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• W1489739311 abstract "The evolutionary origin of biological diversity and the causes of present-day species distribution patterns have long intrigued biogeographers. Most studies that have considered these issues in the Neotropics have tested alternative historical hypotheses (e.g. Hubert & Renno, 2006; Quijada-Mascareñas et al., 2007). These involve changes in dispersal barriers and dispersal routes as a result of changes in climate (as in the Pleistocene refuge hypothesis, which posits that rain forests were fragmented to refugia separated by savanna) or in geography (as in hypotheses proposing that the distributions of land and water have changed as a result of changes in sea level or river systems). Although previously it was common to think that the alternative explanations were mutually exclusive, recent studies recognize that several of them may have played a role in creating patterns of species distribution and diversity. In general, dispersal routes are emphasized in studies aiming to reconstruct how the species of interest have dispersed from their centres of origin to their present ranges. Dispersal barriers are emphasized in vicariance models, in which allopatric genetic differentiation is assumed to have taken place after ancestral species ranges were interrupted by barriers. Species dispersal can be prevented either because the species is physically unable to cross the barrier, or because the species is adapted to a different kind of habitat and does not thrive in the habitat available in the barrier. Hubert & Renno (2006) used species distribution data and a reconstruction of the geological history of South America to draw conclusions about the evolution of South American freshwater fishes. Quijada-Mascareñas et al. (2007) used genetic information on the Neotropical rattlesnake to infer what dispersal routes this savanna species could have used to reach its present distribution, which skirts the Amazonian rain forest area. Both studies made statements about what constitutes a dispersal barrier for the species of interest. For the freshwater fishes, dry land and salt water were considered dispersal barriers (Hubert & Renno, 2006). For the Neotropical rattlesnake, rain forest was considered a dispersal barrier, and savanna (including dry forests and deforested areas) was considered suitable habitat (Quijada-Mascareñas et al., 2007). These are rather coarse habitat classifications, and ecologically it would be surprising if all rivers provided equally suitable habitat for all fish species, or if all savannas were equally suitable and all rain forests equally unsuitable for the rattlesnake. More detailed information on the present-day environmental conditions may be difficult to obtain, and is certainly difficult to reconstruct for the past. However, would the availability of such data make a difference to biogeographical inferences? If the species of interest are specialized to only some kinds of river, forest or savanna, the possible dispersal barriers in the past might have been more extensive, and the dispersal routes more restricted, than is immediately apparent. South America is a large continent with much variation in rainfall, temperature and geology, and some workers have emphasized the biogeographical and evolutionary importance of variation in these environmental factors even within the apparently uniform Amazonian lowlands (Endler, 1982; Tuomisto & Ruokolainen, 1997; Tuomisto, 2006). There are well-known chemical, physical and biological differences between eutrophic white-water rivers and oligotrophic black-water and clear-water rivers. Some terrestrial habitats such as white sand forests, floodplain forests and swamp forests are known to harbour endemic plant and animal species. Even within the ‘typical’, climatically uniform rain forests it has been observed that few, if any, plant species grow on all kinds of soil (Tuomisto, 2006). Although variations in soil and river water are generally considered to be local-scale phenomena, and hence of less importance than climatic variation at broad spatial scales, they present clear regional patterns: western Amazonia is generally more eutrophic than central, southern and northern Amazonia (Sioli, 1975; Clapperton, 1993; Rossetti et al., 2005). This is because northern and southern Amazonia are largely covered by ancient, quartzitic rocks that yield only small amounts of highly leached sediments to the rivers, which typically have black or clear waters. In contrast, the young Andean mountain chain yields large amounts of freshly eroded material to western Amazonian rivers, which typically have white waters. However, none of these major regions is internally homogeneous. Rock type and degree of volcanic activity vary along the Andes, so different rivers carry mineralogically dissimilar materials. In addition, the sediments gradually lose nutrients as they are weathered in situ or are again eroded by the river and redeposited further downstream. Differentiation in sediment properties is reflected in the acidity, nutrient content, potential productivity and other properties of both river waters and soils. These, in turn, are of great importance for plants and aquatic animals, and through their impact on plant productivity may also affect terrestrial animals. In combination with the regionally and temporally variable climate, such differences lead to a large number of different past and present habitats. Western Amazonia has also been affected by marine incursions in the Miocene (Rossetti et al., 2005). A marine incursion is an obvious dispersal barrier for freshwater and terrestrial biota, and may hence have promoted vicariant speciation in Miocene Amazonia (Hubert & Renno, 2006). Marine sediments are also geochemically quite different from sediments of continental origin, and forests growing on the (semi)marine Pebas formation are floristically clearly distinct from forests growing on river terraces in Peruvian Amazonia (e.g. Salovaara et al., 2004). Palaeoarches (subterranean uplifted structures of various kinds) have often been discussed as dispersal barriers, but they may not have emerged high enough to act as such for terrestrial species (Rossetti et al., 2005). If they have formed water divides, they may have prevented the dispersal of aquatic species (Hubert & Renno, 2006), and if they coincide with geochemical boundaries, they may also appear to be of consequence for terrestrial species. However, in the latter case the biogeographical importance of the arch itself might be minimal in comparison to that of the geochemical differences. So, we may ask: to what degree do the present-day species distributions in Amazonia (or South America in general) reflect the ancient presence of dispersal barriers or dispersal routes that have long since disappeared, and to what degree do they conform to the modern distributions of suitable habitat? The marine incursion was an obvious dispersal barrier in the Miocene; however, its ecological impact did not cease when the waters receded, because it permanently altered soil properties and river water characteristics. A similar process can be expected when river-transported sediments of a new kind are introduced to an area, for example when volcanic activity starts in a previously non-volcanic region. Hubert & Renno (2006) interpreted the biogeographical patterns of freshwater fishes in the light of the geological history of South America with an emphasis on dispersal barriers and dispersal routes. However, the placement of the Amazonian rivers in their area cladogram is also consistent with the idea that geochemistry is important. The Amazonian rivers were divided into two clades: the western Amazonian rivers in one, and the central, northern and southern rivers in the other. It would be interesting to conduct supplementary analyses that take into account the geochemical characteristics of each river system to clarify the relative explanatory powers of geochemical similarity and the degree of historical connectivity for faunal similarity among rivers. Quijada-Mascareñas et al. (2007) found their genetic results to support the scenario that the Neotropical rattlesnake dispersed from north to south across the present-day Amazonian rain forest. Because the rattlesnake is a savanna species, they inferred that a savanna corridor must have existed in the Pleistocene. Is this conclusion warranted? The answer depends on how accurate the snake distribution data actually are in terms of geographical location and habitat. Some areas within Amazonia receive relatively little rain even at present, and some rain forest types have a savanna-like structure for edaphic reasons. Arguments based on the absence of a species from a large area are always problematic, especially in poorly studied areas such as Amazonia. A few populations of the Neotropical rattlesnake are known to occur in Amazonia, but Quijada-Mascareñas et al. (2007) did not include these in their genetic analyses, nor did they mention what kind of vegetation these populations inhabit. It is conceivable that the species occurs also in as yet undocumented sites in Amazonia, and that the dispersal barrier formed by the rain forest is not as hard as it seems on the basis of the currently available distribution map. Distinguishing between vicariance and shared environmental patterns is not easy (Endler, 1982; Tuomisto & Ruokolainen, 1997; Ackerly, 2003). Different taxa may show similar area cladograms because speciation has happened through vicariance, and all resident species experienced a breakup of their ancestral ranges in a similar sequence when the various dispersal barriers were formed. However, a similar biogeographical pattern may also arise if different parts of the area of interest have different environmental conditions, which themselves originated in a historical sequence of geological and climatic events. After the formation of each habitat, the plant and animal species that lived in the area (or colonized it from nearby areas) might adapt to it by undergoing genetic change or even speciation (Endler, 1982; Ackerly, 2003). A full understanding of evolution and biogeography seems to require more than studies on when and where geological and climatic processes may have given rise to obvious dispersal barriers (Wiens & Donoghue, 2004). Dispersal barriers may not always be obvious, so the present environmental requirements of the species of interest need to be known in some detail in order to infer where suitable habitats occur at present, where suitable habitats have potentially occurred in the past, and what actually constitutes a dispersal barrier. A discrepancy between present-day environments and species distributions suggests that the impact of past dispersal barriers or other historical factors still lingers. However, without sufficient ecological data, the discrepancy may not be apparent, or it may be interpreted incorrectly. Reconstructions of the past may be very different depending on the degree to which present ecological information is taken into account before the observed patterns are explained with historical events. Editor: Dov Sax" @default.
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• W1489739311 date "2007-04-12" @default.
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• W1489739311 title "Interpreting the biogeography of South America" @default.
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