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72(1), 102-112,
2008 2008 Review ISSN 0717-652X
Julin Monge-Njera
Biologa Tropical, Universidad de Costa Rica, 2060 San Jos, Costa Rica; biologia.tropical@ucr.ac.cr,
julianmonge@gmail.com
ABSTRACT
Ecological biogeography studies the factors that define the spatial distribution of species in the present time. This review
summarises recent contributions on ecological biogeography. Most recent articles report environmental factors such as
temperature, humidity and salinity as key elements in the ecological biogeography of many species (followed by other
organisms and genetic characteristics). Molecular data indicate that some unexplainable ranges are artifacts caused by
taxonomic misidentification (several species erroneously classified as a single species). Island biogeography theory is
often adequate for conservation management, and the new neutral model of ecological biogeography does not fit all
species on which it has been tested. Global warming leads to range expansions, dispersal events, and new invasions. Until
now, most experimental work has been done on temperate ecosystems. In the 21st century, tropical biogeographers
should do landmark contributions by doing field, laboratory and simulation experiments about species ranges and
community biogeography.
RESUMEN
La biogeografa ecolgica estudia los factores que definen la actual distribucin espacial de los organismos. Esta revisin
resume los aportes ms recientes en ese campo. La mayora de los artculos recientes informan que factores ambientales
como temperatura, humedad y salinidad son elementos clave en la biogeografa ecolgica de muchas especies (seguidos de
interacciones con otros organismos, y caractersticas genticas). La biologa molecular est develando que algunas
distribuciones incomprensibles son en realidad el resultado de identificaciones taxonmicas incorrectas en que varias
especies eran consideradas una sola especie. La teora de biogeografa de islas a menudo es aplicable a la conservacin,
pero el nuevo modelo neutralista de biogeografa ecolgica no calza con algunos de los organismos a los cuales se ha
aplicado. El calentamiento planetario probablemente producir ampliaciones de mbito, dispersiones y nuevas invasiones.
Hasta ahora, la mayora del trabajo experimental en este campo se ha hecho en ecosistemas templados, por lo que en el
siglo XXI, la biogeografa tropical debe hacer contribuciones significativas mediante simulaciones y experimentos de
campo y de laboratorio, sobre los mbitos de distribucin geogrfica y la biogeografa de comunidades.
PALABRAS CLAVES: Revisin, biogeografa ecolgica, templada versus tropical, calentamiento global.
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for Madagascan lemurs (Lehman et al. 2006). Like in of California (Cintra-Buenrostro et al. 2005) and
other groups, overriding biotic factors are less tintinnids in the Mediterranean (Krsinic & Grbec
frequently cited, but examples include birds (wood 2006). In the case of Indian Ocean Holothuroids,
warblers in North America) and dwarf chameleons in ranges correlate with dispersal ability (Samyn &
South Africa, whose spatial distribution is clearly Tallon 2005), and in South African ascidians, the
defined by vegetation (Kelly & Hutto 2005, Tolley et correlation is with sociality (Primo & Vzquez 2004).
al. 2006). In contrast with the above examples, no abiotic or
biotic factors have been found to explain the
EC O L O G I C A L FA C TO R S T H AT E X P L A I N ORGANIC ecological biogeography of hydrothermal vent
DISTRIBUTION IN AQUATIC HABITATS communities in the Pacific (Govenar et al. 2005).
In freshwater ecosystems, water level and tempe-
rature often affect organismic distributions. The EXAMPLES OF GEOGRAPHIC RANGES RECONSIDERED AFTER
recent increase in temperature has allowed MOLECULAR ANALYSIS
nonindigenous and invasive water plants to expand Molecular techniques are now frequently used to
their ranges in Sweden (Larson 2006). In Mediterranean understand the ecological biogeography of all groups
rivers, 21% of the distribution of caddisflies is of organisms, and have shown that taxonomic
explained by a combination of ecological factors misidentification originated some cases of
(Bonada et al. 2005). Wetland bird distribution in unexplainable distributional ranges, for example in
Canad is strongly defined by the water level Alopiinae snails, whose distribution fits genetics, not
(Desgranges et al. 2006), and temperature draws the morphology (De Weerd et al. 2004). Unexplainable
limits for the range of Trachemys scripta turtles bird ranges also proved to result from misclassification
(Willette et al. 2005). of convergent taxa in woodpeckers (Moore et al. 2006)
The sea is probably the biome where the ecological and Accipitridae hawks (Do Amaral et al. 2006).
