Formation of the Isthmus of Panama.

The formation of the Isthmus of Panama stands as one of the greatest natural events of the Cenozoic, driving profound biotic transformations on land and in the oceans. Some recent studies suggest that the Isthmus formed many millions of years earlier than the widely recognized age of approximately 3 million years ago (Ma), a result that if true would revolutionize our understanding of environmental, ecological, and evolutionary change across the Americas. To bring clarity to the question of when the Isthmus of Panama formed, we provide an exhaustive review and reanalysis of geological, paleontological, and molecular records. These independent lines of evidence converge upon a cohesive narrative of gradually emerging land and constricting seaways, with formation of the Isthmus of Panama sensu stricto around 2.8 Ma. The evidence used to support an older isthmus is inconclusive, and we caution against the uncritical acceptance of an isthmus before the Pliocene.

Historical biogeography of the Isthmus of Panama.

About 3 million years ago (Ma), the Isthmus of Panama joined the Americas, forming a land bridge over which inhabitants of each America invaded the other-the Great American Biotic Interchange. These invasions transformed land ecosystems in South and Middle America. Humans invading from Asia over 12000 years ago killed most mammals over 44 kg, again transforming tropical American ecosystems. As a sea barrier, the isthmus induced divergent environmental change off its two coasts-creating contrasting ecosystems through differential extinction and diversification. Approximately 65 Ma invading marsupials and ungulates of North American ancestry, and xenarthrans of uncertain provenance replaced nearly all South America's non-volant mammals. There is no geological evidence for a land bridge at that time. Together with rodents and primates crossing from Africa 42 to 30 Ma, South America's mammals evolved in isolation until the interchange's first heralds less than 10 Ma. Its carnivores were ineffective marsupials. Meanwhile, North America was invaded by more competitive Eurasian mammals. The Americas had comparable expanses of tropical forest 55 Ma; later, climate change confined North American tropical forest to a far smaller area. When the isthmus formed, North American carnivores replaced their marsupial counterparts. Although invaders crossed in both directions, North American mammals spread widely, diversified greatly, and steadily replaced South American open-country counterparts, unused to effective predators. Invading South American mammals were less successful. South America's birds, bats, and smaller rainforest mammals, equally isolated, mostly survived invasion. Its vegetation, enriched by many overseas invaders, remained intact. This vegetation resists herbivory effectively. When climate permitted, South America's rainforest, with its bats, birds and mammals, spread to Mexico. Present-day tropical American vegetation is largely zoned by trade-offs between exploiting well-watered settings versus surviving droughts, exploiting fertile versus coping with poor soil, and exploiting lowland warmth versus coping with cooler altitudes. At the start of the Miocene, a common marine biota extended from Trinidad to Ecuador and western Mexico, which evolved in isolation from the Indo-Pacific until the Pleistocene. The seaway between the Americas began shoaling over 12 Ma. About 10 Ma the land bridge was briefly near-complete, allowing some interchange of land mammals between the continents. By 7 Ma, the rising sill had split deeper-water populations. Sea temperature, salinity and sedimentary carbon content had begun to increase in the Southern Caribbean, but not the Pacific. By 4 Ma, the seaway's narrowing began to extinguish Caribbean upwellings. By 2 Ma, upwellings remained only along Venezuela; Caribbean plankton, suspension-feeding molluscs and their predators had declined sharply, largely replaced by bottom-dwelling corals and calcareous algae and magnificent coral reefs. Closing the seaway extinguished the Eastern Pacific's reef corals (successors recolonized from the Indo-Pacific 6000 years ago), whereas many molluscs of productive waters that once thrived in the Caribbean now survive only in the Eastern Pacific. The present-day productive Eastern Pacific, with few, small coral reefs and a plankton-based ecosystem contrasts with the Caribbean, whose clear water favours expansive coral reefs and bottom-dwelling primary producers. These ecosystems reflect the trade-off between fast growth and effective defence with attendant longevity. Overfishing with new technologies during the last few centuries, however, has caused population crashes of ever-smaller marine animals, devastating Caribbean ecosystems.

A spatially explicit neutral model of beta-diversity in tropical forests.

To represent species turnover in tropical rain forest, we use a neutral model where a tree's fate is not affected by what species it belongs to, seeds disperse a limited distance from their parents, and speciation is in equilibrium with random extinction. We calculate the similarity function, the probability F(r) that two trees separated by a distance r belong to the same species, assuming that the dispersal kernel P(r), the distribution of seeds about their parents and the prospects of mortality and reproduction, are the same for all trees regardless of their species. If P(r) is radially symmetric Gaussian with mean-square dispersal distance sigma, F(r) can be expressed in closed form. If P(r) is a radially symmetric Cauchy distribution, then, in two-dimensional space, F(r) is proportional to 1/r for large r. Analytical results are compared with individual-based simulations, and the relevance to field observations is discussed.

Beta-diversity in tropical forest trees.

The high alpha-diversity of tropical forests has been amply documented, but beta-diversity-how species composition changes with distance-has seldom been studied. We present quantitative estimates of beta-diversity for tropical trees by comparing species composition of plots in lowland terra firme forest in Panama, Ecuador, and Peru. We compare observations with predictions derived from a neutral model in which habitat is uniform and only dispersal and speciation influence species turnover. We find that beta-diversity is higher in Panama than in western Amazonia and that patterns in both areas are inconsistent with the neutral model. In Panama, habitat variation appears to increase species turnover relative to Amazonia, where unexpectedly low turnover over great distances suggests that population densities of some species are bounded by as yet unidentified processes. At intermediate scales in both regions, observations can be matched by theory, suggesting that dispersal limitation, with speciation, influences species turnover.