed a long-term global geobotanical program to further the syntaxonomical
study of the Earth following the European phytosociological methodology of BRAUN-BLANQUET (1964). To
this method they incorporated the new synphytosociological analyses proposed by RIVAS-MARTÍNEZ (1976),
and shortly afterwards by GEHÚ & RIVAS-MARTÍNEZ
(1982), as well as recent bioclimatic and biogeographical approaches which were being successfully tested in
Europe and North Africa, and in whose scientific and
methodological debates with Tüxen, Gehú, Bolós, Quézel and Tahktajan, they played an active role.
Primo Yúfera, at that time general secretary of the
Higher Council for Scientific Investigation (CSIC) in
the mid-1970s, convened various directors of the Institutes in the CSIC, including Salvador Rivas-Martínez,
to present research proposals which could be used in
due course for the scientific commemoration of the
fifth centenary of the discovery of America in 1992.
They wasted no time in adapting part of their global
geobotanical program to the wildest possible subject
they could imagine, in keeping with their dual facets of
botanists and climbers at that time: the “Phytosociological and bioclimatological study of the Andes in the
Viceroyalties of Peru and Nueva Granada”. The CSIC,
23
and particularly what was then the Institute of Hispanic
Culture, sparingly but reliably financed the project in
the early years (1977-1982), thus providing a stimulus
and a springboard for many. Rivas-Martínez has since
worked to share what he colloquially termed the
“American adventure” by promoting and organizing,
together with his good friend Manuel Costa, countless
trips and studies in North America, Africa, China, Australia and Tasmania. Moreover Manuel Costa has traveled independently and has an in-depth knowledge of
Central America, Mexico, the east of North America
and Tibet. His last major project, still incomplete after
a dozen years, is currently the geobotanical study of
Venezuela, where, with the help of a large group of
first-rate young professional Venezuelan botanists and
soil scientists, he is directing a series of doctoral theses
on the ground, covering large parts of the territory, four
of which have already been judged “cum laude” at the
University of Valencia. On several occasions accompanying Manuel Costa and his team, Rivas-Martínez has
had the opportunity to increase his knowledge of the
vegetation and the bioclimate of Venezuela and, above
all, to study the regions of Alto Orinoco, La Guayana
and the Gran Sabana.
Gonzalo Navarro, Ph.D., a native of Madrid, now
lives in Cochabamba, after completing his education
and writing an extensive doctoral thesis entitled “Flora
and vegetation of the Sistema Ibérico in Soria”, which
was directed by Rivas-Martínez. Shortly after, as a result of an exploratory trip through the Andes, Gonzalo
Navarro took the irrevocable decision to live and research in Bolivia for most of the year. He thus applied
for a leave of absence from his position as permanent
teacher in the institute of secondary education in Guadalajara (Spain), which he had won through public examination. A little later he continued his connections as
a part-time associate professor in the Department of
Botany at the Faculty of Pharmacy of the Complutense
University, at that time headed by Rivas-Martínez, with
the undertaking that during his winter stay in Madrid
he would teach the optional subject “Biogeography of
South America”, conduct research into Andean, Chacoan and Amazonian ecosystems, and that at the University of Cochabamba, with which there was an interuniversity agreement, he would train young students
and graduates in geobotany so that under his direction
they could begin studying their doctoral thesis on Bolivian subjects, and that thanks to the Mutis grant system, they could spend a long period of training in Madrid studying the subjects for their doctorate. Gonzalo
Navarro amply and generously fulfilled this commitment, which continued successfully maintained for almost a decade. Of the six Mutis grant-holders who were in Madrid, two are already doctors. Unfortunately in
2004, after the requisite yearly democratic and secret
departmental vote for the renewal of his post as associate professor, he found himself to be one vote short,
and was rejected. Fortunately we his friends still enjoy
the privilege of continuing to work and collaborate
with him.
In the last 20 years, Gonzalo Navarro has conducted a continuous, profound and methodical investigation into the Bolivian Andes, and the areas of Beni,
24
Chaco, Cerrado, Yungas and Amazonia, as well as in a
more general manner in the United States, Mexico and
over almost all of South America from Patagonia to the
Venezuelan Caribbean. On many expeditions and journeys he has used his own resources, and on others, resources obtained in projects and competitions proposed
by organizations such as TNC, WCS, WWF, etc. As a
result of this research, as well as multiple reports and
publications, we should also mention the unrivalled
“Vegetation of Bolivia” published in 2002, in the book
“Ecological geography of Bolivia” [ISBN 99905-0225-0] and the astounding “Vegetation map of Bolivia
at a scale of 1:250.000” published by Nature Conservancy [ISBN 978-99954-0-168-9]. For all these reasons he has achieved the distinction on his own merits
of becoming the maximum geobotanical authority on
Bolivia and a key reference in the neotropics of South
America.
Ángel Penas, professor at the University of León,
began his partnership with Rivas-Martínez 30 years
ago in an unforgettable geobotanical research project in
the Cordillera Cantábrica mountains, conducted with
Tomás E. Díaz, José Antonio F. Prieto and Javier Loidi, and which culminated with the publication of a
book on the vegetation in the Picos de Europa mountains. Since then their professional relationship and friendship has continued to grow, particularly through the
publications in Itinera Geobotánica and the Habitats
projects. However their main point in common is their
mutual interest in the study of the vegetation of America and Africa. In January 1996 they successfully investigated, in the company of Leonard Llorens, the southeastern tropical and oreadic temperate vegetation of
South Africa; in America they undertook two major
geobotanical expeditions, one in September 1994, with
Gonzalo Navarro and Francisco Amich, in which they
toured the Pampa, Monte Argentino and Patagonia making inventories, concluding in the Mediterranean Andean forests of Austrocedrus chilensis and in the Valdivian temperate forests of Nothofagus; the second was a
fruitful and extensive geobotanical transect in April
1995, in the company of Gonzalo Navarro, Federico
Fernández and Daniel Sánchez Mata, where they successfully studied the tropical south of the United States: Texas, Tamaulipas, Chihuahua and Sonora, as well
as the continental Mediterranean territories of the Great
Basin and the oceanic Mediterranean territories of California. However Ángel Penas most important research
activity in South America, apart from his geobotanical
excursions to Colombia, Venezuela, Peru, Chile and
Argentina, was in Brazil, where only between 1992 and
2006 he personally directed six doctoral theses on the
region of Parana, judged cum laude in the University of
León, and still found time to travel extensively and study numerous aspects of Amazonia, La Catinga, Cerrado Oriental, Mata Atlántica and the mesophytic Pampa.
Salvador Rivas-Martínez began his study of tropical
and Andean flora and vegetation in 1961 on the occasion of a major mountaineering expedition to the Andes in Peru (Apolobamba massif and Cordillera Blanca). Between 1965 and 1976 he continued these studies
intermittently in Colombia, Ecuador, Bolivia, Chile and
Argentina, taking advantage of other mountaineering
expeditions and various botanical and professional expeditions. However, it was between 1977 and 1990,
owing to several well-organized projects financed by
Spain, when he was able uninterruptedly to develop the
“Phytosociological and bioclimatic study of the Andes
of the Viceroyalties of Perú and Nueva Granada”,
which from 1985 became known as “Bioclimatic and
cultivation belts of Perú” thanks to another better financed project. During those 13 years he made more
than 20 botanical expeditions around a large part of the
Andes, Amazonia, Chaco, the Pacific coasts and deserts, the Pampa, Patagonia and Valdivia, where he
took 3,000 phytosociological inventories, compiled climate data for more than 2,000 South American weather
stations, and mapped the vegetation and bioclimate of
most of the territories accessible by road. Manuel Costa, as well as many of his disciples, colleagues from the
Complutense University such as Carlos Arnaiz, Paloma
Cantó, Ana Crespo, Antonio Galán, Javier Loidi, Gonzalo Navarro and Conchita Sáenz, also took part in some of these journeys. From the outset he was lucky
enough on almost all these campaigns to enlist the collaboration and valuable help of Oscar Tovar, the eminent Peruvian agrostologist and botanist, Professor of
Botany at the University of San Marcos in Lima, as
well as the support, teachings and opinions of Stephan
Beck, Ángel Cabrera, José Cuatrecasas, Ramón Ferreira, Raúl Lara, Máximo Lieberman, Eduardo Martínez
Carretero, Eric Oberdorfer, Salvador Rivas Goday, Fidel Roig and Paul Seibert. Unfortunately, two years
before the official conclusion of this last project, highlevel political differences between Spain and Peru, unrelated to any personal considerations, led to the abrupt
termination of the agreement and to the project’s financing. As it was not possible to redirect the South American research program to Spain, it had to be postponed. In view of these difficulties he took the decision
to spend two sabbatical years in the United States, in
the Missouri Botanical Garden, in order to devote himself completely to the study of the vegetation of North
America and finally to formulate the bioclimatology at
the global level. Luck would have it that Gonzalo Navarro, a young doctor with considerable training in taxonomy and geobotany, had moved to Bolivia and with
great enthusiasm and effort began to study in great
detail and with marked success the oreadic and flatlandic vegetation of Bolivia. This circumstances led Rivas
Martínez in 1994 to return to South America to work
with Navarro, who had by now become a considerable
expert and specialist, and together they designed the
biogeographical synthesis of South America (1994),
which has today, after many years, been improved and
updated with the help and mutual responsibility of Ángel Penas and Manuel Costa, as well as with the collaboration and advice of other botanists with extensive
experience in the territory: J. Amigo, A. Butzke, S. del
Río, A. Galán, J. Guevara, J. Izco, E. Martínez Carretero, O. Rangel, F. Roig (†), D. Sánchez-Mata, L.G. Sancho, P. Soriano and O. Tovar (†); as well as with the
experience of S. Rivas Sáenz, expert and co-author of
the program of computerized bioclimatic cartography.
At the same time a detailed review was made of the
bibliography which has enabled us to compare our ob-
servations with those of other authors, of both general
and specific works, practically all of which are listed in
the bibliographical references.
Among the authors with a general character, there
are some who must necessarily be mentioned due to
their importance from both the methodological standpoint and for the proposals they formulate in relation to
the biogeography of South America: ALCARAZ, F.
(1999), ÁLVAREZ, M., RAMÍREZ, C.& DEIL, U. (2008),
AUBRÉVILLE, A. (1970), BRAUN-BLANQUET, J. (1964),
CABRERA Á. L. & WILLINK A. (1973), CHEVALIER, A.
