Jordi F. Pagès

Combined effects of fragmentation and herbivory on Posidonia oceanica seagrass ecosystems

Summary 1. Fragmentation is a major agent for seagrass meadow decline, yet little is known about how it interacts with processes like herbivory, an important functional driver of seagrass meadows. The interaction with external stressors like fragmentation could exacerbate the effects of internal ecosystem drivers like herbivory, with distinct implications for ecosystem management. 2. We used manipulative field experiments to assess these interactive effects in two Posidonia oceanica seagrass meadows. We monitored replicated plots in small and large patches in two meadows suffering fragmentation with and without herbivores (using exclusion cages) to test whether fragment size and herbivory could act together to alter ecosystem functioning. We measured changes in defoliation rates, primary production, canopy height and nutrient content in all plots after 4 months of herbivore exclusion. 3. Our results show that herbivores increased defoliation rates resulting in reduced primary production, nutrient content and canopy structure (canopy height). Patch size (fragment) on its own also reduced primary production, nutrient content and canopy structure. We also observed significant additive interactions between herbivores and fragmentation on canopy structure and production responses. In addition, small patches showed nutrient limitation but were able to accumulate more carbohydrate reserves, probably due to a higher light availability. This may explain why small patches can persist under significant herbivore pressure. 4. Synthesis. While fragmentation has already been identified as an important external agent of seagrass decline, the combination of fragmentation and herbivory can seriously exacerbate structural losses and affect primary production, profoundly compromising the role of seagrasses as habitatforming ecosystems. These interactions between external stressors and internal drivers may result in large unexpected consequences that may flow on to the rest of the ecosystem.

Matrix composition and patch edges influence plant–herbivore interactions in marine landscapes

Summary The functioning of ecosystems can be strongly driven by landscape attributes. Despite its importance, however, our understanding of how landscape influences ecosystem function derives mostly from species richness and abundance patterns, with few studies assessing how these relate to actual functional rates. We examined the influence of landscape attributes on the rates of herbivory in seagrass meadows, where herbivory has been identified as a key process structuring these relatively simple systems. The study was conducted in three representative Posidonia oceanica meadows. The principal herbivores in these meadows are the fish Sarpa salpa and the sea urchin Paracentrotus lividus, and we hypothesized that differences in their interaction with landscape attributes would significantly influence herbivory rates. We measured herbivore abundance, herbivory rates, primary production and plant quality (C:N) in seagrass patches embedded either in rock or in sand (matrix attribute), in patches either near or far from a rocky reef (distance attribute) and at the edges and interior of meadows. Our results show that matrix and meadow edges significantly affected the actual levels of herbivory. Herbivory rates were higher in seagrass patches embedded in a rocky matrix compared to those on sand, and herbivory at the centre of seagrass meadows was higher than at the edges. In contrast, patch distance to rocky reefs did not affect herbivory. Neither herbivore abundance nor food quality explained the patterns across different landscape attributes. This suggests that variation in herbivory across the landscape may be related much more to behavioural differences between species in their evaluation of risk, movement and food preference in relation to the landscape structure. Our results indicate that richness and abundance patterns may mask critical interactions between landscape attributes and species responses, which result in considerable heterogeneity in the way key functional processes like herbivory are distributed across the ecosystem mosaic.

Should we sync? Seascape‐level genetic and ecological factors determine seagrass flowering patterns

Spatial and temporal heterogeneity in flowering occur in many plant species with abiotic pollination and may confer fitness advantages through mechanisms such as predator satiation or pollination efficiency. Environmental factors such as light quality or quantity and temperature play an important role in inducing synchronization on wide geographic scales. On a smaller geographic scale, external factors such as resource availability and herbivory are theorized to trigger flowering, while genetic factors may also play an important role. In this study, we assessed the importance of ecological and genetic factors in shaping seascape-level spatial heterogeneity in flowering of the seagrass Posidonia oceanica. By investigating spatially close sites (

The Mediterranean benthic herbivores show diverse responses to extreme storm disturbances.

