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Natural grassland ecosystems are increasingly threatened by excessive loadings of nutrients and by the presence of species bred for high productivity. By manipulating grazing regimes and nutrient availability, agricultural practices facilitate the establishment and spread of certain forage plant species outside managed landscapes, challenging local biodiversity. The ecological success of some species in the invaded range sometimes seems to be associated with the symbiosis with foliar fungal endophytes. Symbiotic fungi may increase the competitiveness of host species, but also the resistance to herbivory through the production of toxic secondary compounds such as alkaloids. While progress has been made in understanding how soil nutrients modulate other benefits offered by fungal endophytes to plants (e.g. stress tolerance, competitive ability, etc), the consequences for a higher trophic level (i.e. herbivores) and the potential feedbacks on plant invasion have not been explored yet.We explored the relative and interactive importance of soil nitrogen (N) and phosphorus (P) in modulating the interaction of the invasive grass tall fescue—associated with fungal endophytes—and native herbivores in a natural grassland. We hypothesized that N and P nutrients modulate differentially leaf quality traits, namely nutritional value and fungal alkaloid contents, determining the level of damage by native insect herbivores on the exotic tall fescue.We found that only the addition of P significantly increased native caterpillar density in the field, which corresponded to a concomitant increase in leaf damage. In contrast to expectations, the concentration of the alkaloid ergovaline in leaves was not strongly related to N. It was the level of soil P which dictated the concentration of the element (P) in the leaves and reduced the level of defence against herbivores in this endophyte‐symbiotic species. Then, herbivore performance increased, and plants were more prone to be attacked.Synthesis. Our study indicates a strong control of soil P fertility on the triple interaction among plants, fungal endophytes and native herbivores. This highlights the potential role of increased soil nutrients on the invasion spread of endophyte‐symbiotic forage plants in natural grasslands. [ABSTRACT FROM AUTHOR]

Introduced ungulates can alter understory structure and composition posing a serious threat to forest biodiversity. Yet how large-herbivore impacts in forested regions vary along major environmental gradients remains little explored. If ungulate effects shift with habitat conditions, then management could be tailored to protect most vulnerable forests. We tested the hypothesis that the extent of livestock impact on understory vegetation increases with habitat moisture across Nothofagus dombeyi forests in Nahuel Huapi National Park, NW Patagonia, Argentina. Understory composition and species diversity were compared for paired sites ( N = 5), which were historically used by cattle or remained free of livestock for more than 50 yr, and were located along a regional precipitation gradient (1500–2800 mm/yr). Long-term cattle presence reduced the cover of sub-canopy trees, shrubs and bamboo by 57–83%, and increased the relative cover of ground-layer herbs, regardless of habitat moisture. Livestock effects on species composition increased towards the wettest forests, which contained more species exclusive to either browsed or unbrowsed sites. Livestock presence increased species richness ( α diversity) and within-site heterogeneity ( β diversity) in some locations, but mostly reduced species evenness (30%) throughout the moisture gradient. Species turnover at the gradient scale was lower across browsed sites than for livestock-free sites. Our results indicate that the historical presence of domestic cattle induced region-wide changes in understory communities, highlighting the vulnerable nature of the local flora to ungulate disturbance. The greater impact of livestock browsing on the species composition of wetter forests was consistent with the role of plant growth–defence trade-offs along resource gradients. We suggest that the erosion of understory vegetation attributed to domestic herbivores in Patagonian beech forests can be mitigated by adjusting current animal stocks, while moister forests should be given the highest conservation priority. [ABSTRACT FROM AUTHOR]

1. Endemic herbivory can influence forest ecosystem function, but how annual productivity consumption relates to seasonal resource utilisation by folivore guilds remains poorly understood. 2. Monthly changes in leaf damage and foliage traits were monitored in 'dry' and 'wet' Nothofagus pumilio (Fagales: Nothofagaceae) deciduous forests in northern Patagonia, Argentina. Herbivore-induced leaf abscission was assessed and foliar productivity consumption was measured in the canopy and in litterfall harvests. 3. Seasonal damage ranged from 8% to 32% in dry forest, but remained below 5% in wet forest although foliar quality was higher in the latter. In dry forest, dominant guilds were temporally separated; leaf miners consumed younger foliage in spring to early summer, whereas leaf tiers prevailed in late summer to autumn. In wet forest, damage created by external chewers was concentrated in early summer. 4. Insect damage induced premature leaf abscission, especially in dry forest. Although foliar production in wet forest doubled that in dry forest, the percentage of productivity lost to folivores was higher in dry (14-20%) than in wet (1.2-1.8%) forest. 5. The overall greater impact of herbivory in dry forest canopies countered the expectation that consumption would increase with plant productivity and nutritional quality. Lower temperatures and a shorter growing season are likely to constrain folivory in wet forest stands. [ABSTRACT FROM AUTHOR]

Predators are thought to play a key role in controlling herbivory, thus having positive indirect effects on plants. However, evidence for terrestrial trophic cascades is still fragmentary, perhaps due to variation in top-down forces created by environmental heterogeneity. We examined the magnitude of predation effects on foliar damage by chewing insects and mean leaf size, by excluding birds from saplings in ‘dry’ and ‘wet’ Nothofagus pumilio forests in the northern Patagonian Andes, Argentina. The experiment lasted 2 years encompassing a severe drought during the La Niña phase of a strong El Niño/Southern Oscillation event, which was followed by unusually high background folivory levels. Insect damage was consistently higher in wet than in dry forest saplings. In the drought year (1999), bird exclusion increased folivory rates in both forests but did not affect tree leaf size. In the ensuing season (2000), leaf damage was generally twice as high as in the drought year. As a result, bird exclusion not only increased the extent of folivory but also significantly decreased sapling leaf size. The latter effect was stronger in the wet forest, suggesting compensation of leaf area loss by dry forest saplings. Overall, the magnitude of predator indirect effects depended on the response variable measured. Insectivorous birds were more effective at reducing folivory than at facilitating leaf area growth. Our results indicate that bird-initiated trophic cascades protect N. pumilio saplings from insect damage even during years with above-normal herbivory, and also support the view that large-scale climatic events influence the strength of trophic cascades. [ABSTRACT FROM AUTHOR]

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment. [ABSTRACT FROM AUTHOR]

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 (2000–2002) 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 (2001–2002), 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. [ABSTRACT FROM AUTHOR]