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Such clustering is likely to be a consequence of pollen diagrams containing relatively high percentages of Podocarpus and Hedyosmum in association with Arecaceae, Ericaceae, Ilex, Myrsine, Myrtaceae and Weinmannia Figs.

In Lake Pata, at ca. A similar mechanism was also proposed much earlier as an explanation for the occurrence of Andean forest elements, such as Alnus, Podocarpus, making up new floristic associations with lowland taxa in Ecuadorian Amazonia 31 , 32 involving Alnus, Hedyosmum and Podocarpus site 49, San Juan Bosco and 50, Mera, ca 300 m elev. Higher percentages of Alnus during the LGM in marine sediments of site 53 Amazon River Delta are likely to represent long-distance pollen signatures exported by the Amazon Basin during high precipitation events prevailing during the LGM 9 , 23 , 31 , 32. This threshold, although considered a background effect in Colombian Andean lakes 79 , can hypothetically be interpreted as active migration of this arboreal taxon into lowlands of the Brazilian Amazon. Outlier cluster points such as sites 27 and 31 in southeastern coastal Brazil and sites 32 and 39, in the Cerrado region of central Brazil, are again signaling floristic similarities brought about by cooling and higher humidity levels, which characterize most of the South American lowlands during the LGM. On the other hand, sites 49 and 50 San Juan Bosco and Mera, in Ecuador are outside this cluster by an artifact caused only by the presence of Ericaceae pollen, indicative of cool and moist environments. Such connectivity between the Andes and the Amazon could have only been possible if one assumes a large LGM rainforest cover within most of the Amazon Basin, dominated by cold- and humid-adapted taxa, a scenario that has recently been proposed based on a biome modelling analysis for paleovegetation cover 107.

All pollen spectra related to this connectivity are interpreted as a consequence of downward migration of high-elevation taxa into the Amazonian lowlands during the LGM, forming non-analog plant assemblages whose key taxa are now found with disjunct distributions in the Neblina region of the Upper Rio Negro, in Brazil 17. We hypothesize that this large exposed area was vegetated by cold and humid successional forests as a consequence of the downward migration of montane taxa into the coastal lowlands, a scenario supported by pollen records sites 17 and 30 from modern mangrove areas in southeastern Brazil 52 , 60. A similar mechanism of rainforest and mangrove vegetation belts expanding and contracting in synchrony with sea level oscillations has been observed for the mid Holocene of coastal northeastern Brazil 108. Despite the apparent connection in terms of habitat suitability, the absence of Araucaria in the LGM pollen records of Chile might represent a significant retraction of Araucaria araucana Mol. Koch, the endemic Chilean species, in synchrony with a hypothetical expansion of Araucaria angustifolia into Uruguay and northern Argentina, especially onto the exposed continental shelf. Drimys winteri JR Forst. Another possible branch of this connection is suggested by higher habitat suitability during the LGM displayed by Araucaria, Ericaceae, Symplocos, Weinmannia and Arecaceae in northern Argentina, Paraguay and Uruguay, south of the Chaco region.

Noteworthy is the presence, within this cluster, of the Brazilian site of Pato Branco 15 , relatively close to the Andes and the borderline site Sao Francisco de Assis 12 in coastal Brazil, at 91 m elevation which can be explained by forest spectra dominated by Podocarpus. The absence of Ericaceae and Symplocos in most of these Andean sites can be hypothesized to be a consequence of paramo vegetation expansion downslope during the LGM, generating pollen spectra dominated by grasses and few arboreal elements 68 , 69. This LGM scenario closely resembles the cold migration corridors for multiple montane forest refugia proposed for the Heinrich Stage 1 climatic phase, ca. The disjunct pattern is likely to represent a relatively isolated forest ecosystem created by the Andean geographical barrier containing Drimys, Ericaceae, Myrtaceae, Podocarpus and Weinmannia. The modern landscape of this area, between sites 9 and 10, is characterized by the southern Chile endemic Araucaria araucana as the dominant taxa in the emergent forest canopy, a species which surprisingly is not present in the LGM pollen records. A tentative explanation relies on the absence of palynological studies covering this glacial phase in these higher southern latitudes. It may also be that this taxon migrated to ice-free areas without lake basins or peatbogs to capture a local pollen history.

Analysis of modern distributions as influenced by former climates: Are relicts still there? We approached the question of whether relict populations can still be found in present-day landscapes by investigating the modern distributions of 137 species belonging to Andean genera occurring in the Atlantic domain and evaluated their ecological niches by means of SDM MaxEnt. Each map Figs. The same applies to Clethra brasiliensis, C. There are two possible hypothesess that could explain the absence of these species in database. The first could represented biased caused by field sampling and taxonomical identification methodologies. The second, and very plausible alternative explanation invokes habitat loss caused by unsuitable climatic change after the LGM.