& EMBERGER, L. (1937), DARLINGTON, P.J. (1957),
DIELS, L. & MATTICK, F. (1958), DRUDE, O. (1890),
ENGLER, A. (1924), GENTRY, A. (1982), GOOD, R.
(1974), GRISEBACH, A. (1872), HAYEK, A. (1926),
HUBER, O. & RIINA, R. (1997, 2003), HUECK, K. &
SEIBERT, P. (1972, 1981), JOSSE, C., CUESTA, F., NAVARRO, G., BARRENA, V., CABRERA, E., CHACÓN-MORENO, E., FERREIRA, W., PERALVO, M., SAITO, J. & TOVAR, A. (2008), JOSSE, C., NAVARRO, G., COMER, P.,
EVANS, R., FABER-LAGENDOEN, D., FELLOWS, M., KITTEL, G., MENARD, S., PYNE, M., REID, M., SCHULZ,
K., SNOW, K. & TEAGUE, J. (2003), JOSSE, C., NAVARRO, G., ENCARNACIÓN, F., TOVAR, A., COMER, P., FERREIRA, W., RODRÍGUEZ, F., SAITO, J., SANJURJO, J.,
DYSON, J., RUBIN DE CELIS, E., ZÁRATE, R., CHANG, J.,
AHUITE, M., VARGAS, C., PAREDES, F., CASTRO, W.,
MACO, J. & REÁTEGUI, F. (2007), LABUNTSOVA, M.A.
(1969), LAVRENKO, E. M. (1964), LLORENTE BOUSQUETS, J. & MORRONE, J. J. (EDS.) (2001), LUTEYN, J.
(1999), MORELLO, J. (1958), MORRONE, J. J. (2000,
2001A, 2001B), OZENDA, P. (1964), POSADAS, P. E.; ESTÉVEZ, J. M. & MORRONE, J. J. (1997), PRADO, D.E. &.
GIBBS, P. E. (1993), RAVEN, P.H. (1963), RAVEN, P.H.
& AXELROD, D.J. (1974), RICARDI, M., GAVIRIA, J. &
ESTRADA, J. (2001), RIKLI, M. (1913, 1934), RIVASMARTÍNEZ, S. (1976, 1994, 1997, 2001, 2004, 2005),
RIVAS-MARTÍNEZ, S & RIVAS-SAÉNZ (2009), RIVASMARTÍNEZ, S. & COLS. (2007, 2011), RIVAS-MARTÍNEZ,
S. & NAVARRO, G. (1998), RIVAS-MARTÍNEZ, S., RIVAS-SÁENZ, S., PENAS, A., NAVARRO, G. & COSTA, M.
& COLS. (2011), RIVAS-MARTÍNEZ, S., SÁNCHEZ-MATA, D. & COSTA, M. (1999), RIVAS-MARTÍNEZ, S. &
TOVAR O. (1983), SCHMITHÜSEN, J. (1961), SCHWABE,
G. H. (1968), SMITH, A.C. & JOHNSTON, I.M. (1945),
TAKHTAJAN, A. (1988), THORNE, R.F. (1963), DEIL, U.,
ÁLVAREZ, M., BAUER, E.V. & RAMÍREZ, C. (2011),
UDVARDY, M.D.F. (1975), WALTER, H. (1898) and
WALTER, H. & STRAKA, H. (1970).
We have also based this work on the studies carried
out in territories which –although extensive– are more
specific than the previous ones, by different authors
which we mention below, for their work done in various South American countries. From Venezuela and
the Guayanas we should highlight the works of
ATAROFF, M. & SARMIENTO, L. (2003), AYMARD, G.
(2003), CASTROVIEJO, S. & LÓPEZ, G. (1985), COSTA,
M.; CEGARRA, J.; LUGO, L.; GUEVARA J.R.; CARRERO,
O.; LOZADA, J. & SORIANO, P. (2008), COSTA, M.; CEGARRA, A.; LUGO, L.; LOZADA, J.; GUEVARA, J. & SORIANO, P. (2007), CUELLO, N. & CLEEF. A.M. (2009 a,
b), DUNO DE ESTEFANO, R., AYMARD, G. & HUBER, O.
(2007), ESTRADA SÁNCHEZ, J.C. (2003), GALÁN DE
25
MERA, A. (2007), GALÁN DE MERA, A., GONZÁLEZ, A.,
MORALES, R., OLTRA, B. & VICENTE ORELLANA, J.A.
(2006), GUEVARA, J. R., CARRERO, O., HERNÁNDEZ, C.
& COSTA, M. (2007), HUBER, O. & ALARCÓN, C.
(1988), HUECK, K. (1960), LOZADA, J, GUEVARA, J.R.,
SORIANO, P. & COSTA, M. (2006), MAGUIRE, B. (1970,
1972, 1979), MINISTERIO DEL AMBIENTE Y DE LOS RECURSOS NATURALES (1985), MONASTERIO, M. & REYES, S. (1980), PITTIER, H. (1920, 1935), RÖHL, E.
(1946), SARMIENTO, G. & MONASTERIO, M. (1969),
STEYERMARK, J. A. (1974, 1979) and SUSACH CAMPALANS, F. (1989).
In Colombia those of CLEEF, A. M. (1979, 1981)),
CLEEF, A.M. & RANGEL, J.O. (1986), CLEEF, A.M.,
RANGEL, J.O. & ARELLANO, H. (2008), CLEEF, A.M.;
RANGEL, J.O. & SALAMANCA, S. (1983, 2003), CORTÉS
B. R. & FRANCO R., P. (1997), CUATRECASAS, J. (1934,
1958), DUGAND, A. (1970), HAMMEN, T. VAN DER,
RANGEL CH., J.O. & CLEEF, A. M. (EDS.). (2005),
KLOOSTERMAN, E.H.; CLEEF, A.M. & SALAMANCA, S.
(2003), PINTO-ZÁRATE, J.H. & RANGEL-CH., J.O. (2010
a, b), RANGEL-CH., J.O. (2004, 2008), RANGEL-CH.,
J.O. (ED.) (1995, 1997, 2000, 2007, 2008, 2009, 2010 a,
b), RANGEL-CH., J.O. & AGUIRRE, J. (1983), RANGELCH., J.O. & ARELLANO-P., H. (2009), RANGEL-CH.,
J.O. & ARIZA, N.C. (2000), RANGEL-CH., J.O., CLEEF,
A.M. & ARELLANO, H. (2008), RANGEL-CH., J.O.,
CLEEF, A.M. & SALAMANCA, S. (2005), RANGEL-CH.,
J.O., GARAY-P., H. & AVELLA, A. (2010), RANGELCH., J.O., SÁNCHEZ, D. & ARIZA, N.C. (2005), RIVERA
DÍAZ, O. & FERNÁNDEZ-ALONSO, J. L. (2003), SALAMANCA S., CLEEF, A.M. & RANGEL, J.O. (2003), SÁNCHEZ, R. & RANGEL, J.O. (1990) and WITTE, H.J.J.
(1995).
In Peru and Bolivia we should mention the works
of DE LA BARRA, N. (2003), GALÁN DE MERA, A.
(1995, 1999), GALÁN DE MERA, A., BALDEÓN, S., BELTRÁN, H., BENAVENTE, M. & GÓMEZ, J. (2004), GALÁN
DE MERA, A., CÁCERES, C. & GONZÁLEZ, A. (2003),
GALÁN DE MERA, A., ROSA, M.V. & CÁCERES, C.
(2002), GALÁN DE MERA, A. & VICENTE ORELLANA,
J.A (1996, 2006), GUTTE, P. (1980, 1986, 1988), MOLINA, J.A., NAVARRO, G., DE LA BARRA, N. &
LUMBRERAS, A. (2007).NAVARRO, G. (1993, 1997,
2003), NAVARRO, G. & FERREIRA, W. (2007, 2009),
NAVARRO, G. & MALDONADO, M. (2002), NAVARRO,
G., MOLINA, J. A. & DE LA BARRA, N. (2005), RAMÍREZ, C. & BECK, S. (1981), RIVAS-MARTÍNEZ, S. &
TOVAR, O. (1982), SEIBERT, P. & MENHOFER, X. (1991,
1992, 1993), WEBERBAUER, A. (1945) and WILLIAMS,
L. (1945).
In Ecuador the works by ACOSTA-SOLIS, M. (1966,
1984), AGUIRRE, Z., COOMBES, L. & RAMSAY, P. M.
(2001), MADSEN, J.E., COTTON, E. & BALSLEV, H.
(EDS.) (2002), CAÑADAS, L. (1983), GREHAN, J. R.
2001, IZCO, J., PULGAR, I., AGUIRRE, Z. & SANTÍN, F.
(2004), JOHNSTON, M.P. & P.H. RAVEN (1973), MOSCOL OLIVERA, MARCELA C. & CLEEF, A. M. (2009),
PÚLGAR, I., IZCO, J. & JADÁN, O. (2010), QUINTANILLA,
V.G. (1983), SIERRA, R. (ED.) (1999), SIERRA, R., CICERÓN, C., PALACIOS, W. & VALENCIA, R. (1999), TERÁN,
E. (1979), TERNEUS, E. (2002) and ULLOA, C. & JØRGENSEN, P.M. (1993).
26
In Chile the works carried out by AMIGO J. (2009),
AMIGO, J., IZCO, J. &.RODRÍGUEZ-GUITIÁN, M.A.
(2007), AMIGO J. & RAMÍREZ, C. (1998), AMIGO, J.,
RAMÍREZ, C. & GARCÍA QUINTANILLA, L. (2004, 2007),
AMIGO, J., SAN MARTÍN, J. & GARCÍA QUINTANILLA, L.
(2000), ETAYO, J. & SANCHO, L.G. (2008), HILDEBRAND-VOGEL, R. (1984), HILDEBRAND-VOGEL, R.,
GODOY, R. & VOGEL, A. (1990), LUEBERT, F. & GAJARDO, R. (2005), LUEBERT, F. & P PLISCOFF. (2006),
MÉNDEZ, E. (2007), MÉNDEZ, E. & AMBROSETTI, A.J.
(1985), NAVARRO, G. & RIVAS-MARTÍNEZ, S. (2005),
OBERDORFER, E. (1960), PINTO, R. & LUEBERT, F.
(2009), RAMÍREZ, C., SAN MARTÍN, C., CONTRERAS, D.
& SAN MARTÍN, J. (1994) and RUTHSATZ, B. (1995).
In Argentina, Paraguay, Uruguay and Brasil, the
works published by BOLÓS, O., CERVI, A.C. & HATSCHBACH, G. (1991), BUTZKE, A. (1997), CABIDO, M.