biogeography of organisms has been more intensively Examples of novel genetic results include the finding
studied. Maybe the reason is that, in comparison with that mycorrhizal fungi are not the host generalist,
forests and other terrestrial ecosystems, the ocean is species-poor group that was previously believed
a physically simpler medium. Abiotic factors known (Fitter 2005) and the discovery of the strong host
to define organismic regions in the ocean are specialization of plant pathogenic fungi in North
temperature, substrate, currents, light and nutrients, America (Johnson et al. 2005).
among others. For example, temperature explains the Genetics also show that the range recoveries of some
distribution of fish larvae in Jalisco, Mexico (Navarro- groups, such as the red oaks of the USA, are endangered
Rodrguez et al. 2004); Archaeobacteria in North because recent populations growing on old pasture
America and Crimea (Knittel et al. 2005); seabirds in lands have low genetic diversity (Gerwein & Kesseli
the Pacific (Smith & Hyrenbach 2003); sea stars in the 2006). A similar problem affects damselflies in England
Gulf of California (Cintra-Buenrostro et al. 2005); (Watts et al. 2005), shrews in Tanzania (Stanley &
dinoflagellates cysts in the Artic (Matthiessen et al. Olson 2005) and prong horn antelopes in the USA
2005); coccolithophores worldwide (Baumann et al. (Stephen et al. 2005). In contrast, the ravens of the
2005) and red tide precursor organisms in the Pacific world appear to be one species with a climate-defined
(Sierra-Beltrn et al. 2004). biogeography and large genetic variation (Omland et
Substrate characteristics define the distribution of al. 2006).
gastropods in Britain (Grahame et al. 2006); Barriers are the defining element of ecological
polychaetes in Canada (Quijon & Snelgrove 2005); biogeography, and recent genetic work shows that
echinoderms in Colombia (Neira & Cantera 2005) and barriers can be ethological in the case of the Artic
chondricthyan fish in Britain (Ellis et al. 2005). charr (Adams et al. 2006) and the Mallorcan midwife
Japanese ostracods match ocean currents in their toad (Kraaijeveld-Smit et al. 2006), but also climatic
distribution (Ogoh & Ohmiya 2005), and the same is (for marmots in the USA, Floyd et al. 2005) or
true for sponges worldwide (Woerheide et al. 2005), topographic (for North American salamanders, Liu
and for bacterioplankton and nutrients in the Baltic et al. 2006).
Sea (Sipura et al. 2005). Genetics are also applied to understand the
An important biotic factor is food, for sea-birds in distribution of freshwater organisms. Physical barriers
Beringia (Piatt & Springer 2003), sea-stars in the Gulf affect Thioalkalivibrio bacteria worldwide (Foti et
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al. 2006), and several fish species: the bass in the 2005); forest and sedge land in Australia (Driscoll
USA (Cooke & Philipp 2006), sardines in Brazil (Paiva 2005); Pinus populations in China (Chianget al. 2006);
et al. 2006), the Atlantic Salmon in Russia (Primmer et land snails in Greece and New Zealand (Barker 2005,
al. 2006) and Australian Leiopotherapon fish (Bostock Triantis et al. 2005); dung beetles in Namibia (Sole et
et al. 2006). On the other hand, a gene pool analysis al. 2005); fish in Hungarian streams (Eros & Grossman
of brown trout found that introduced individuals, 2005); seabirds in Australia (Priddel et al. 2006); birds
which had a particular genetic constitution, failed to of prey, in the Mediterranean and Macaronesian
significantly establish in lake Tinnsjo, Norway, even Archipelagos (Donazar et al. 2005); birds in Britain
though the lake is within the range of the species and Ireland (Russell et al. 2006), and Howler Monkeys
(Heggenes et al. 2006). In the Japanese and Korean in Venezuela (Feeley & Terborgh 2006). By contrast,
sections of its range, the bluegill sunfish population the model could not explain the island biogeography
is characterized by a much reduced genetic variability of Australasian land snails, possibly because it
that may represent a bottleneck effect (Kawamura et neglects in situ speciation (Cameron et al. 2005), and
al. 2006). In the case of the Pearly Mussel in the USA, forest-savanna mosaics in Africa (Hovestadt et al.
