Principles of niche expansion.

Niche expansion is attained by adaptations in two generalized phenotypical traits-niche position and niche width. This gives room for a wide range of conceptual ways of niche filling. The niche variation hypothesis reduces the range by predicting that expansion occurs by increasing variation in niche position, which has been debated on empirical and theoretical grounds as also other options seem possible. Here, we propose a general theory of niche expansion. We review empirical data and show with an eco-evolutionary model how resource diversity and a trade-off in resource acquisition steer niche evolution consistent with observations. We show that the range can be reduced to a discrete set of two orthogonal ways of niche filling, through (1) strict phenotypical differentiation in niche position or (2) strict individual generalization. When individual generalization is costly, niche expansion undergoes a shift from (2) to (1) at a point where the resource diversity becomes sufficiently large. Otherwise, niche expansion always follows (2), consistent with earlier results. We show that this either-or response can operate at both evolutionary and short-term time scales. This reduces the principles of niche expansion under environmental change to a notion of orthogonality, dictated by resource diversity and a resource-acquisition trade-off.