Ecology's cruel dilemma, phylogenetic trait evolution and the assembly of Serengeti plant communities

P>1. Ecologists debate the importance of neutral versus niche-based explanations for patterns of species coexistence and whether small-scale data can inform ecological understanding of communities, referred to by McNaughton [Ecological Monographs, 1983, 53, 291] as 'ecology's cruel dilemma.' Research on phylogenetic relationships, traits and species co-occurrence has attempted to address this topic, with results considerably mixed.2. We address the hypothesis that plant community assembly is influenced by trait similarity across ecological gradients and this affects mean phylogenetic distance (MPD) of species within sites. We analysed specific leaf area (SLA), maximum plant height and phylogenetic relationships among Serengeti grasses, a system ideally suited to study community assembly because of an ecological gradient in which the dominant plant stress shifts from drought to light competition.3. Phylogenetic community assembly theory predicts that MPD would be lowest (under-dispersed) at dry sites and greatest (over-dispersed) at sites with higher rainfall. Similarly, theory predicts that low soil nutrient concentrations should filter intolerant species, so that MPD is expected to be under-dispersed at infertile, low-elevation sites and over-dispersed at fertile, higher-elevation sites. However, as gradients of rainfall and soil fertility run counter to one another across the Serengeti, it was unclear how this covariation would influence MPD.4. Surprisingly, traits showed different evolutionary patterns: SLA displayed convergent evolution while maximum plant height displayed Brownian evolution across the phylogeny. As predicted, statistically under-dispersed assemblages occurred at lower rainfall, infertile sites while statistically over-dispersed assemblages occurred at higher rainfall, fertile sites. However, the pattern across all plots was weak, with most plots showing no statistical pattern of MPD.5. Multivariate analyses using structural equation modelling, which statistically controlled for covariation among environmental effects, revealed complex direct and indirect effects of environmental variation on MPD, including offsetting direct effects of SLA and maximum plant height due to their different patterns of trait evolution.6. Synthesis. Spatially counteracting gradients of moisture and soil fertility across the Serengeti, combined with contrasting patterns of trait evolution, obscured the relationship between MPD and any single environmental variable. Our study shows that integrating trait and phylogenetic relationships across ecological gradients yields considerable insight into the ecological mechanisms that determine community composition, but that multivariate techniques may be required to appropriately reveal such patterns.