no isolation by distance has been found (Grobler et 2005).
al. 2006). General recent developments in island biogeography
On marine ecosystems, genetics have shown that refer to the z index, curvilinear pattern, life history
ranges can be defined by both abiotic and biotic and leaf lifespan. Low z values correlate with high
barriers. Abiotic barriers were identified in English colonization ability; a multi-species metapopulation
snails, for which a cliff acts as the barrier (Grahame et model indicated that the relationship of z with
al. 2006), and for Pacific snails (Meyer et al. 2005). important parameters can be predicted only if either
Mussels in the Indian Ridge and the Atlantic Ridge the dispersal ability or the power of establishment is
are limited spatially by endosymbiosis (McKiness & known (Hovestadt & Poethke 2005). He et al. (2005)
Cavanaugh 2005); and again, as in vertebrates, strange presented a new model for local-regional species
ranges may also represent taxonomical richness in ecological islands that does not require
misidentifications, for example in Atlantic amphipods species interactions to produce the curvilinear pattern.
(Kelly et al. 2006). In other cases, traditionally Using island biogeography theory, they found that a
recognized barriers such as the East Pacific Barrier high extinction rate produces a curvilinear pattern of
and the Bahamas barrier do not significantly impair local-region relationships, while a high colonization
gene flow for some fish species (Robertsonet al. 2004, rate produces a linear pattern. Equilibrium models need
Taylor & Hellberg 2006). to consider life history traits, because one island can
be in equilibrium and have barriers for one organism,
ISLAND BIOGEOGRAPHY but not for others (Shepherd & Brantley 2005). Finally,
One aspect of ecological biogeography, island it has been stated that better biogeography models
biogeography, received much attention at the end of for vegetation should include leaf lifespan (Zhang &
the 20th century, first on theoretical grounds, then on Luo 2004).
its potential application to the problem of
conservation of organisms in islands of protected ISLAND BIOGEOGRAPHY AND CONSERVATION
nature in an anthropologically changed world. Island biogeography is often applied in conservation
The MacArthur and Wilson model of island for hypothesis testing, finding ways for wildlife-
biogeography (MacArthur & Wilson 1967) was based human coexistence, and prediction.
on common sense principles: islands that are big and A long term (40 years) study in Japan found that
close to species sources should have more species, biodiversity in islands of habitat in a matrix of non-
and diversity results from the equilibrium of species habitat can be summarized with the phi-coefficient,
addition and extinction. Maybe for that reason, most which has the advantage, over other indices, that it
recent literature reports that such principles hold for can be combined with the Chi-square test to
real islands. Examples include bacteria and shrubs in statistically test hypotheses (Yasuda et al. 2005).
Spain (Maestre & Cortina 2005, Reche et al. 2005); Regarding coexistence, urban water reservoirs can
diatoms in Antarctica (Van de Vijver et al. 2005); conserve the bulk of tropical molluscan biota in
vascular plants in North America (McMaster 2005); Southeast Asia, especially when the pH is near 7.3
calcareous grasslands in Belgium (Bisteau & Mahy and the habitat includes rocks (Clements et al. 2006).
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Recibido: 13.08.07
Aceptado: 15.04.08
112