Above- and below-ground competition effects of two heathland species: implications for growth and response to herbivory in birch saplings

We examined experimentally the effect of competition from two common heathland plant species (Calluna vulgaris or Molinia caerulea) on Betula pubescens saplings subjected to simulated mammalian browsing damage. We tested two hypotheses: that B. pubescens saplings alter their growth allocation in response to different patterns of competition from the two species in order to maximise resource acquisition; and that, when only B. pubescens saplings are damaged, herbivory reduces its ability to compete with both species. In an ex-situ experiment we grew B. pubescens saplings in the presence of below- or above- and below-ground interactions from C. vulgaris or M. caerulea. Saplings were also subjected to simulated browsing by clipping (50% of current year's growth), either pre-senescence or at bud-burst. We measured the morphology and dry mass allocation response of the saplings over a period of two years. We found that competition reduced sapling dry mass by approximately 50%, but C. vulgaris reduced dry mass to a greater extent than did M. caerulea. The total competition intensity of C. vulgaris was greater than that of M. caerulea, due to an apparent facilitative effect of M. caerulea shoots on birch growth. Saplings compensated for browsing damage, resulting in no difference in dry mass one year after damage. However, sapling morphological responses to browsing damage were dependent on the competing species. Despite the large competitive effect of below-ground interactions, saplings did not increase allocation to root growth as predicted. Additionally, in response to above-ground interactions from M. caeruela, saplings increased allocation to root growth. This study highlights the importance of patterns, as well as intensity, of competition in determining plant responses to inter-specific interactions. Sapling responses did not follow the pattern predicted by the balanced-growth hypothesis. We suggest that this is due to the multi-functionality of plant component parts and the balance between competitive and facilitative effects of interacting plants.