Abstract

Three prominent explanations have been proposed to explain the dramatic differences in species richness across regions and elevations, (i) time for speciation, (ii) diversification rates, and (iii) ecological limits. But the relative importance of these explanations and, especially, their interplay and possible synthesis remain largely elusive. Integrating diversification analyses, null models, and geographic information systems, I study avian richness across regions and elevations of the New World. My results reveal that even though the three explanations are differentially important (with ecological limits playing the dominant role), each contributes uniquely to the formation of richness gradients. Further, my results reveal the likely interplay between the explanations. They indicate that ecological limits hinder the diversification process, such that the accumulation of species within a region gradually slows down over time. Yet, it does not seem to converge toward a hard ceiling on regional richness. Instead, species-rich regions show suppressed, but continued, diversification, coupled with signatures of possible competition (esp. Neotropical lowlands). Conversely, species-poor, newly-colonized regions show fast diversification and weak to no signs of competition (esp. Nearctic highlands). These results held across five families of birds, across grid cells, biomes, and elevations. Together, my findings begin to illuminate the rich, yet highly consistent, interplay of the mechanisms that together shape richness gradients in the New World, including the most species-rich biodiversity hotspots on the planet, the Andes and the Amazon. [Biogeography; community; competition; macroevolution; phylogenetics; richness gradient.]

Highlights

  • When Humboldt traveled the New World, he was astonished by the dramatic differences in species richness across regions and elevations, from the lowlands of the tropics to the mountaintops of the temperate (Humboldt and Bonpland 1807)

  • I refrained from analyzing all birds, given that the avian phylogeny has lately been in flux (Jetz et al 2012; Jarvis et al 2014; Hedges et al 2015; Prum et al 2015), and the effects of unresolved phylogenetic relationships would be further exacerbated by the dramatic differences in the quality of the geographic and elevational data available for different parts of the avian phylogeny (IOC World Bird List v8.1, IUCN 2018)

  • Species richness declined from the tropics toward the temperate and from lowlands toward highlands (Fig. 1)

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Summary

Introduction

When Humboldt traveled the New World, he was astonished by the dramatic differences in species richness across regions and elevations, from the lowlands of the tropics to the mountaintops of the temperate (Humboldt and Bonpland 1807). The explanations conflict in multiple respects, such that it is possible to derive their diagnostic predictions (Table 1), and each has been supported and challenged by empirical evidence It is well-documented that regions which have been colonized for a long-time tend to harbor more species, presumably because they have had more time for speciation and the accumulation of richness (Ricklefs 2006; McGuire et al 2014). Interactions between the mechanisms remain largely unresolved It has been theorized (Simpson 1953; Sepkoski 1981; Van Valen 1985; Stroud and Losos 2016) that regional diversification declines over time, as species gradually accumulate within a region. This process can be modulated by historical changes in MACHAC—DIVERSITY DYNAMICS IN BIRDS

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