(1985), CABIDO, M. & ACOSTA, A. (1986), CABIDO, M.,
ACOSTA, A. & DÍAZ, S. (1990), CABRERA, Á. L. (1971),
CABRERA, Á. L. (1976), DIESEL, S. (2005), EITEN, G.
(1972, 1983), ESKUCHE, U. (1984, 2005), FAGGI, A.M.
(1985), FIASCHI, P & J.R. PIRANI. (2009), FONTANA,
S.L. (2005), GALÁN DE MERA, A. & NAVARRO G.
(1992), GANDULLO, R. & FAGGI, A. M. (2003, 2005),
GANDULLO, R. & SCHMID, P. (2001), GIBBS, P. E.,
LEITAO FILHO, H. & SHEPHERD, G. (1983), KEGLER, A.
(2005), KEGLER, A., DIESEL, S., WASUM, R.A., HERRERO, L., DEL RÍO, S. & PENAS, A. (2010), LEÓN, R.J.C. &
BURKART, S.E. (1988), LEWIS, J:P:, COLLANTES, M:B:,
PIRE, E.F., CARNEVALE, N.J., BOCCANELLI, S.I., STOFELLA, S.L. & PRADO, D.E. (1985), MARTÍNEZ CARRETERO, E. (1993, 1995, 2000, 2001, 2006), MÉNDEZ, E., E.
MARTÍNEZ CARRETERO & I. PERALTA, I. (2006), NAVARRO, G., MOLINA, J. A. & PÉREZ DE MOLAS, L. (2006),
PRANCE, G. T. (1977, 1978, 1979), RATTER, J. A., LEITAO-FILHO, H. DE F., ARGENT, G., GIBBS, P. E., SEMIR,
J., SHEPHERD, G. & TAMASHIRO, J. (1988), REITZ, P. R.
(1961), RIZZINI, C. T. (1979), ROIG, F.A. (1972, 1998),
ROIG, F.A., ANCHORENA, J., DOLLENZ, O., FAGGI, A.M.
& MÉNDEZ, E. (1985), ROIG, F.A., DOLLENZ, O., &
MÉNDEZ, E. (1985), ROIG, F.A.& FAGGI, A. (1985),
RUTHSATZ, B. (1977), SAMPAIO, A. J. DE (1934), SCUR
L. (2005), STUTZ DE ORTEGA, L. C. (1983, 1984, 1986,
1987, 1990), VELOSO H. P. (1946, 1948), VELOSO H. P.
& KLEIN, R. M. (1957, 1959) and WASUM, R. A.
(2005).
And finally we should indicate that we have analysed proposals for territories other than South America
but which bear a relation with South America for floristic or other reasons, among which we should mention: CANO CARMONA , E., VELOZ, A. & CANO ORTÍZ,
A. (2010), CONTRERAS MEDINA., R.; LUNA VEGA, I. &
MORRONE, J. J. (1999), EMMERICH, K. H. (1988), GALÁN DE MERA, A. (2005), LAUER, W. (1968), LUNA VEGA, I.; MORRONE, J. J.; ALCÁNTARA AYALA, O. & ESPINOSA ORGANISTA, D. (2001), MATTICK, F. (1964),
DEIL, U. (1994, 1999). We have also particularly taken
into account the bioclimatic typology of RIVAS-MARTÍNEZ, S. (2004), RIVAS-MARTÍNEZ, S. & RIVAS SÁENZ,
S. (2009) and RIVAS-MARTÍNEZ, S., RIVAS SÁENZ, S. &
PENAS, A. (2011).
Results
A. Biogeogeographical typology of South America
up to provincial level
In South America and from Honduras to the Antarctic Peninsula we recognise: 1 kingdom, 3 subkingdoms, 4 superegions, 13 regions and 53 biogeographic
provinces (see Anex Map).
The biogeographical denominations, as is traditional, are place names and adjectives which express
territories, history or toponyms. Generally these denominations are given in Spanish and translated into
English.
B. NEOTROPICAL-AUSTROAMERICAN Kingdom
[Reino NEOTROPICAL-AUSTROAMERICANO]
Ba. NEOTROPICAL Subkingdom
[Subreino NEOTROPICAL]
Bab. CARIBBEAN-NEOGRANADIAN Superegion
[Superregión CARIBEÑA-NEOGRANADINA]
9. CARIBBEAN-MESOAMERICAN Region]
[Región CARIBEÑA-MESOAMERICANA]
9.3. Lesser Antillean Province
[Provincia Antillana Menor]
9.5. Chiapan-Honduran Province]
[Provincia Chiapaneca-Hondureña]
9.6. Panamanian-Costa Rican Province
[Provincia Panameña-Costarricense]
10. NEOGRANADIAN Region
[Región NEOGRANADINA]
10.1. Guajiran-Caribbean Province
[Provincia Guajireña-Caribeña]
10.2. Llaneran Province
[Provincia Llanera]
10.3. Colombian Andean Province
[Provincia Andina Colombiana]
10.4. Cordobesa-Lower Magdalena Province
[Provincia Cordobesa-Bajomagdalena]
10.5. Colombian Pacific Province
[Provincia Pacífico Colombiana]
10.6. Guayaquilian-Ecuadorean Province
[Provincia Guayaquileña-Ecuatoriana]
10.7. Insular Galapagos Province
[Provincia Islas Galápagos]
Bac. AMAZONIAN-GUYANAN Superegion
[Superregión AMAZÓNICA-GUAYANENSE]
11. GUYANAN-ORINOQUIAN Region
[Región GUAYANA-ORINOQUENSE]
11.1. Guyanan Province
[Provincia Guayanense]
11.2. Deltaic Orinoquian Province
[Provincia Orinoquense Deltaica]
11.3. Guaviarean-Orinoquian Province
[Provincia Guaviareña-Orinoquense]
11.4. Tepuyan Province
[Provincia Tepuyana]
11.5. Guyanese Brazilian Province
[Provincia Brasileña Guayanense]
12. AMAZONIAN Region
[Región AMAZÓNICA]
12.1. West Amazonian Province
[Provincia Amazónica Occidental]
12.2. North Amazonian Province
[Provincia Amazónica Septentrional]
12.3. Deltaic Amazonian Province
[Provincia Amazónica Deltaica]
12.4. Southwest Amazonian Province
[Provincia Amazónica Suroccidental]
12.5. Central Amazonian Province
[Provincia Amazónica Central]
Bad. CHACOAN-BRAZILIAN Superegion
[Superregión CHAQUEÑA-BRASILEÑA]
13. BRAZILIAN-PARANENSE Region
[Región BRASILEÑA-PARANAENSE]
13.1. Brazilian Atlantic Province
[Provincia Atlántica Brasileña]
13.2. Paranense Province
[Provincia Paranaense]
13.3. Catingan Province
[Provincia Catinguense]
13.4. Tocantins Province
[Provincia Tocantinense]
13.5. East Cerrado Province
[Provincia Cerradense Oriental]
13.6. West Cerrado Province
[Provincia Cerradense Occidental]
13.7. Pantanalian Province
[Provincia Pantanalense]
13.8. Benian Province
[Provincia Beniana]
14. CHACOAN Region
[Región CHAQUEÑA]
14.1. North Chacoan Province
[Provincia Chaqueña Septentrional]
14.2. South Chacoan Province
[Provincia Chaqueña Meridional]
Bae. TROPICAL SOUTH ANDEAN Superegion
[Superregión SURANDINA TROPICAL]
15. TROPICAL SOUTH ANDEAN Region
[Región SURANDINA TROPICAL]
15.1. Desertic Peruvian-Ecuadorean Province
[Provincia Peruana-Ecuatoriana Desértica]
15.2. Mesophytic Punenian Province
[Provincia Puneña Mesofítica]
15.3. Xerophytic Punenian Province
[Provincia Puneña Xerofítica]
15.4. Bolivian-Tucumanan Province
[Provincia Boliviana-Tucumana]
15.5. Yungenian Province
[Provincia Yungueña]
16. HYPERDESERTIC TROPICAL PACIFIC Region
[Región PACÍFICA TROPICAL HIPERDESÉRTICA]
16.1. Hyperdesertic North Peruvian Province
[Provincia Norperuana Hiperdesértica]
16.2. Hyperdesertic Tropical Chilean-Arequipan
Province
[Provincia Chilena-Arequipeña Tropical
Hiperdesértica]
Bb. AUSTROAMERICAN Subkingdom
[Subreino AUSTROAMERICANO]
17. PAMPEAN Region
[Región PAMPEANA]
17.1. Mesophytic Pampean Province
[Provincia Pampeana Mesofítica]
17.2. Xerophytic Pampean Province
[Provincia Pampeana Xerofítica]
27
18. MIDDLE CHILEAN-PATAGONIAN Region
[Región MESOCHILENA-PATAGÓNICA]
18.1. Desertic Mediterranean Chilean Province
[Provincia Chilena Mediterránea Desértica]
18.2. Central Chilean Province
[Provincia Chilena Central]
18.3. Mediterranean Andean Province
[Provincia Andina Mediterránea]
18.4. Argentine Monte Province
[Provincia Monte Argentino]
18.5. North Patagonian Province
[Provincia Patagónica Septentrional]
18.6. South Patagonian Province
[Provincia Patagónica Meridional]
19. VALDIVEAN-MAGELLANIAN Region
[Región VALDIVIANA-MAGALLÁNICA]
19.1. Valdivean Province
[Provincia Valdiviana]
19.2. Temperate Magellanian Province
[Provincia Magallánica Templada]
19.3. Boreal Austromagellanian Province
[Provincia Austromagallánica Boreal]
19.4. Insular Falkland Province
[Provincia Islas Malvinas]
19.5. Insular Juan Fernandez Province
[Provincia Islas Juan Fernández]
Bc. CIRCUMANTARCTIC Subkingdom
[Subreino CIRCUNANTÁRTICO]
20. INSULAR ANTARCTIC Region
[Región ANTÁRTICA INSULAR]
20.1. Insular Atlantical Antarctic Province
[Provincia Islas Antárticas Atlánticas]
21. CONTINENTAL ANTARCTIC Region
[Región ANTÁRTICA CONTINENTAL]
21.1. West Antarctic Province
[Provincia Antártica Occidental]
21.2. East Antarctic Province
[Provincia Antártica Oriental]
B. Descriptions of biogeographic units
Amazonian: South-American equatorial and Atlantic
southern eutropical biogeographic region (12), pluvial and tropical mesophytic infra-lower thermotropical bioclimates and with few rare exceptions submesophytic, with varzeal compensation, on soils and
waters enriched with ions and nutrients. To the north
it borders on the hyperoligotrophic soils and waters
with Guyanan-Orinoquian Region; to the south on
the infra-thermotropical pluviseasonal lower mesophytic and submesophytic border with the BrazilianParanense Region and, at the headwaters of the Andean mountains with the yungas of the Tropical
South Andean Region which have a height of between 500-1000 m. The Amazonian Region comprises five biogeographic provinces: 12.1. West Amazonian, 12.2. North Amazonian, 12.3. Deltaic Amazonian, 12.4. Southwest Amazonian, 12.5 and Central
Amazonian. [Región Amazónica]
Amazonian-Guyanan: South-American biogeographical superegion (Bac), with tropical macrobioclimate formed by the Guyanan-Orinoquian and Amazonian Regions (11+12). [Superregión AmazónicaGuayanense].
