A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.

Biotic interactions are believed to play a role in the origin and maintenance of species diversity, and multiple hypotheses link the latitudinal diversity gradient to a presumed gradient in the importance of biotic interactions. Here we address whether biotic interactions are more important at low latitudes, finding support for this hypothesis from a wide range of interactions. Some of the best-supported examples are higher herbivory and insect predation in the tropics, and predominantly tropical mutualisms such as cleaning symbioses and ant-plant interactions. For studies that included tropical regions, biotic interactions were never more important at high latitudes. Although our results support the hypothesis that biotic interactions are more important in the tropics, additional research is needed, including latitudinal comparisons of rates of molecular evolution for genes involved in biotic interactions, estimates of gradients in interaction strength, and phylogenetic comparisons of the traits that mediate biotic interactions. 245Annu. Rev. Ecol. Evol. Syst. 2009.40:245-269. Downloaded from www.annualreviews.org by University of Sussex on 10/02/12. For personal use only.

The evolutionary dynamics underlying the latitudinal gradient in biodiversity have been controversial for over a century. Using a spatially explicit approach that incorporates not only origination and extinction but immigration, a global analysis of genera and subgenera of marine bivalves over the past 11 million years supports an "out of the tropics" model, in which taxa preferentially originate in the tropics and expand toward the poles without losing their tropical presence. The tropics are thus both a cradle and a museum of biodiversity, contrary to the conceptual dichotomy dominant since 1974; a tropical diversity crisis would thus have profound evolutionary effects at all latitudes.