T. 2006. Trophic and non-trophic pathways mediate apparent competition through post-dispersal seed predation in a Patagonian mixed forest. Á Oikos 113: 469 Á480.Post-dispersal predation can be a major source of seed loss in temperate forests. Little is known, however, about how predator-mediated indirect interactions such as apparent competition alter survival patterns of canopy tree seeds. Understorey plants may enhance tree seed predation by providing sheltered habitat to granivores (non-trophic pathway). In addition, occurrence of different tree seeds in mixed patches may lead to short-term apparent competition between seed types, because of the granivores' foraging response to changes in food patch quality (trophic pathway). We hypothesised that understorey bamboo cover and mixing of seed species in food patches would both increase tree seed predation in a Nothofagus dombeyi(/Austrocedrus chilensis forest in northern Patagonia, Argentina. Seed removal experiments were conducted for three consecutive years (2000 Á2002) differing in overall granivory rates. Seed patch encounter and seed removal rates were consistently higher for the larger and more nutritious Austrocedrus seeds than for the smaller Nothofagus seeds. Seed removal was greater beneath bamboo than in open areas. This apparent competition pathway was stronger in a low-predation year (2000) than in high-predation years (2001 Á2002), suggesting a shift in microhabitat use by rodents. Patch composition had a significant, though weaker, impact on seed survival across study years, whereas seed density per patch enhanced encounter rates but did not influence seed removal. Removal of the less-preferred Nothofagus seeds increased in the presence of Austrocedrus seeds, but the reciprocal indirect effect was not observed. However, this non-reciprocal apparent competition between seed species was only significant in the high-predation years. Our study shows that granivore-mediated indirect effects can arise through different interaction pathways, affecting seed survival patterns according to the predator's preference for alternative seed types. Moreover, results indicate that the occurrence and relative strength of trophic vs non-trophic pathways of apparent competition may change under contrasting predation scenarios.

Plant cover plays a major role in shaping the nature of recruitment microsites through direct (resource mediated) and indirect (consumer mediated) interactions. Understorey plants may differentially affect seedling establishment, thus contributing to regeneration-niche separation among canopy tree species. We examined patterns of early tree seedling survival resulting from interactive effects of understorey bamboo (Chusquea culeou) and resident consumers in a mixed temperate Patagonian forest, Argentina. Newly germinated seedlings of Nothofagus dombeyi and Austrocedrus chilensis were planted in bamboo thickets and non-bamboo patches, with or without small-vertebrate exclosures. We found species-specific patterns of seedling survival in relation to bamboo cover. Nothofagus survival was generally low but increased under bamboo, irrespective of cage treatment. Desiccation stress accounted for most Nothofagus mortality in open, non-bamboo areas. In contrast, Austrocedrus survival was highest in non-bamboo microsites, as most seedlings beneath bamboo were killed by small vertebrates through direct consumption or non-trophic physical damage. There was little evidence for a negative impact of bamboo on tree seedling survival attributable to resource competition. The balance of simultaneous positive and negative interactions implied that bamboo presence facilitated Nothofagus early establishment but inhibited Austrocedrus recruitment via apparent competition. These results illustrate the potential for dominant understorey plants to promote microsite segregation during early stages of recruitment between tree seedlings having different susceptibilities to water stress and herbivory. We recognise, however, that patterns of bamboo-seedling interactions may be conditional on moisture levels and consumer activity during establishment. Hence, both biotic and abiotic heterogeneity in understorey environments should be incorporated into conceptual models of regeneration dynamics and tree coexistence in forest communities.

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.

The factors and processes that regulate Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) natural regeneration in southwestern Argentina are unknown. We investigated (i) the germination responses of P. menziesii seeds to variations in the radiation and thermal environments imposed by canopy cover and litter, (ii) the effects of litter on seedling emergence, and (iii) the effects of understory vegetation on seedling performance and seed and seedling predation. Seed germination was high in darkness and under canopies. Under a combination of canopy types and litter covers, germination showed a negative correlation with temperature between 23 and 40°C. Germination was not altered by changes in red/far-red ratio. These results suggest that the presence of an overstory canopy promotes P. menziesii seed germination compared with a no-canopy situation (e.g., a clearcut). Litter inhibited seedling emergence by reducing water availability and by acting as a mechanical barrier. Understory vegetation (established saplings) reduced seedling growth and survival. However, our results suggest that direct competition between seedlings and understory vegetation is not the principal force regulating regeneration; indirect effects appear to be of paramount importance because P. menziesii seed and seedling consumption (presumably by rodents) is greatly encouraged by the presence of dense vegetation patches in the understory.

Summary1. Episodic mass flowering and subsequent die-off of bamboo understories may generate rare opportunities for tree regeneration by altering consumer-seedling interactions as much as by increasing light availability to seedlings. We hypothesized that bamboo dieback interacts with canopy neighbourhood composition in creating recruitment microsites for tree seedling species with varied shade tolerance and susceptibility to herbivory. 2. We conducted a 2-year experiment in a Patagonian mixed forest altered by extensive, but patchy dieback of the bamboo Chusquea culeou. Newly emerged seedlings of Nothofagus alpina (more shade-tolerant) and Nothofagus dombeyi (less shade-tolerant) were planted in conspecific and heterospecific canopy neighbourhoods, with either a flowered (dead) or non-flowered (live) bamboo understorey. Seedlings were placed inside and outside mesh cages to assess mortality from vertebrate or invertebrate consumers. 3. Vertebrate exclusion increased seedling survival regardless of bamboo condition. Seedling loss to invertebrates decreased with bamboo die-off, resulting in higher survival of N. alpina in dead than in live patches. In contrast, bamboo die-off increased N. dombeyi mortality by wilting, which counteracted the benefits of seedling release from consumers. Bamboo die-off increased light availability and enhanced seedling growth for both species. 4. Nothofagus alpina seedlings were less damaged or killed by invertebrates under heterospecific canopies than under conspecifics (associational resistance), whereas N. dombeyi performance was unaffected by neighbourhood composition. Bamboo die-off did not change seedling performance patterns observed across canopy neighbourhoods with live bamboo understories. 5. Synthesis. Gaps created by bamboo die-off can exert both positive and negative, species-specific effects on the likelihood of tree seedling establishment. We conclude that infrequent understorey disturbances coupled with canopy neighbourhood effects mediated by seedling herbivores may drive gap-phase succession within old-growth forests.