28
Argentinian Montean: Biogeographic province of the
Middle Chilean-Patagonian Region (18.4). Non-tropical, desertic and xeric Mediterranean mostly subtropical South America. [Provincia Monte Argentino].
Austroamerican: Non-tropical South-American subkingdom (Bb), covering a wide territory usually
south of parallel 30ºS, both Mediterranean (Middle
Chilean-Patagonian Region) and temperate or boreal
macrobioclimates (Pampean and Valdivean-Magellanian Region). To the north it borders on the tropical
macrobioclimate with Brazilian-Paranense, Chacoan,
Tropical South Andean and Hyperdesertic Tropical
Pacific regions. It has gondwanic paleorelationships
and later ones with the Neozelandic-Australian Kingdom and the Circumantarctic Subkingdom. Three regions are recognized: Pampean (17), Middle ChileanPatagonian (18) and Valdivean-Magellanian (19)
[Subreino Austroamericano].
Austrocircummediterranean: Meridional pluricontinental geographic area with a gondwanic origin, and
Mediterranean macrobioclimate, potentially made up
of the Austro-American, South-African and Australian evergreen climatophilous forests, semideserts
and deserts. Territories with Mediterranean macrobioclimate of the southern hemisphere [Área Austrocircunmediterránea]
Austrocircumtemperate: Meridional pluricontinental
geographic area with a gondwanic origin, with temperate bioclimate, potentially formed by the AustroAmerican, Australian-New-Zealander and South-African warm mostly evergreen and gymnospermic
climatophilous forests. Territories with temperate
macrobioclimate of the southern hemisphere [Área
Austrocircuntemplada].
Austrocoldtemperate: Meridional pluricontinental
wide geographic area with a gondwanic origin, mostly located south of parallel 35ºS, formed by cold and
temperate latitudinal and altitudinal zones, to the
south of the subtropical boundary. Territories with
temperate macrobioclimate of the southern hemisphere, often with deciduous forest, with a thermicity
index lower than It<200. [Área Austrocriotemplada].
Benian: Biogeographic province of the Brazilian-Paranense Region (13.8). Tropical South America; low
and middle basin of Mamore and Beni rivers, mostly
infra-thermotropical humid. [Provincia Beniana].
Bolivian-Tucumanan: Biogeographic province of the
Tropical South Andean Region (15.4). Tropical South America; eastern Andes of Bolivia and Argentina
as far as Tucuman above the Chacoan Region. [Provincia Boliviana-Tucumana].
Boreal Austromagellanian: Austro-American biogeographic province (19.3), with boreal hyperoceanic
bioclimate, in the south of the Valdivean-Magellanian Region [Provincia Austromagallánica Boreal].
Brazilian Atlantic: Biogeographic province of the
Brazilian-Paranense Region (13.1). Infra-thermotro-
pical coastal Braziliantropical. South America south
of San Francisco river. [Provincia Atlántica Brasileña].
Brazilian-Paranense: Atlantic South-American biogeographic region (13), located in amazonic and rioplatensean basin. Meridional equatorial (Ceará, Maranhao), meridional eutropical (Bahía, Planalto, Mato
Grosso, Rondonia, Pantanal and Beni), and meridional subtropical to the east of the Paraguay-Paraná river (parallel 30ºS), whose bioclimate is mostly tropical pluviseasonal submesophytic, as well as tropical
pluvial, at the south of the tropic of Capricorn, and
tropical xeric in the Catinga-Sertao (basin of the San
Francisco river). The Brazilian-Paranense region includes eight biogeographic provinces: 13.1. Brazilian
Atlantic, 13.2. Paranense, 13.3. Catingan, 13.4. Tocantins, 13.5. East Cerrado, 13.6. West Cerrado,
13.7. Pantanalian and 13.8 Benian [Región Brasileña-Paranaense].
Caribbean-Mesoamerican:Mesoamerican biogeographic region (9), with tropical bioclimate, chiefly pluviseasonal, to a lesser extent pluvial or xeric and rarely desertic in some islands or Antilleancoasts. It includes all the Caribbean islands, the Florida and Yucatan peninsulas, the rainy territories of Veracruz and
all over Central America to the north of Darien. The
Caribbean-Mesoamerican Region comprises six biogeographic provinces: 9.1. Floridian, 9.2. Cuban, 9.3.
Lesser Antillean, 9.4. Veracruzenian-Yucatecan, 9.5.
Chiapan-Honduran and 9.6. Panamanian-Costa Rican
[Región Caribeña-Mesoamericana]
Caribbean-Neogranadian: Caribbean and North South-American biogeographic superegion (Bab), with
tropical macrobioclimate formed by the CaribbeanMesoamerican and Neogranadian Regions (9+10).
Caribbean and New Granada. [Superregión Caribeña-Neogranadina].
Catingan: Biogeographic province of the BrazilianParanense Region (13.3) mostly tropical xeric. Tropical South America. Catinga. [Provincia Catinguense].
Central Amazonian: Biogeographic province of the
Amazonian Region (12.5); it has been also called
Madeira-Tapajoz. Tropical South America. Central
Amazon. [Provincia Amazónica Central].
Central Chilean: Biogeographic province of the
Middle Chilean-Patagonian Region (18.2). Mediterranean non-tropical South America. [Provincia Chilena Central].
Chacoan: Atlantic tropical South-American biogeographic region (14), located to the northwest of Paraguay river (Paranense biogeographic province) with
dry and semiarid tropical xeric bioclimate. Its central
axis is the Pilcomayo river when it reaches the plain
after flowing across the tropical Andes in the Xerophytic Punenian and Bolivian-Tucumanan Provinces.
Towards the north it borders on Santa Cruz de la Sierra with Chiquitanía, West Cerrado Province, and
towards the northeast with the Pantanal. Towards the
south, when the tropical macrobioclimate disappears
and evolves to temperate, it come into contact with
the Pampean Region (Xerophytic Pampean Province)
and when it becomes Mediterranean xeric or desertic
it is replaced by the subdesertic vegetation of the Argentinean Montean, the start of the North Patagonian
Province of the Middle Chilean-Patagonian Region.
The Chacoan Region comprises two biogeographic
provinces: 14.1. North Chacoan and 14.2 South Chacoan. [Region Chaqueña].
Chacoan-Brazilian: South-American biogeographic
superegion (Bad) with tropical macrobioclimate made up of Brazilian-Paranense and Chacoan Regions
(13 + 14). [Superregión Chaqueña-Brasileña].
Chiapan-Honduran : Biogeographic province of the
Caribbean-Mesoamerican Region (9.5). Central America. [Provincia Chiapaneca-Hondureña].
Circumantarctic: Biogeographic subkingdom (Bc),
with polar macrobioclimate, exceptionally boreal hyperoceanic in some subtemperate isles, hypergelid to
a large extent made up of the Antarctic Continent and
the islands and archipelagos peripheral to the Antarctic Peninsula and the Antarctic Continent (Continental Antarctic Region), as well as the islands far
from the continent, immersed in the westerly winds
and the icy waters of the Antarctic Convergence (Insular Antarctic Region: South Orkney, South Georgia, South Sandwich, Bouvet, Prince Edward, Crozet,
Kerguélen, McDonald, Macquarie, Balleny, Scott,
etc.). The Circumantarctic Subkingdom comprises
two biogeographic regions: 20. Insular Antarctic and
21. Continental Antarctic. [Subreino Circunantártico].
Colombian Andean: Biogeographic province of the
Neogranadian Region (10.3). Tropical South America of the Andes from Cotopaxi volcano in Ecuador
next to Quito to the Andes of Mérida, Eastern and
Central Range and Cauca Valley, extending from infratropical xeric in the inner Magdalena deep valleys
to cryorotropical pluvial in the snowy high mountains. It has been also named Andina Paramuna due
to its cold and humid high plains enriched with espeletineans (frailejones). [Provincia Andina Colombiana].
Colombian Pacific: Biogeographic province of the
Neogranadian Region (10.5). Tropical South America. [Provincia Pacífico Colombiana].
Continental Antarctic: Biogeographic region belonging to the Circumantarctic Subkingdom (21) with
pergelid polar bioclimate and scarce suprapolar bioclimate Tp<20. Because of its orography, bioclimate,
geography and biota, two great territories can be
identified on the Antarctic Continent, to the west and
east of the Transantarctic Mountains, to which we
confer the provincial biogeographic level. 21.1. West
Antarctic Province: this extends along the coast toward the west and south from the Filchner and Ronne
ice shelves to follow the coasts close to the Ross Ice
29
Shelf (20ºW-160º E: West Antarctic or Lesser Antarctic), which includes the Ellsworth Mountains with
the Vinson Peak (5140 m) culminating at the top of
Antarctica (West Antarctic Province). 21.2 East Antarctic Province: this extends along the coast towards
the east from the Usarp mountains on Oates Coast
(160º E) to the Ekström ice field on Princess Martha
Coast (20º W: East Antarctic or Greater Antarctic),
which includes as its culminating part the Dome Argus (4030 m); to the west of the French station of
Charcot (2435 m) an ice glacier thickness of 4776
has been measured; the record for the coldest temperature on the surface of the Earth (-89.4º C) is held
by the Russian station of Vostok (3488 m). The Antarctic Continent, with more than 13.5 million square
kilometres, was the center of the Gondwana supercontinent 180 million years ago, after which a set of
subcontinents: South America, Africa, India, Australia and New Zealand separat and slid toward the
north, leaving the Antarctica more or less fixed
around the South Pole, isolated and perhaps partly or
wholly under ice cover for 40 million years. Today,
98 percent of the surface of Antarctica is covered by
a thick ice cover of about 2.5 km of thickness on average. There are also wide flat glacial floating planes
(ice shelf), some of 1000 km in length in protected
coastal areas (Ross & Ronne Ice Shelves). The annual precipitation exceeds 600 mm in some hyperoceanic windward coastal or insular adjacent localities, while many extreme continental areas inside the
center-east have less than 30 mm. The dominant bioclimate is polar pergelid and only upper suprapolar
(Tp<20) in the marked hyperoceanic coasts and attached islands (Graham Land at the end of the Antarctic Peninsula and surrounding islands, especially
in sunny places with environmental humidity). Only
two vascular plants are known, with distribution
Austro-American linked to coastal habitats with
freshwater or very little brackish and seasonal humidity available, nevertheless there are several hundred lichen species, bryophytes, fungi and protophytes, especially on well-exposed rock habitats
where melt water or cryptoprecipitations are available at least some time during the summer. Among
birds and mammals that breed on the continent or
adjacent islands, always in low continental sea-land
stations, it is worth mentioning five species of penguins, more than a dozen bird and six seal species.