Table 1 depicts the results of data integration validating LGM connections across South America with the presence of modern relicts and strengthened by a composite map of their geographical distributions Fig. Podocarpus and Weinmmania, and to a lesser degree Escallonia and Berberis, appear to be the main drivers in delimiting relicts which can be explained by their long-distance dispersal of pollen and seeds. Table 1 Taxa occurrences in modern relicts and LGM fossil pollen records in relation to proposed connectivity patterns between the Andean and Atlantic domains. This same pattern of forest formation can be applied to the exposed Amazonian portion of the South American continental shelf based on LGM pollen spectra present in offshore marine sediments site 53 indicating the presence of a cool and moist tropical forest. Its floristic composition was dominated by the montane taxa Podocarpus, Weinmannia, Myrica, Symplocos, Hedyosmum and Alnus, also suggestive of a downslope migration into the lowlands. A recent analysis of pollen signatures transported by Amazonian rivers to its delta 109 suggests nearby terrigenous sources as the main component in marine sediments, which implies that cold and humid forests were thriving in fluvial valleys of eastern Amazonia and by consequence on the exposed coastal shelf during the LGM. A similar scenario of large scale regional vegetation coverage was reported for the LGM of the exposed Southeast Asia-Australasia continental shelf with the predominance of rainforest in wetter areas, resulting in a greater area of rainforest than nowadays 110.

During the LGM these taxa, currently restricted to cool and humid montane ecosystems, thrived sympatrically along with Cerrado elements such as Antonia ovata Pohl, Byrsonima L. Another significant example is given by Araucaria and Podocarpus thriving sympatrically during the LGM with Cerrado elements at Lagoa Santa in south Central Brazil 18 and during most of the last glacial cycle ca. It is worth mentioning that SDM and modern occurrences especially of Podocarpus and Hedyosmum in the tepui region of southern Venezuela and northern Brazilian Amazonia sites 51—52, Lagoa da Pata depict possible former connections between these areas under a scenario of 1100 m downslope migration 23 , 116 of cold-adapted taxa, forming plant assemblages of warm- and cold-adapted taxa living sympatrically 23 , 106 , 117 and without modern counterparts. Former climatic mechanisms and northward displacement of cold air masses Our overall data confirm that during the Last Glacial Maximum, cold fronts were intensified and could have reached farther north than at present, thus affecting northeastern Brazil. Because the resulting equator-pole temperature gradient was therefore larger, transient systems had probably higher intensity thus causing convergence of humidity precisely where the mean humidity zonal stream flow was high 13 , 15 , 120 , 121. In this period, in the regions near the equator, the changes in humidity and temperature were smaller and, at high latitudes, larger, with lower temperatures and less humidity. Circulations such as Rossby waves were likely to be intensified and amplified, which resulted in the propagation of cold fronts and associated cold air masses to the equatorial region 13 , 97 , 98 , 122.

This scenario is corroborated by recent cases of the incursion of cold waves from the high latitudes of the southern hemisphere that even cross the equator because of the break in the geostrophic balance caused by the Andes 123 , 124 , 125 , 126 , 127 , 128. The pressure gradient is maintained high temperature contrast with decreasing Coriolis force decreasing wind 13 , thus enhancing the cold air mass acceleration northwards. Genetic and additional palynological evidence for a late Quaternary Andean-Atlantic connection A wide array of evidence for historical connections between Andean and Atlantic ecosystems has been provided by genetic analyses in certain groups of animals, such as birds 101 , 129 , 130 , 131 , 132 , 133 , frogs 134 and rodents 135. Among these studies, there is significant support for a major disjunction between the Andes and the Atlantic Forest between 1 Ma and 0. Other analyses also based on genetic flow between closely related bird species with Andean-Atlantic disjunct distributions focused on the LGM as a more recent decisive moment of connectivity, under humid phases and expansion of forests in both the Cerrado and Chaco routes 129.

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Окончил Белорусский государственный университет, факультет международных отношений г. Минск, Республика Беларусь , и аспирантуру Института мировой экономики и международных отношений Российской академии наук г. Москва, Российская Федерация.

Кандидат экономических наук. Является автором нескольких научных работ в области инвестиций и инновационного развития, а также одним из авторов национального доклада «Инновационное развитие — основа модернизации экономики России» [1].

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