Vertical stratification patterns of tropical forest vertebrates: a meta-analysis.

Tropical forests harbour the highest levels of terrestrial biodiversity and represent some of the most complex ecosystems on Earth, with a significant portion of this diversity above ground. Although the vertical dimension is a central aspect of the ecology of forest communities, there is little consensus as to prominence, evenness, and consistency of community-level stratification from ground to canopy. Here, we gather the results of 62 studies across the tropics to synthesise and assess broad patterns of vertical stratification of abundance and richness in vertebrates, the best studied taxonomic group for which results have not been collated previously. Our review of the literature yielded sufficient data for bats, small mammals, birds and amphibians. We show that variation in the stratification of abundance and richness exists within and among all taxa considered. Bat richness stratification was variable among studies, although bat abundance was weighted towards the canopy. Both bird richness and abundance stratification were variable, with no overriding pattern. On the contrary, both amphibians and small mammals showed consistent patterns of decline in abundance and richness towards the canopy. We descriptively characterise research trends in drivers of stratification cited or investigated within studies, finding local habitat structure and food distribution/foraging to be the most commonly attributed drivers. Further, we analyse the influence of macroecological variables on stratification patterns, finding latitude and elevation to be key predictors of bird stratification in particular. Prominent differences among taxa are likely due to taxon-specific interactions with local drivers such as vertical habitat structure, food distribution, and vertical climate gradients, which may vary considerably across macroecological gradients such as elevation and biogeographic realm. Our study showcases the complexity with which animal communities organise within tropical forest ecosystems, while demonstrating the canopy as a critical niche space for tropical vertebrates, thereby highlighting the inherent vulnerability of tropical vertebrate communities to forest loss and canopy disturbance. We recognise that analyses were constrained due to variation in study designs and methods which produced a variety of abundance and richness metrics recorded across different arrangements of vertical strata. We therefore suggest the application of best practices for data reporting and highlight the significant effort required to fill research gaps in terms of under-sampled regions, taxa, and environments.