Despite having been able to detect warmer and
colder periods in Antarctica over the past 150,000
years and correlate them with the concentrations of
carbon dioxide –an increased greenhouse-gas effect
in the Earth– in the last 50 years the increase of CO2
from 280 ppm to 370 ppm (global warming) does not
seem to have had a significant impact on the current
increase in temperature in East Antarctica. [Región
Antártica Continental].
Cordobesa-Lower Magdalena: Biogeographic province of the Neogranadian Region (10.4). Xeric to
pluvial infra-thermotropical Caribbean tropical South
America of the plains, hills and dams of the Sinu,
Magdalena and Cauca basins from Barrancabermeja
and Cáceres to the swamps close to Plato. [Provincia
Cordobesa-Bajomagdalena].
30
Deltaic Amazonian: Biogeographic province of the
Amazonian Region (12.3), infratropical pluvial and
pluviseasonal mesophytic. Tropical South America.
Amazonian Delta. [Provincia Amazónica Deltaica].
Deltaic Orinoquian: Biogeographic province of the
Guyanan-Orinoquian Region (11.2), infratropical
pluvial. Tropical South America. Orinoquian Delta.
[Provincia Orinoquense Deltaica].
Desertic Mediterranean Chilean: Biogeographic
province of the Middle Chilean-Patagonian Region
(18.1). Nontropical South America. Toward the south
of Antofagasta (24º C) the coastal and interior hyperdeserts, deserts and semideserts are already Mediterranean (winter rainfall) until Region V (32ºS), and
belong to the Desertic Mediterranean Chilean Province which, from the extreme hyperdeserts (Io 0.00.1) lacking in vascular climatophilous vegetal cover
to the III Region (26ºS), continues northwards with
the acute and moderate deserts with arid cacti such as
Eulychnia tenuis and Copiapoa marginata. North of
Vallenar in the IV Region (29ºS) the arid deserts (Io
0.4-0.9) with large cacti of Eulychnia breviflora appear; and reaching up until north of Coquimbo (30º
S), the semideserts (Io 1.0-1.9) with large Cactaceae,
rosulate puya and shrubs of Eulychnia acida, Echinopsis skottsbergii, Echinopsis litoralis, Puya chilensis and Lithraea caustica, preamble to the Mediterranean pluviseasonal sclerophylous forests and chaparrals of the class Lithraeo-Cryptocaryetea, typical of
the Central Chilean Province (18.2). [Provincia Chilena Mediterránea Desértica].
Desertic Peruvian-Ecuadorean. Biogeographic province of the Tropical South Andean Region (15.1).
Deserts and semideserts (Io 0.4-2.0) of the tropical
mountains and coasts of the Andean western slopes,
from the coast of Manta Bay in Ecuador (1º S), that
together with the Tumbes Peruvian coast up to Machala, with the short tree Loxopterigium huasango,
constitute two biogeographic districts: Desertic Coastal Ecuadorean and Tumbesian, in Peru from El Alto
in Piura (4º 10’ S) towards the south leaves the littoral to cover the interior deserts and semideserts and
the western Andean foothills up until Tarata in Tacna
and the borders of Chile (18º S). In contrast, the ultrahyperarid and hyperarid hyperdeserts (Io 0.0-0.4)
belong to the Hyperdesertic Tropical Pacific Region
(16) extending along the coasts, ridges and mountains ranges from Talara (4º 30’ S) in Peru to Antofagasta (24º S) in Chile. Tropical South America. [Provincia Peruana-Ecuatoriana Desértica].
East Antarctic: Biogeographic province of the Continental Antarctic Region (21.2), polar pergelid and
scarcely upper suprapolar (Tp<10) in sun exposedcoasts. East Antarctic. This could be also named Greater Antarctic. [Provincia Antártica Oriental].
East Cerrado: Biogeographic province of the Brazilian-Paranense Region (13.5). Tropical South America. Eastern Cerrado. [Provincia Cerradense Oriental].
Guajiran-Caribbean: Biogeographic province of the
Neogranadian Region (10.1). Tropical South America. Caribbean from Cartagena to the Paria Peninsula,
with the Guajira, Maracaibo, Falcón, Barquisimeto,
Valencia, Caracas and coastal Venezuelan mountains. To the south it borders with the Llaneran
(10.2), Colombian Andean (10.3) and Cordobesa-Lower Magdalena (10.4) biogeographical provinces.
[Provincia Guajireña-Caribeña].
Guaviarean-Orinoquian : Biogeographic province of
the Guyanan-Orinoquian Region (11.3). Tropical
South America. [Provincia Guaviareña-Orinoquense].
Guayaquilian-Ecuadorean: Biogeographic province
belonging to the Neogranadian Region (10.6). Pacific
equatorial tropical South America. It extends along
the coast from the Guayaquil Gulf (3ºS) up the Ancon Bay on the Colombian border (1º 30’N), excluding a desertic narrow coastal territory with mangroves going from the Puná Island to the Manta Bay.
In contrast, the coast is always hyperarid and ultrahyperarid with cold sea waters and without mangroves extending from latitude 4º 10’ S up to 24ºS to
the Hyperdesertic Tropical Pacific Region (16)]. The
Andean ranges of the Colombian Andean Province
(10.3), with the typical frailejones, extend as far as
Ecuador up to the snow covered volcanoes of Cotopaxi (5897 m) and Iliniza (5383 m) (1ºS). South of
this latitude the mountain chain continues the
Guayaquilian-Ecuadorean Province, whose highest
peak is the Chimborazo (6310 m). In the high humid
or even hyperhumid supra-cryorotropical high
mountains there are short and tall pajonales and
scrubland climatophilous communities, as well as the
perennifolius meso-micro-cloud-forests which become serial shrubby pajonales when damaged. In the
upper semiarid-subhumid infra-mesotropical belts,
mostly on the pacific slope, the natural matured
vegetation, seriously damaged by agriculture, corresponds to deciduous and semideciduous micro-mesoforests that in the hottest areas can be thorny or doliform. The xeric mesotropical deciduous forests with
small doliform trees and cacti are also visible at the
south of the province (6ºS) next to the Abra Porculla
(2145 m) and Bagua low hot valley, in the Marañón
basin. Eastwards from the Napo to the Pastaza rivers
it borders the humid lower thermotropical pluvial
belt with the West Amazonian Province (12.1) and at
the Condor Mountain Range and the Chinchipe basin, the humid-hyperhumid meso-lower supratropical
pluvial belt with the lauroid and Podocarpus humid
cloud forests of the Yungenian Province (15.5)
coming from the Eastern Peruvian Andes. [Provincia
Guayaquileña-Ecuatoriana].
Guyanan: Biogeographic province of the GuyananOrinoquian Region (11.1). Tropical South America.
[Provincia Guayanense].
Guyanan-Orinoquian: Equatorial and Atlantic northern eutropical south American biogeographic region
(11), with pluvial and pluviseasonal bioclimate, ex-
tending over the whole of the hard Guyanese Shield
and the surrounding sandy fluvial and windy sand
deposits which give rise to soils and waters which are
extraordinarily poor in nutrients and bases such as
Old Guyana, Vichada, Guaviare, Vaupés and Roraima territories. To the north and south it borders on
the Neogranadian and Amazonian Regions respectively, both of which have rich soils and waters. The
Guyanan-Orinoquian Region comprises five biogeographic provinces: 11.1. Guyanan, 11.2. Deltaic Orinoquian, 11.3. Guaviarean-Orinoquian , 11.4. Tepuyan and 11.5. Guyanese Brazilian . [Región Guayana-Orinoquense].
Guyanese Brazilian : Biogeographic province of the
Guyanan-Orinoquian Region (11.5). Tropical South
America. [Provincia Brasileña Guayanense].
Hyperdesertic North Peruvian : Biogeographic province of the Hyperdesertic Tropical Pacific Region
(16.1). Tropical South America. Hyperdesertic northern Peru. [Provincia Norperuana Hiperdesértica].
Hyperdesertic Tropical Chilean-Arequipan: Biogeographic province of the Hyperdesertic Tropical Pacific Region (16.2). Tropical South America. Hyperdesertic Tropical Chile and Arequipa, ultrahyperarid
and hyperarid coasts, ridges and mountain ranges of
Northern Arequipa, Moquegua and Tacna Peruvian
departments. [Provincia Chilena-Arequipeña Tropical Hiperdesértica].
Hyperdesertic Tropical Pacific: South American tropical biogeographic region (16), extending from 4º
10’ S in the hyperarid coastal of Talara (Peru) to the
extreme ultrahyperarid tropical deserts of Antofagasta (24ºS). To this region belongs the extreme
marked and moderate hyperdeserts, thermo-mesotropical hyperarid and ultrahyperarid (It > 320; Io <
0.4), which in the Hyperdesertic Tropical ChileanArequipan Province can reach almost 3000 meters,
with the moderate hyperdeserts of columnar cactus
of Browningia candelaris. The rocky or clayey hyperdeserts, without occasional hydric contribution or
fog, lack vascular vegetation cover, but the misty deserts with unmeasurable drizzled moisture during
some weeks in year, have different types of vegetation depending on the topography and substrate
(coastal ridges), such as the aerohygrophile communities of Tillandsia sp. pl. which grow on several
substrates and dune communities, or those than grow
on deep sandy soils formed by numerous radicant,
geophytic or terophytic plants. In the hyperdeserts
the natural matured vegetation corresponds to very
open formations of different cactus, some of them
crassiarborescents columnar microphanerophytics
with very low growth such as Neoraimondia arequipensis. The Hyperdesertic Tropical Pacific Region
comprises two biogeographic provinces: 16.1. Hyperdesertic North Peruvian and 16.2. Hyperdesertic
Tropical Chilean-Arequipan. However it has few floristic and vegetational relationships with the deserts
and semideserts of the Desertic Peruvian-Ecuadorean
biogeographic province (15.1). [Región Pacífica Tropical Hiperdesértica].
