Ecospace: A unified framework for understanding variation in terrestrial biodiversity

Understanding patterns in biodiversity is a core ambition in ecological research. Existing ecological theories focusing on individual species, populations, communities, or niches aid in understanding the determinants of biodiversity patterns, yet very few general models for biodiversity have emerged from simplistic approaches. We propose that a systematic, low-dimensional representation of environmental space with building blocks adopted from gradient, niche, metapopulation and assembly theory may unite old and new aspects of biodiversity theory and improve our understanding of variation in terrestrial biodiversity.We propose the term ecospace to cover the local conditions and resources underlying diversity. Our definition of ecospace encompasses abiotic position, biotic expansion and spatiotemporal continuity, which all affect the biodiversity of a biotope (α-diversity). Position refers to placement along abiotic gradients such as temperature, soil pH and fertility, leading to environmental filtering known from classical community theory. Expansion represents the build-up and diversification of organic matter that are not strictly given by position. Continuity refers to the spatiotemporal extension of position and expansion.Biodiversity is scale dependent. The contribution of one biotope to large scale diversity must be estimated by considering its unique contribution to the species richness of the surrounding landscape or region or to the biodiversity of the entire planet. In addition to the relationship between ecospace and biotope richness (α-diversity), we also propose a relation between the uniqueness of the biotope ecospace and the unique contribution of species to the surrounding larger-scale richness.Whereas the impacts of ecospace position and continuity on biodiversity have been studied in isolation, studies comparing or combining them are rare. Furthermore, biotic expansion has never been fully developed as a determinant of biodiversity, ignoring the megadiverse carbon-depending groups of insects and fungi. Precursors of the ecospace concept have been presented over the last 70 years, but they were never fully developed conceptually for terrestrial biodiversity or applied to prediction of biodiversity.Ecospace unites classical and – at times – contradicting theories such as niche theory, island biogeography theory and a suite of community assembly theories into one framework for further development of a general theory of terrestrial biodiversity.