You'd better walk alone: Changes in forest composition affect pollination efficiency and pre-dispersal cone damage in Iberian Juniperus thurifera forests.

Changes in land-use patterns are a major driver of global environmental change. Cessation of traditional land-use practices has led to forest expansion and shifts in forest composition. Consequently, former monospecific forests maintained by traditional management are progressing towards mixed forests. However, knowledge is scarce on how the presence of other tree species will affect reproduction of formerly dominant species. We explored this question in the wind-pollinated tree Juniperus thurifera. We hypothesised that the presence of heterospecific trees would have a negative effect on cone production and on the proportion of cones attacked by specialised predators. We assessed the relative importance of forest composition on cone production, seed development and pre-dispersal cone damage on nine paired pure and mixed J. thurifera forests in three regions across the Iberian Peninsula. The effects of forest composition on crop size, cone and seed characteristics, as well as damage by pre-dispersal arthropods were tested using mixed models. Cone production was lower and seed abortion higher in mixed forests, suggesting higher pollination failure. In contrast, cone damage by arthropods was higher in pure forests, supporting the hypothesis that presence of non-host plants reduces damage rates. However, the response of each arthropod to forest composition was species-specific and the relative rates of cone damage varied depending on individual tree crops. Larger crop sizes in pure forests compensated for the higher cone damage rates, leading to a higher net production of sound seeds compared to mixed forests. This study indicates that ongoing changes in forest composition after land abandonment may impact tree reproduction.

Efficiency of pollination and satiation of predators determine reproductive output in Iberian Juniperus thurifera woodlands.

Fruit production in animal-dispersed plants has a strong influence on fitness because large crops increase the number of seeds dispersed by frugivores. Large crops are costly, and environmental control of plant resources is likely play a role in shaping temporal and spatial variations in seed production, particularly in fluctuating environments such as the Mediterranean. The number of fruits that start to develop and the proportion of viable seeds produced are also linked to the number of flowers formed and the efficiency of pollination in wind-pollinated plants. Finally, large fruit displays also attract seed predators, having a negative effect on seed output. We assessed the relative impact of environmental conditions on fruit production, and their combined effect on seed production, abortion and seed loss through three predispersal predators in Juniperus thurifera L., sampling 14 populations across the Iberian Peninsula. Wetter than average conditions during flowering and early fruit development led to larger crop sizes; this effect was amplified at tree level, with the most productive trees during more favourable years yielding fruits with more viable seeds and less empty and aborted seeds. In addition, large crops satiated the less mobile seed predator. The other two predispersal predators responded to plant traits, the presence of other seed predators and environmental conditions, but did not show a satiation response to the current-year crop. Our large-scale study on a dioecious, wind-pollinated Mediterranean juniper indicates that pollination efficiency and satiation of seed predators, mediated by environmental conditions, are important determinants of reproductive output in this juniper species.

The geographic selection mosaic for squirrels, crossbills and Aleppo pine.

The interactions between many species are structured in a geographic mosaic of populations among which selection is divergent. Here we tested the hypothesis that such a geographic selection mosaic arises for common crossbills (Loxia curvirostra) feeding on seeds in the cones of Aleppo pine (Pinus halepensis) because of geographic variation in the occurrence of European red squirrels (Sciurus vulgaris). On the Iberian Peninsula, Sciurus exerted directional selection favouring larger cones with larger scales, which has caused cones there to be larger than in the Balearic Islands where Sciurus are absent. Moreover, cones on the Iberian Peninsula are so large that they are apparently little used by the relatively small-billed crossbills on the Peninsula; selection by Sciurus seems to have made the cones so difficult to feed on that crossbills rely mostly on the seeds of other conifers. Where crossbills are present but Sciurus are absent (Mallorca Island), cones were smaller as a result of relaxation of selection by Sciurus. However, cones on Mallorca had proportionally thicker scales in comparison to where both Sciurus and crossbills are absent (Ibiza Island), presumably as an adaptation against crossbill predation. Here crossbills specialize on Aleppo pine, have relatively large bills and have apparently coevolved in an arms race with Aleppo pine. These results suggest that Sciurus has influenced both the geographic selection mosaics for crossbills and conifers and the adaptive radiation of crossbills in Eurasia much like Tamiasciurus has done in the North America.