Universality in biodiversity patterns: variation in species–temperature and species-productivity relationships reveals a prominent role of productivity in diversity gradients

Temperature and productivity appear as universal positive large-scale correlates of species richness. However, the strength and the shape of species–temperature (STR) and species–productivity (SPR) relationships vary widely, and the causes of this variation are poorly known. We analysed 1) published species richness data for multiple taxa sampled in various regions and 2) different clades within vertebrate classes globally, to test for the effects of spatial scale and characteristics of examined taxa and regions on the strength and direction of STRs and SPRs. There are striking differences in the variation of the relationships among types of data, between ectotherms and endotherms and also between STRs and SPRs. Some sources of this variation are of statistical nature (e.g. the relationships are stronger if the range of temperature or productivity variation is wider), but non-statistical sources are more important and illuminate the processes responsible for the origin of biodiversity patterns. The SPRs are generally stronger and less variable than STRs, and SPR variation is weakly related to the explored factors – the SPRs are stronger in warmer regions in ectotherms, while clade size is the only factor consistently affecting the strength of the SPR in endotherms. In contrast, STRs are weaker and more variable, and this variation is linked to region characteristics – most importantly, STRs are stronger in the regions where temperature positively correlates with productivity, indicating that productivity plays a role even in the STRs. The effect of temperature on species richness is thus complex and context-dependent, while productivity is a more universal driver of species richness patterns, largely independent of particular characteristics of given region or taxon. Productivity thus appears as the main proximate driver of species richness patterns, probably due to its effect on the limits of the number of viable populations which can coexist in a given environment.