Limits of pine forest distribution at the treeline: herbivory matters
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Abstract
Herbivores can affect future forest composition by feeding selectivity. At temperature-sensitive treelines, herbivory can exacerbate or constrain climate-driven distributional shifts in tree species. This study analyses the impact of herbivory in a Mediterranean treeline of widespread Pinus sylvestris and P. nigra pinewoods, testing whether herbivory damage reinforces or inhibits the climatic responses of these trees. We used naturally occurring sapling pairs of similar size and age of both species, thereby isolating plant characteristics from environmental effects in herbivore behaviour. Herbivory damage by ungulates proved higher than that caused by insects in saplings of both species. Low plant density and extreme abiotic conditions at the treeline could in part be responsible for the observed low incidence of insect herbivory. Ungulates preferred P. sylvestris over P. nigra, implying heavier browsing damage for a large number of P. sylvestris saplings, suffering reduced internode growth as a consequence. In addition, P. sylvestris could not compensate height-growth reductions due to browsing with higher growth rate than P. nigra. In fact, P. sylvestris showed similar or lower relative height growth with respect to P. nigra. Under a scenario of increasing aridity and maintenance of ungulate populations, the upward migration of P. sylvestris in its southern range could be restricted by higher drought vulnerability than P. nigra, a situation exacerbated by ungulate herbivory. Our results indicate that ungulate herbivory reinforces climatic response of coexisting P. sylvestris and P. nigra at treeline, favouring a potential change in community dominance towards Mediterranean P. nigra.
Keywords
Ungulate browsing Insect herbivory Treeline dynamics Mediterranean mountain Pinus sylvestris Pinus nigraNotes
Acknowledgments
We thank the Consejería de Medioambiente (Andalusian Government) and the direction of Sierra de Baza Natural Park for facilities to carry out the study. We also wish to thank Otilia Romera for field assistance and Luis Cayuela for his help with mixed effects analysis. David Nesbitt checked the English of this article. This study was supported by the coordinated Ministerio de Educación y Ciencia (Spanish Government) Project DINAMED (CGL2005-05830-C03), GESBOME (P06-RNM-1890), and by grant FPU-MEC (AP2005-1561) to A. H.
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