Inconsistent relationships between area, heterogeneity and plant species richness in temperate farmed landscapes

Relationships between area, heterogeneity and species richness are fundamental concepts in ecology yet questions remain about how area and heterogeneity tradeoff (AHTO) to constrain biodiversity. Although there is growing evidence for unimodal heterogeneity diversity relationships (HDR's) and an AHTO, tests of the concept and consequences for species richness across a landscape-scale gradient of human-modified ecosystems are rare. Using data from a national (Wales) field survey we analysed relationships between environmental heterogeneity and plant species richness (α and γ). We used ordination to produce a composite metric of heterogeneity and compared this to commonly used metrics. We used niche hypervolumes to categorise the breadth of plant species' ecological preferences and analysed relationships between species richness, niche width and heterogeneity. The HDR was unimodal with α diversity at the smallest scale and positive with α and γ diversity (non-linear) at the 1 km scale although in low intensity landscapes the HDR with γ diversity was unimodal. There was a unimodal relationship between habitat diversity and γ diversity. Land use intensity was unimodally related to diversity. There were significant interactions between niche width and heterogeneity. Richness of broad niche species increased with heterogeneity with flattening of the curve at higher levels. Narrow niche species were rare and mostly unresponsive. The expected decline in narrow niche species with increasing heterogeneity was not found although they did decline with land-use intensity. Using a unique dataset, an analysis of a large-scale mosaic of ecosystems found that the shape of the HDR varies with land use intensity, the heterogeneity metric, spatial scale, diversity type and niche width. Although heterogeneity can increase species richness, there may be tradeoffs at higher heterogeneity. A fundamental constraint on realising the benefit of heterogeneity is the low availability of narrower niche species in local species pools in modified landscapes.