31
Insular Antarctic: Biogeographic region belonging to
the Circumantarctic Subkingdom (20) with polar hyperoceanic, oceanic and pergelid bioclimate, and exceptionally boreal hyperoceanic only formed by isles
located in the austral glaciated sea around the Antarctic Continental (Continental Antarctic Region),
linked by the permanent or temporary iced sea all the
year (banquisa) and the meeting between the ice waters (-2ºC) and temperate ones (0º to 3ºC) of the Antarctic Convergence (about 1600 km of the coast),
adjacent to the circumpolar current, driven by the
west wind drift. Inside the region we accept three
oceanic groups of islands or archipelagos at the biogeographic province level: 20.1 Insular Atlantic Antarctic Province (20ºE-80ºW): Bouvet Island, South
Georgia Islands, Diego Ramírez Islands, South
Sandwich Islands and South Shetland Islands; 20.2
Insular Indian Antarctic Province (20ºE-140ºE): Crozet Island, Prince Edward Island, Heard Island, Kerguélen Islands and McDonald Islands and 20.3. Insular Pacific Antarctic Province (140ºE-80ºW):
Balleny Islands, Macquarie Islands, Peter Island and
Scott Islands. The terrestrial and oceanic-terrestrial
biota is quite poor and with gondwanic lineage, but
with influences of the nearby large continents and
islands: South America, Africa, Australia, New Zealand and Tasmania. [Región Antártica Insular].
Insular Atlantical Antarctic: Biogeographic insular
province of the Insular Antarctic Region (20.1). Atlantic Antarctic archipelagos and islands: South
Georgia, Diego Ramirez, Bouvet, South Sandwich
and South Shetland (20º E-80º W) with thermo-suprapolar and pergelid bioclimates. [Provincia Islas
Antárticas Atlánticas].
Insular Falkland: Biogeographic insular province of
the Valdivean-Magellanian Region (19.4). Atlantic
boreal (antiboreal) South America. [Provincia Islas
Malvinas].
Insular Galapagos: Biogeographic insular province
of the Neogranadian Region (10.7). Xeric-desertic
pacific equatorial tropical South America. [Provincia
Islas Galápagos].
Insular Juan Fernandez: Biogeographic insular province of the Valdivean-Magellanian Region (19.5).
Mediterranean and temperate subtropical pacific non
tropical South America. [Provincia Islas Juan
Fernández].
Lesser Antillean: Biogeographic province of the Caribbean-Mesoamerican Region (9.3). Antilles Islands. [Provincia Antillana Menor].
Llaneran: Biogeographic province of the Neogranadian Region (10.2). Xeric and pluviseasonal infratropical Atlantic tropical South America of the plains,
hills, rivers and dams of water in rich soils from the
river Meta to Guajira river, with hydrophytic herbaceous and humid wooded natural permanent savanna
to pluviseasonal evergreen or deciduous rich soil forest vegetation climax. [Provincia Llanera].
32
Mediterranean Andean: Biogeographic province of
the Middle Chilean-Patagonian Region (18.3). Nontropical South America. Mediterranean Andes. [Provincia Andina Mediterránea].
Mesophytic Pampean: Biogeographic province of the
Pampean Region (17.1). Subhumid and humid temperate Atlantic (30º-40º S) South America. [Provincia Pampeana Mesofítica].
Mesophytic Punenian: Biogeographic province of the
Tropical South Andean Region (15.2). Tropical
mesophytic and hygrophytic subhumid to hyperhumid puna of South America. [Provincia Puneña Mesofítica].
Middle Chilean-Patagonian: Pacific and Atlantic
non-tropical South American biogeographic region
(18), all with Mediterranean macrobioclimate. The
region obliquely crosses the subcontinent from one
ocean to another (Pacific coast: 24º-38º S, Atlantic
coast 40º-52º S); across a broad Mediterranean
bridge in the high Andes (approx. 30º-35ºS). Its
northern Andean mountain zone corresponds to the
puna floristic district Cuyano (MARTÍNEZ CARRETERO 1995). From the extreme ultrahyperarid thermo-mesomediterranean hyperdeserts of Atacama
(24ºS) and the central Chilean deciduous and sclerophyllous dry to subhumid thermo-mesomediterranean micro-mesoforests (38ºS), it reaches the oromediterranean belt of the cordillera with the climatophilous scrubland and “pajonales” of the Mediterranean Andean Province and, in the high Andes, the
climatophilous cryoromediterranean dry-humid pulvinate vegetation. Beyond the cordillera appears the
meso-megascrubland, more or less thermic semiaridarid of the Argentine Monte biogeographic Province,
and continues towards the south, on the eastern side
of the cordillera, with northern or southern Patagonian supra-oromediterranean semiarid-dry dwarf
shrubby vegetation. As far as the coast in Rio Gallegos (51º 50’ S), appears the temperate xeric macrobioclimate with the productive grasslands in summer
and, more toward the southwest the remains of the
old micro-mesoforests of Nothofagus, particularly
when the ombrotype is subhumid or humid with the
Valdivean-Magellanian mesoforest of Wintero-Nothofagetea class. The Middle Chilean-Patagonian Region comprises six biogeographic provinces: 18.1.
Desertic Mediterranean Chilean, 18.2. Central Chilean, 18.3. Mediterranean Andean, 18.4. Argentine
Monte, 18.5. North Patagonian and 18.6. South Patagonian. [Región Mesochilena-Patagónica].
Neotropical: Biogeographic subkingdom of America
(Ba), with tropical macrobioclimate; that is to say, all
territories of the equatorial and eutropical latitudinal
belts (0º-23º N & S), as well as the subtropical belts
(23º a 35º N & S) having tropical macrobioclimate.
Five biogeographic superegions are recognized in
this subkingdom: Baa. Mexican (Mexican Xerophytic and Madrean (regions: 7+8), Bab. CaribbeanNeogranadian (regions: 9+10), Bac. Amazonian-Guyanan: (regions: 11+12), Bad. Chacoan-Brazilian (regions: 13+14) and Bae. Tropical South Andean (regions: 15+16). [Subreino Neotropical].
Neotropical-Austroamerican: American and Antarctic Biogeographic Kingdom (B) established by three
broad subkingdoms having distinct bioclimates: Ba.
Neotropical (tropical bioclimates: North, Central and
South America); Bb. Austro-American (temperate,
mediterranean and boreal bioclimates): Bc. Circumantarctic (polar bioclimates: Antarctic and related archipelagos). [Reino Neotropical-Austroamericano]
North Amazonian: Biogeographic province of the
Amazonian Region (12.2); it has been also named
Roraima. Tropical South America. Western Amazon.
[Provincia Amazónica Septentrional].
North Chacoan: Biogeographic province of the Chacoan Region (14.1). Tropical South America. Northern Chaco. [Provincia Chaqueña Septentrional].
North Patagonian: Biogeographic province of the
Middle Chilean-Patagonian Region (18.5). Non-tropical Mediterranean South America. [Provincia Patagónica Septentrional].
Neogranadian: Neotropical biogeographic region
(10) cover a wide northern territory of South America from the inner Guayaquil Bay and the Andean Peruvian-Ecuadorean mountains south of Loja (6ºS) to
the Gulf of Panama in the Pacific and the Gulf of Darien, the Guajira (12º 30’ N) and Paria (11ºN) peninsulas in Caribbean Sea (10ºN). To the east and south
it borders on the Guyanan-Orinoquian (11), Amazonian (14) and Tropical South Andean (15) regions.
The Neogranadian Region (see provinces text) comprises seven biogeographic provinces: 10.1. Guaji
ran-Caribbean, 10.2. Llaneran, 10.3. Colombian Andean, 10.4. Cordobesa-Lower Magdalena, 10.5.
Colombian Pacific, 10.6. Guayaquilian-Ecuadorean
and 10.7. Insular Galapagos. [Región Neogranadina]
Pampean: Non-tropical Atlantic Temperate SouthAmerican biogeographic region (17) belonging to the
Austro-American Subkingdom. All the region has a
temperate macrobioclimate (temperate oceanic and
xeric), located between 30ºS and 40ºS parallels and
chiefly in the low basins of the Paraná and Uruguay
rivers which come from the tropical Brazilian-Paranense Region (approx. 30ºS). A considerable part of
the Pampean Region territory, except for its internal
and surrounding mountains ranges: Ventana, Carape,
San Luis and Córdoba (2884 m), has a very recent
origin (Holocene). For this reason the native flora
and particularly the arborescent one is very poor.
Towards the south and west of the rioplatensean territorial region the bioclimate changes to Mediterranean xeric or desertic and consequently to the wide
Middle Chilean-Patagonian Region, always across
the Argentine Monte Province. The Pampean Region
comprises two biogeographic provinces: 17.1. Mesophytic Pampean and 17.2. Xerophytic Pampean. [Región Pampeana]
Panamanian-Costa Rican: Biogeographic province
of the Caribbean-Mesoamerican Region (9.6). Central America. [Provincia Panameña-Costarricense].
Pantanalian: Biogeographic province of the Brazilian-Paranense Region (13.7). Tropical South America. The Pantanal of South America, mostly covered
with hydrophytic herbaceous and permanent or temporary flooded wooded natural permanent savanna.
[Provincia Pantanalense].
Paranense: Biogeographic province of the BrazilianParanense Region (13.2). Tropical South America.
[Provincia Paranaense].
South Chacoan: Biogeographic province of the Chacoan Region (14.2). Tropical South America. Southern Chaco. [Provincia Chaqueña Meridional].
South Patagonian: Biogeographic province of the
Middle Chilean-Patagonian Region (18.6). Non.tropical Mediterranean South America. [Provincia Patagónica Meridional].
Southwest Amazonian: Biogeographic province of
the Amazonian Region (12.4); it has been also named
Acre and Madre de Dios. Tropical South America.
South-western Amazon. [Provincia Amazónica Suroccidental].
Temperate Magellanian: Biogeographic province of
the Valdivean-Magellanian Region (19.2). Humid-ultrahyperhumid temperate non-tropical Pacific South
America.[Provincia Magallánica Templada].
