Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns Ó Springer Science+Business Media B.V. 2006 and, if it existed, how it could condition the fitness consequences of both types of satiation.

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Summary1. The spatial distribution of dispersed seeds results from the combined action of the caching strategies followed by different granivores. Hence, it is essential to study the factors that influence seed predation and caching decisions to achieve a better understanding of the dispersal process. 2. In this study, we document how seed dispersal and the spatial patterns of natural recruitment are linked to the strategies used by granivores to protect their cached seeds from pilferage. We present a theoretical model showing that those strategies may convey benefits for both seed cachers and plants.3. We studied the relationships among seed production, seed predation ⁄ caching, cache pilferage and plant recruitment in a savanna-like landscape of oaks dispersed by scatter-hoarding rodents. 4. Our results show that acorn-dispersing rodents were concentrated under the canopies of scattered oaks, where the theft of cached acorns increased by 77% as compared to that of the surrounding open landscape. Acorns were thus cached selectively in the open areas to reduce pilferage; in fact, none of the few seeds cached beneath tree canopies survived predation by granivores (pilferage + recovery). Meanwhile, some acorns cached in the surrounding open areas were neither pilfered nor recovered and then recruited successfully. Accordingly, natural recruitment of newly emerged seedlings was higher outside than under canopies, suggesting that rodent caching strategies have direct implications for the directed dispersal of oaks. 5. Synthesis. The spatial patterns of seed dispersal shape the fitness of both the plant because they influence dispersal and recruitment efficiency, and the granivores that cache and predate its seeds because they influence their foraging efficiency. Cache protection strategies reduce pilferage significantly and enhance seed recovery rates by the cache owner. At the same time, more seeds remain dispersed and unrecovered. Thus, cache protection strategies can provide net benefits to the plant in terms of effective directed dispersal.

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Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one-sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate Á rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus ) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa . The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short-term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse-plants for the establishment of Holm oak seedlings in Mediterranean areas.

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1. Body size in parasitic insects can be subjected to contrasting selective pressures, especially if they complete their development within a single host. On the one hand, a larger body size is associated with a higher fitness. On the other hand, the host offers a discrete amount of resources, thus constraining the evolution of a disproportionate body size.2. The present study used the weevil Curculio elephas as a study model. Larvae develop within a single acorn, feeding on its cotyledons, and larval body size is strongly related to individual fitness.3. The relationship between larval and acorn size was negatively exponential. Larval growth was constrained in small acorns, which did not provide enough food for the weevils to attain their potential size. Larval size increased and levelled off in acorns over a certain size (inflexion point), in which cotyledons were rarely depleted. When there were more than one larva per acorn, a larger acorn was necessary to avoid food depletion.4. The results show that C. elephas larvae are sometimes endoparasitic, living on the edge of host holding capacity. If they were smaller they could avoid food depletion more easily, but the fitness benefits linked to a larger size have probably promoted body size increase. The strong negative effects of conspecific competition may have possibly influenced female strategy of laying a single egg per seed. 5. Being larger and fitter, but always within the limits of the available host sizes, may be one main evolutionary dilemma in endoparasites.Confl icts over Curculio larvae body size 305

Large-scale forest fragmentation can increase interpopulation genetic differentiation and erode the genetic variability of remnant plant populations. In this study, we analyze the extent of clonality and the genetic variability and structure within a holm oak (Quercus ilex) population from Central Spain at 3 patches showing different degrees of fragmentation. For this purpose, we have typed 191 individuals (105 adults and 86 saplings) at 9 microsatellite loci. Microsatellite markers revealed an extensive clonal structure in this species, with most analyzed clumps constituting a single "genet", which in some cases extended over a considerable area (up to 318 m(2)). The maximum distance between "ramets" tended to be higher in the extremely fragmented patch, suggesting that intensive management and environmental perturbation has favored clonal propagation. We have also found evidence that fragmentation has contributed to reduce genetic variability and increase genetic differentiation in holm oak saplings, indicating that the younger cohorts are suffering some negative genetic consequences of long-term population fragmentation. Finally, analyses of fine spatial genetic structure have revealed significant kinship structures up to 20-50 m that were particularly patent in the 2 less fragmented patches. Overall, our findings point to long-term genetic shifts in population structure of holm oaks in fragmented landscapes; however, further research is required on pollen dispersal and gene flow in this species.