Functional trait space and the latitudinal diversity gradient

Abstract

The processes causing the latitudinal gradient in species richness remain elusive. Ecological theories for the origin of biodiversity gradients, such as competitive exclusion, neutral dynamics, and environmental filtering, make predictions for how functional diversity should vary at the alpha (within local assemblages), beta (among assemblages), and gamma (regional pool) scales. We test these predictions by quantifying hypervolumes constructed from functional traits representing major axes of plant strategy variation (specific leaf area, plant height, and seed mass) in tree assemblages spanning the temperate and tropical New World. Alpha-scale trait volume decreases with absolute latitude and is often lower than sampling expectation, consistent with environmental filtering theory. Beta-scale overlap decays with geographic distance fastest in the temperate zone, again consistent with environmental filtering theory. In contrast, gamma-scale trait space shows a hump-shaped relationship with absolute latitude, consistent with no theory. Furthermore, the overall temperate trait hypervolume was larger than the overall tropical hypervolume, indicating that the temperate zone permits a wider range of trait combinations or that niche packing is stronger in the tropical zone. Although there are limitations in the data, our analyses suggest that multiple processes have shaped trait diversity in trees, reflecting no consistent support for any one theory. S pecies richness increases toward the equator (1, 2) in major clades of both extant and extinct species of plants and animals (3, 4). The generality of the pattern hints at a correspondingly general explanation, yet the latitudinal gradient in species richness remains one of ecology's greatest unsolved puzzles. Long-running debates over the causes of the latitudinal gradient of species richness have focused on ecological, evolutionary, and geographic explanations (5-10). Although there has been some progress (11), it is also increasingly clear that there are numerous obstacles to understanding the primary drivers of the latitudinal gradient, including an ever-increasing number of hypotheses (12, 13), challenges in clearly separating their interdependencies (14, 15), and difficulties in rigorously falsifying their assumptions and predictions (16).More powerful tests of biodiversity theories need to move beyond species richness and instead explicitly focus on the mechanisms generating the gradient, by recasting the theories in terms of other measures of diversity, such as functional diversity (17-19). For example, explanations that assume species richness is limited by resource availability have often focused on the strength of species interactions, life history differences, and environmental constraints on how species pack into niche space (20). Evolutionary hypotheses have focused on differences in diversification rates, as well as the influence of species interactions on diversification rates (9). These interaction-based explanations implicitly refer to the degree of ecol...

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