Tepuyan: Biogeographic province of the GuyananOrinoquian Region (11.4), mostly meso-supratropical pluvial. Tropical South America. Tepuys. [Provincia Tepuyana].
Tocantins: Biogeographic province of the BrazilianParanense Region (13.4), mostly infratropical pluviseasonal. Tropical South America. Tocantins River.
[Provincia Tocantinense].
Tropical South Andean: South American biogeographical superegion (Bae), with tropical macrobioclimate formed by the Tropical South Andean and Hyperdesertic Tropical Pacific Regions (15+16). Tropical western coast and Andes. [Superregión Surandina Tropical]
Tropical South Andean: South American biogeographic region (15) belonging to the tropical Andes of
the southern hemisphere, extending from the yungas
and humid punas of the Amazon and Cajamarca departments and South Piura semideserts, as well as the
arid Ecuadorean coasts at the south of parallel (1ºS),
to the cardonales and Chilean-Argentinian tropical
desertic and semidesertic punas of the Cerro las
Tórtolas (6120 m) near the Agua Negra Pass (4735
m) (30ºS). To the north and west it borders on the
Neogranadian Region, inside Guayaquil Bay. To the
east, between the 4ºS and 17ºS parallels, below the
Andean yungas (600-900 m) the pluvial and mesophytic pluviseasonal humid-hyperhumid warm infrathermotropical (It>640) rainforests belong to the
Amazonian Region. In Santa Cruz de la Sierra (Bolivia) the subhumid pluviseasonal forests correspond
to the West Cerrado (Brazilian-Paranense Region)
33
and connect with the Tropical South Andean along a
narrow band coming from the Chiquitanía. The contact with the tropical xeric Chacoan Region is established slightly towards the south, to finish at the
south of the 30ºS parallel in the Mediterranean
mountains and plains of the Middle Chilean-Patagonian Region of the provinces: Mediterranean Andean, Argentine Monte and Central Chilean. The
Tropical South Andean Region comprises five biogeographic provinces: 15.1. Desertic Peruvian-Ecuadorean, 15.5. Yungenian, 15.2. Mesophytic Punenian, 15.3. Xerophytic Punenian and 15.4. BolivianTucumanan. [Región Surandina Tropical].
Valdivean: Biogeographic province of the ValdiveanMagellanian Region (19.1), from mesotemperate to
cryorotemperate subhumid to ultrahyperhumid. Nontropical temperate and boreal South America. [Provincia Valdiviana].
Valdivean-Magellanian: Pacific and Austro-Atlantic
South-American biogeographic region (19), extending southwards from the Araucanía Region IX in
Chile (38ºS) and Rio Gallegos (51º50’S) in Argentina, with woodland and forest of Wintero-Nothofagetea vegetation class. Its macrobioclimate is temperate
and boreal. The boundary between the temperate and
boreal macrobioclimates is located in the middle of
Isla Grande in Tierra de Fuego and the adjacent continental territory of the Strait of Magellan at parallel
53º30’, as well as in Queen Adelaide Archipelago in
the Pacific (52ºS). The Falkland (Malvinas) islands
with boreal hyperoceanic bioclimate constitute a biogeographic province of this region. The extreme hyperoceanic, temperate and Mediterranean subhumidhumid Juan Fernandez Islands with a high number of
endemisms with gondwanic and magellanic origin
are also included in this region. The Valdivean-Magellanian Region comprises five biogeographic provinces: 19.1. Valdivean, 19.2. Temperate Magellanian, 19.3. Antiboreal Magellanian, 19.4. Insular Falkland and 19.5. Insular Juan Fernandez. [Región Valdiviana-Magallánica].
West Amazonian: Biogeographic province of the
Amazonian Region (12.1); also named Loreto Province. Tropical South America. Western Amazon.
[Provincia Amazónica Occidental].
West Antarctic: Biogeographic province of the Continental Antarctic Region (21.1), polar pergelid and
scarcely upper suprapolar (Tp <20) in the coast and
adjacent islands to Palmer Land in the Antarctic
Peninsula. West Antarctic. It is also known as Lesser
Antarctic. [Provincia Antártica Occidental].
West Cerrado: Biogeographic province of the Brazi
lian-Paranense Region (13.6). Tropical South America. Western Cerrado. [Provincia Cerradense Occidental].
Xerophytic Pampean: Biogeographic province of the
Pampean Region (17.2). Non-tropical temperate Atlantic South America. Xerophytic Pampa [Provincia
Pampeana Xerofítica].
34
Xerophytic Punenian: Biogeographic province of the
Tropical South Andean Region (15.3). Tropical
South America. Xerophytic Puna. Semiarid, dry and
subhumid ombrotypes, supra-, oro-, cryorotropical
thermotypes. [Provincia Puneña Xerofítica].
Yungenian: Biogeographic province of the Tropical
South Andean Region (15.5). Tropical South America. Eastern tropical Andean territories, where it is
frequent to find humid, cloudy and evergreen shiny
leaved meso-macroforest as matured vegetation,
mainly in the humid-hyperhumid thermo-supratropical belt. Yunga. [Provincia Yungueña].
References
The most significant and understandable vegetational publications that we know and used for Biogeographic of South America initial advance are disted in the
references.
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Acosta-Solis, M. 1984. Los páramos andinos del Ecuador.
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Aguirre, Z., Madsen, JE., Cotton, E. & Balslev, H. (eds.)
2002. Botánica austroecuatoriana. Estudios sobre los recursos vegetales en las provincias de Oro, Loja y ZamoraChinchipe. Abya-Yala (edit.), Quito.484 pp.
Alcaraz, F. 1999. Manual de teoría y práctica de Geobotánica.Universidad de Murcia. 401 pp.
Alvarez, M., Ramírez, C. & Deil U. 2008. Ecología y distribución de Hydrocotyle cryptocarpa Speg. en Sudamérica. Gayana Bot. 65: 138-143
Amigo, J. 2009. La flora y la vegetación de Chile: un territorio
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Amigo, J., Izco, J. &.Rodríguez-Guitián, MA. 2007. Rasgos
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Amigo J. & Ramírez, C. 1998. A bioclimatic classification of
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Amigo, J., Ramírez, C. & García Quintanilla, L. 2004. The
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Amigo, J., Ramírez, C. & García Quintanilla, L. 2007. Mantle
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G. San Matías
iq
nt
Abrolh os Ba nk
u
-20°
C. Frío
e la P
l ata
S O U T H
ca
-50°
-40°
-30°
-20°
-10°
-110°
-100°
-90°
-40°
A T L A N T
I C
Gulf of San Jorge
West Falkland
A rg e nt in e
Basin
Falkland Is.
O C E A
N
East Falkland
´s St r.
Santa Inés I.
rn
kb u
le
l Coc
ag
Ca na
Be
al
C an
-80°
-50°
Tierra del Staten
Fuego
I.
Sou th Geo
C. Horn
Sou th Orkn
Shetland Is.
-60°
-30°
L. de los Patos
Valdés Peninsula
21.1
-10°
10°
A T L A N T I C
Guyana
G. of Guayaquil
ANTARCTICA
-20°
N O R T H
Tobago
Trinidad
s
Bo
liv
ia
Staten I.
om
-30°
ch
S e a
C. de la Aguja
-70°
-60°
ANTARCTICA
-50°
rgia
w i c h I s.
n
bur
ock
C
l
a
gle
Can
ea
B
al
Can
19.3
Nicaragua
-40°
ey Is.
-40°
Sa
nd
Str.
19.4
Tr
en
-50°
Guadeloupe
Dominica
Martinique
St. Lucía
Barbados
s
19.2
C a r i b b e a n
a la
Panama
-40°
Falkland Is.
-60°
au
CIRCUMAN
10°
S O U T H
lan
ca
Argentine
Basin
West Falkland
-50°
-50°
Gu
at em
A T L A N T
I C
Gulf of San Jorge
-70°
C. Gracias a Dios
Mar del Plata
Comodoro
Rivadavia
-80°
Honduras
n
(
!
du ras
de
ro
Ba
hía
B
18.6
Santa Inés I.
-40°
-40°
55. Hyperoceanic [Hiperoceánico]
ate
Pl
EAN
ND
ellan´s
Mag
-50°
AUSTROAM ERICAN Subking
dom
Polar [Polar]
n
HA
UT n
S O gi o
A L er e
IC up
S
San Valentín
4058
#
os I.
Madre de Di
-30°
-10°
Boreal [Boreal]
aco
d e la
Valdés Peninsula
I.
Wellington
-30°
-20°
31. Xeric [Xérico]
33. Oceanic [Oceánico]
A
OP
-20°
Chonos
insula
Taitao Pen
-20°
(
!
(
!
G. San Matías
18.5
as
Gulf of Pen
CHACOAN-BRAZILIAN Sup
eregion
Colora
do
Ch
ub
u
Chiloé I.
Arch ip ié la go
-10°
-30°
P
!
w i c h I s.
TR
-10°
NEOTROPICAL Subkingdom
19.1
(
!
Bahía Blanca
Neg
(
!
Puerto Montt
R i s e
-40°
Montevideo
R ío
(
!
La Plata
nd
-40°
-50°
Temperate [Templado]
Ch
P
!
t
C h
i l
e
-60°
22. Hyperdesertic-Oceanic [Hiperdesértico-Oceánico]
24. Desertic-Oceanic [Desértico-Oceánico]
26. Xeric-Oceanic [Xérico-Oceánico]
28. Pluviseasonal-Oceanic [Pluviestacional-Oceánico]
o
18.4
0°
-70°
ío s
Buenos Aires
17.2
18.2
10°
Valdivia
0°
-80°
Mediterranean [Mediterráneo]
(
!
AMAZONIAN-GUYANAN Superegion
-90°
11. Hyperdesertic [Hiperdesértico]
12. Desertic [Desértico]
13. Xeric [Xérico]
14. Pluviseasonal [Pluviestacional]
15. Pluvial [Pluvial]
L. de los Patos
Pelotas
(
!
-100°
Tropical [Tropical]
Pôrto Alegre
L. Mar
Chiquita Santa Fé!( Paraná
Santiago
-40°
CARIBBEAN-NOVOGRANATENSEAN Superegion
55
-20°
Curitiba!(
ua
y
-50°
P
!
Concepción
10°
46
-50°
(
!
18.3
Mt. Aconcagua
# 6960
-40°
33
Campinas
13.2
Asunción
(
!
26
(
!
(
!
-30°
31
Juiz de Fora Campos
(
!
ra
ná
Pa
(
!
