Limited phenotypic plasticity in range-edge populations: a comparison of co-occurring populations of two Agrimonia species with different geographical distributions.

Increased importance of genetic drift and selection for stress resistance have been predicted to lead to a reduction in the degree of phenotypic plasticity in populations at margins of a species' geographical range, relative to those in the centre. We examined the effect of population positioning within the species range on degree of active morphological plasticity to vegetation shade. Importantly, we discriminated between active, size-independent morphological adjustments in response to shade and passive changes in morphology caused by the dependence of morphological traits on plant size, as only the former are considered to be adaptive. Two closely related and ecologically similar Agrimonia species were examined in the same geographical location, where one species reaches the edge of its distribution (Agrimonia pilosa) and the other does not (A. eupatoria). Plasticity to light availability is likely to be advantageous for both species as they occupy habitats with variable light conditions. However, we hypothesised that high levels of environmental stress should lead to reduced active plasticity in marginal compared with more central populations. Agrimonia eupatoria exhibited active adjustments in leaf morphology in response to tree shade, and in elongation of stems and inflorescences in response to herbaceous shade. In contrast, A. pilosa exhibited very limited active plasticity. High active plasticity allowed A. eupatoria to retain constant shoot growth in a wide range of light conditions, while the lack of active plasticity in A. pilosa resulted in a strong dependence of shoot growth on light availability. We propose that high levels of environmental stress in marginal areas of a species' range may lead to a significant reduction in the degree of active plasticity. Our results clearly indicate that discrimination between active and passive plasticity is crucial for reaching valid conclusions about differences in adaptive plasticity between marginal and non-marginal populations.