The mid-domain effect: It’s not just about space

Ecologists and biogeographers have long sought to understand how and why diversity varies across space. Up until the late 20th century, the dominant role of environmental gradients and historical processes in driving geographical species richness patterns went largely undisputed. However, almost 20 years ago, Colwell & Hurtt (1994) proposed a radical reappraisal of ecological gradient theory that called into question decades of empirical and theoretical research. That controversial idea was later termed the ‘the mid-domain effect’: the simple proposition that in the absence of environmental gradients, the random placement of species ranges within a bounded domain will give rise to greatest range overlap, and thus richness, at the center of the domain (Colwell & Lees, 2000) (Fig. 1a). The implication of this line of reasoning is that the conventional null model of equal species richness regardless of latitude, elevation or depth should be replaced by one where richness peaks at some midpoint in geographical space. Our intention here is to draw attention to a neglected, yet important manifestation of the mid-domain effect, namely the application of mid-domain models (also referred to as geometric constraint models) to non-spatial domains. If individual species have ranges that exist not just in geographical space but also in environmental factors, such as temperature, rainfall, pH, productivity or disturbance, shouldn’t we also expect mid-domain richness peaks along non-spatial gradients? A mid-domain model applied to non-spatial gradients predicts the maximum potential richness for every value of an environmental factor. As with spatial mid-domain models, realized richness would probably be less, but the limits to richness are still predicted to be hump-shaped. Indeed, hump-shaped relationships emerge with remarkably high frequency across various non-spatial gradients. For instance, two of ecology’s most fundamental, albeit controversial theories – the productivity–diversity relationship and the intermediate disturbance hypothesis – predict mid-domain peaks in species richness. However, the potential of non-spatial mid-domain models has gone largely ignored