Catastrophic storms have been observed to be one of the major elements in shaping the standing structure of marine benthic ecosystems. Yet, little is known about the effect of catastrophic storms on ecosystem processes. Specifically, herbivory is the main control mechanism of macrophyte communities in the Mediterranean, with two main key herbivores: the sea urchin Paracentrotus lividus and the fish Sarpa salpa. Consequently, the effects of extreme storm events on these two herbivores (at the population level and on their behaviour) may be critical for the functioning of the ecosystem. With the aim of filling this gap, we took advantage of two parallel studies that were conducted before, during and after an unexpected catastrophic storm event. Specifically, fish and sea urchin abundance were assessed before and after the storm in monitored fixed areas (one site for sea urchin assessment and 3 sites for fish visual transects). Additionally, we investigated the behavioural response to the disturbance of S. salpa fishes that had been tagged with acoustic transmitters. Given their low mobility, sea urchins were severely affected by the storm (ca. 50% losses) with higher losses in those patches with a higher density of sea urchins. This may be due to a limited availability of refuges within each patch. In contrast, fish abundance was not affected, as fish were able to move to protected areas (i.e. deeper) as a result of the high mobility of this species. Our results highlight that catastrophic storms differentially affect the two dominant macroherbivores of rocky macroalgal and seagrass systems due to differences in mobility and escaping strategies. This study emphasises that under catastrophic disturbances, the presence of different responses among the key herbivores of the system may be critical for the maintenance of the herbivory function.

Immanent conditions determine imminent collapses: nutrient regimes define the resilience of macroalgal communities

Predicting where state-changing thresholds lie can be inherently complex in ecosystems characterized by nonlinear dynamics. Unpacking the mechanisms underlying these transitions can help considerably reduce this unpredictability. We used empirical observations, field and laboratory experiments, and mathematical models to examine how differences in nutrient regimes mediate the capacity of macrophyte communities to sustain sea urchin grazing. In relatively nutrient-rich conditions, macrophyte systems were more resilient to grazing, shifting to barrens beyond 1 800 g m−2 (urchin biomass), more than twice the threshold of nutrient-poor conditions. The mechanisms driving these differences are linked to how nutrients mediate urchin foraging and algal growth: controlled experiments showed that low-nutrient regimes trigger compensatory feeding and reduce plant growth, mechanisms supported by our consumer–resource model. These mechanisms act together to halve macrophyte community resilience. Our study demonstrates that by mediating the underlying drivers, inherent conditions can strongly influence the buffer capacity of nonlinear systems.

Contrasting effects of ocean warming on different components of plant-herbivore interactions

There is increasing uncertainty of how marine ecosystems will respond to rising temperatures. While studies have focused on the impacts of warming on individual species, knowledge of how species interactions are likely to respond is scant. The strength of even simple two-species interactions is influenced by several interacting mechanisms, each potentially changing with temperature. We used controlled experiments to assess how plant-herbivore interactions respond to temperature for three structural dominant macrophytes in the Mediterranean and their principal sea urchin herbivore. Increasing temperature differentially influenced plant-specific growth, sea urchin growth and metabolism, consumption rates and herbivore preferences, but not movement behaviour. Evaluating these empirical observations against conceptual models of plant-herbivore performance, it appears likely that while the strength of herbivory may increase for the tested macroalga, for the two dominant seagrasses, the interaction strength may remain relatively unchanged or even weaken as temperatures rise. These results show a clear set of winners and losers in the warming Mediterranean as the complex factors driving species interactions change.

The richness of small pockets: Decapod species peak in small seagrass patches where fish predators are absent

Patchy landscapes behave differently from continuous ones. Patch size can influence species behaviour, movement, feeding and predation rates, with flow-on consequences for the diversity of species that inhabit these patches. To understand the importance of patchiness on regional species pools, we measured decapod richness and abundance in several seagrass patches with contrasting sizes. Additionally, we evaluated potential drivers of patch-specific species distribution including resource abundance, predator habitat use and the structural complexity of patches. Our results showed a non-random distribution of decapod species: small patches were clear hotspots of diversity and abundance, particularly of larger-bodied epifaunal decapods. Interestingly, these hotspots were characterized by lower nutrient resources, lower canopy height, but also lower predator use. Small fish invertivores such as Coris julis and several species of Symphodus were mostly restricted to large patches. These resident predators may be critical in clumping predation in large patches with consequences for how biodiversity of their prey is distributed across the seascape. Our results highlight the idea that a habitat mosaic with both large and small seagrass patches would potentially bolster biodiversity because preys and predators may seek refuge in patches of different sizes.