33
34
33
31
28
Abrolhos Bank
Pa rag uay
14.1
18.1
15
13.1
Belo Horizonte
-20°
13
24
-40°
13.5
13.7
Cerro Ojos San Miguel
del Salado de Tucumán
(
!
# 6863
an
Arch. de Ju
-10°
22
Salvador
(
!
15.3
13
11
Aracaju
Goiânia
Sucre
16.2
N
O C E A
(
!
(
!
15.4
Antofagasta
13.3
12
34
Iquique!(
I C
P A C I F
Recife
P
!
Santa Cruz
(
!
-20°
(
!
Brasìlia
(
!
(
!
-20°
o
(
!
(
!
L. Poopó
13
14
-10°
-30°
13.6
#
(
!
o
Sã
isc
anc
Fr
or é
Nevado de Ancohuma
6550
P
!
La Paz
Cochabamba
Arequipa
13.4
(
!
L. Titicaca
15.1
s
Ara
M
a
Lima
(
!
ir e
os
Gu
ap
(!
!
P
Cusco
Teresina!(
in
Ar
os
Di
de
e
dr
o
lad
Sa
-50°
Ta
pa
jós
12.4
15.2
C. São Roque
15
or é
M am
-60°
Pôrto Velho
P
Callao
12
Fortaleza
(
!
Natal
s
Tel e
(
!
15.5
-10°
province (provincia)
-70°
á
uan
ip
Huascarán
#6768
velt
Roose
Chimbote
(
!
0°
15
13
(
!
li
Trujillo
0°
Ar
19.5
-80°
12.5
10°
14
(
!
12.3
-40°
13
0°
São Luis
Ma
de
s
ru
Pu
a
Uc ay
(
!
Belém
(
!
Santarém
(
!
a z on
ira
Ju r u á
(
!
-30°
-90°
Br a
n co
Am
n
(
!
Manaus
r añón
Pta. Negra 16.1
Chiclayo
A
zo
ma
-50°
14
Equator
Xi n g u
Ma
Pta. Pariñas
12.2
12.1
-60°
13
10°
Toca ntins
G. of Guayaquil
Putumayo
Na
p
-70°
15
Marajó I.
Japurá
-80°
eR
Cotopaxi
5897
15.1
10.6 # Chimborazo
Guayaquil
6267
(
!
#
10.7
-90°
Gr
Quito!P
o
10.7
Ga lápagos Is.
11.5
Ne g ro
Ca
q uet
á
Authors: S. Riva s Sáenz; Rivas-Martínez, S.; Pe nas, Á; Navarro , G.; Costa, M.
e
C. de San Francisco
1.500 km
BIOCLIMATES
ty n
an
Guaviare
0°
11.1
1.000
Lambert Equal-Area Azimuthal Projection
O C E A N
Cayenne
C. Orange
P
!
500
E ntr
o
10.3
2810 #
11.4 Mt. Roraima
11.3
P
!
Paramaribo
P
!
250
P a m p
a s
lf
Go
Georgetown
P
!
M e ta
0
p a t a g o
n i
a
de
Pa
(
!
11.3
10°
A T L A N T I C
Ciudad
co
rino !(Guayana
a
10.5
O
Cúcuta
Trinidad
11.2
e
10.4
Panamá
P
!
n am
á
9.6
N O R T H
Tobago
P Port of Spain
!
l
10°
(
!
G. of Darién
Canal
5800 !(Maracaibo 10.1
Caracas
P
!
Sierra NevadaL. Maracaibo
( Valencia
!
(
!
de Sta. Marta
Barquisimeto
10.1
10.1
#
(
!
d
Cartagena
San José Panamá
P
!
P
!
L
Barranquilla
St George's
I. Margarita Grenada
C. de la Aguja
á
L. Nicaragua
P
!
with collaboration of:
Javier Amigo (Chile and Argentina), Alindo Butzke (Brazil), Sara del Río (Argentina,
Brazil and Paraguay), Antonio Galán (Peru), 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 (†) (Peru)
na m
Tre
nc
h
10.1
e q ui b o
la
Managua
a le na
tem
a
a gd
Gua
S e a
P
!
Pa
San Salvador !P
P
!
Tegucigalpa
M
9.5
Basse-Terre
o
Guatemala
SALVADOR RIVAS-MARTÍNEZ, GONZALO NAVARRO, ÁNGEL
PENAS & MANUEL COSTA
Guadeloupe
Dominica
Fort-de-France !P Martinique
Castries !P St. Lucía
Kingstown !P 9.3 Barbados
P
!
Bridgetown
St. Vincent
onduras
Co
c
P
!
-40°
C ord
C . . Oc ci
cen de n t
al
tr
C. al
Or
ien
ta l
G . de H
-50°
ur
Co
B. NEOTROPICAL-AUSTROAMERICAN Kingdom [NEOTROPICAL-AUSTROAMERICANO]
Ba. NEOTROPICAL Subkingdom [NEOTROPICAL]
Bab. CARIBBEAN-NEOGRANADIAN Superegion [CARIBEÑA-NEOGRANADINA]
9. CARIBBEAN-MESOAMERICAN Region [CARIBEÑA-MESOAMERICANA]
9.3. Lesser Antillean Province [Antillana Menor]
9.5. Chiapan-Honduran Province [Chiapaneca-Hondureña]
9.6. Panamanian-Costa Rican Province [Panameña-Costarricense]
10. NEOGRANADIAN Region [NEOGRANADINA]
10.1. Guajiran-Caribbean Province [Guajireña-Caribeña]
10.2. Llaneran Province [Llanera]
10.3. Colombian Andean Province [Andina Colombiana]
10.4. Cordobesa-Lower Magdalena Province [Cordobesa-Bajomagdalena]
10.5. Colombian Pacific Province [Pacífico Colombiana]
10.6. Guayaquilean-Ecuadorean Province [Guayaquileña-Ecuatoriana]
10.7. Insular Galapagos Province [Islas Galápagos]
Bac. AMAZONIAN-GUYANAN Superegion [AMAZÓNICA-GUAYANENSE]
11. GUYANAN-ORINOQUIAN Region [GUAYANA-ORINOQUENSE]
11.1. Guyanan Province [Guayanense]
11.2. Deltaic Orinoquian Province [Orinoquense-Deltaica]
11.3. Guaviarean-Orinoquian Province [Guaviareña-Orinoquense]
11.4. Tepuyan Province [Tepuyana]
11.5. Guyanese Brazilian Province [Brasileña Guayanense]
12. AMAZONIAN Region [AMAZÓNICA]
12.1. West Amazonian Province [Amazónica Occidental]
12.2. North Amazonian Province [Amazónica Septentrional]
12.3. Deltaic Amazonian Province [Amazónica Deltaica]
12.4. Southwest Amazonian Province [Amazónica Suroccidental]
12.5. Central Amazonian Province [Amazónica Central]
Bad. CHACOAN-BRAZILIAN Superegion [CHAQUEÑA-BRASILEÑA]
13. BRAZILIAN-PARANENSE Region [BRASILEÑA-PARANAENSE]
13.1. Brazilian Atlantic Province [Atlántica Brasileña]
13.2. Paranense Province [Paranaense]
13.3. Catingan Province [Catinguense]
13.4. Tocantins Province [Tocantinense]
13.5. East Cerrado Province [Cerradense Oriental]
13.6. West Cerrado Province [Cerradense Occidental]
13.7. Pantanalian Province [Pantanalense]
13.8. Benian Province [Beniana]
14. CHACOAN Region [CHAQUEÑA]
14.1. North Chacoan Province [Chaqueña Septentrional]
14.2. South Chacoan Province [Chaqueña Meridional]
Bae. TROPICAL SOUTH ANDEAN Superegion [SURANDINA TROPICAL]
15. TROPICAL SOUTH ANDEAN Region [SURANDINA TROPICAL]
15.1. Desertic Peruvian-Ecuadorean Province [Peruana-Ecuatoriana Desértica]
15.2. Mesophytic Punenian Province [Puneña Mesofítica]
15.3. Xerophytic Punenian Province [Puneña Xerofítica]
15.4. Bolivian-Tucumanan Province [Boliviana-Tucumana]
15.5. Yungenian Province [Yungueña]
16. HYPERDESERTIC TROPICAL PACIFIC Region [PACÍFICA TROPICAL HIPERDESÉRTICA]
16.1. Hyperdesertic North Peruvian Province [Norperuana Hiperdesértica]
16.2. Hyperdesertic Tropical Chilean-Arequipan Province [Chilena-Arequipeña Tropical
Hiperdesértica]
Bb. AUSTROAMERICAN Subkingdom [AUSTROAMERICANO]
17. PAMPEAN Region [PAMPEANA]
17.1. Mesophytic Pampean Province [Pampeana Mesofítica]
17.2. Xerophytic Pampean Province [Pampeana Xerofítica]
18. MIDDLE CHILEAN-PATAGONIAN Region [MESOCHILENA-PATAGÓNICA]
18.1. Desertic Mediterranean Chilean Province [Chilena Mediterránea Desértica]
18.2. Central Chilean Province [Chilena Central]
18.3. Mediterranean Andean Province [Andina Mediterránea]
18.4. Argentine Monte Province [Monte Argentino]
18.5. North Patagonian Province [Patagónica Septentrional]
18.6. South Patagonian Province [Patagónica Meridional]
19. VALDIVEAN-MAGELLANIAN Region [VALDIVIANA-MAGALLÁNICA]
19.1. Valdivean Province [Valdiviana]
19.2. Temperate Magellanian Province [Magallánica Templada]
19.3. Boreal Austromagellanian Province [Austromagallánica Boreal]
19.4. Insular Falkland Province [Islas Malvinas]
19.5. Insular Juan Fernandez Province [Islas Juan Fernández]
Bc. CIRCUMANTARCTIC Subkingdom [CIRCUNANTÁRTICO]
20. INSULAR ANTARCTIC Region [ANTÁRTICA INSULAR]
20.1. Insular Atlantical Antarctic Province [Islas Antárticas Atlánticas]
21. CONTINENTAL ANTARCTIC Region [ANTÁRTICA CONTINENTAL]
21.1. West Antarctic Province [Antártica Occidental]
21.2. East Antarctic Province [Antártica Oriental]
region (region)
-80°
BIOGEOGRAPHIC MAPS OF
THE WORLD: SOUTH AMERICA
Parnaib a
-90°
gua
ia
BIOGEOGRAPHICAL UNITS
-30°
-20°
-10°
Technical cartography: Miguel Álvarez, Ignacio Prieto
© EditAEFA

Biogeographic Map of South America. A preliminary

Transcripción

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