Despite the increasing evidence of drastic and profound changes in many ecosystems, often referred to as regime shifts, we have little ability to understand the processes that provide insurance against such change (resilience). Modelling studies have suggested that increased variance may foreshadow a regime shift, but this requires long-term data and knowledge of the functional links between key processes. Field-based research and ground-truthing is an essential part of the heuristic that marries theoretical and empirical research, but experimental studies of resilience are lagging behind theory, management and policy requirements. Empirically, ecological resilience must be understood in terms of community dynamics and the potential for small shifts in environmental forcing to break the feedbacks that support resilience. Here, we integrate recent theory and empirical data to identify ways we might define and understand potential thresholds in the resilience of nature, and thus the potential for regime shifts, by focusing on the roles of strong and weak interactions, linkages in meta-communities, and positive feedbacks between these and environmental drivers. The challenge to theoretical and field ecologists is to make the shift from hindsight to a more predictive science that is able to assist in the implementation of ecosystem-based management.

Invasive species and bottom-water hypoxia both constitute major global threats to the diversity and integrity of marine ecosystems. These stressors may interact with unexpected consequences, as invasive species that require an initial environmental disturbance to become established can subsequently become important drivers of ecological change. There is recent evidence that improved bottom-water oxygen conditions in coastal areas of the northern Baltic Sea coincide with increased abundances of the invasive polychaetes Marenzelleria spp. Using a reactive-transport model, we demonstrate that the long-term bioirrigation activities of dense Marenzelleria populations have a major impact on sedimentary phosphorus dynamics. This may facilitate the switch from a seasonally hypoxic system back to a normoxic system by reducing the potential for sediment-induced eutrophication in the upper water column. In contrast to short-term laboratory experiments, our simulations, which cover a 10-year period, show that Marenzelleria has the potential to enhance long-term phosphorus retention in muddy sediments. Over time bioirrigation leads to a substantial increase in the iron-bound phosphorus content of sediments while reducing the concentration of labile organic carbon. As surface sediments are maintained oxic, iron oxyhydroxides are able to persist and age into more refractory forms. The model illustrates mechanisms through which Marenzelleria can act as a driver of ecological change, although hypoxic disturbance or natural population declines in native species may be needed for them to initially become established. Invasive species are generally considered to have a negative impact; however, we show here that one of the main recent invaders in the Baltic Sea may provide important ecosystem services. This may be of particular importance in low-diversity systems, where disturbances may dramatically alter ecosystem services due to low functional redundancy. Thus, an environmental problem in one region may be either exacerbated or alleviated by a single species from another region, with potentially ecosystem-wide consequences.

Abstract. Predicting the dynamics of ecosystems requires an understanding of how trophic interactions respond to environmental change. In Antarctic marine ecosystems, food web dynamics are inextricably linked to sea ice conditions that affect the nature and magnitude of primary food sources available to higher trophic levels. Recent attention on the changing sea ice conditions in polar seas highlights the need to better understand how marine food webs respond to changes in such broad-scale environmental drivers. This study investigated the importance of sea ice and advected primary food sources to the structure of benthic food webs in coastal Antarctica. We compared the isotopic composition of several seafloor taxa (including primary producers and invertebrates with a variety of feeding modes) that are widely distributed in the Antarctic. We assessed shifts in the trophic role of numerically dominant benthic omnivores at five coastal Ross Sea locations. These locations vary in primary productivity and food availability, due to their different levels of sea ice cover, and proximity to polynyas and advected primary production. The d 15 N signatures and isotope mixing model results for the bivalves Laternula elliptica and Adamussium colbecki and the urchin Sterechinus neumeyeri indicate a shift from consumption of a higher proportion of detritus at locations with more permanent sea ice in the south to more freshly produced algal material associated with proximity to ice-free water in the north and east. The detrital pathways utilized by many benthic species may act to dampen the impacts of large seasonal fluctuations in the availability of primary production. The limiting relationship between sea ice distribution and in situ primary productivity emphasizes the role of connectivity and spatial subsidies of organic matter in fueling the food web. Our results begin to provide a basis for predicting how benthic ecosystems will respond to changes in sea ice persistence and extent along environmental gradients in the high Antarctic.

Size is a fundamental organismal trait and an important driver of ecosystem functions. Although large individuals may dominate some functions and provide important habitat structuring effects, intra-specific body size effects are rarely investigated in the context of BEF relationships. We used an in situ density manipulation experiment to explore the contribution of large, deep-burrowing bivalves to oxygen and nutrient fluxes across the sediment-water interface. By manipulating bivalve size structure through the removal of large individuals, we held species identity constant, but altered the trait characteristics of the community. The number of large bivalves was the best predictor of ecosystem functioning. Our results highlight that (a) accounting for body size provides important insights into the mechanisms underpinning biodiversity effects on ecosystem function, and (b) if local disturbances are recurrent, preventing individuals from reaching large sizes, the contribution of large adults may be lost, with largely unknown implications for ecosystem functionality.T here is clear evidence that losses in biodiversity reduce the efficiency of ecosystem functions, including productivity and nutrient cycling 1 , but the actual mechanisms that underpin the positive biodiversityecosystem function (BEF) relationships remain an area of intense research 2,3 . A few species (with unique traits) may in fact dominate certain ecosystem processes [4][5][6] , and recent meta-analyses have indeed shown that species identity effects may be as important as richness effects per se 7 . Improving the mechanistic understanding of BEF relationships and allow prediction of the magnitude of change in ecosystem function following the loss of particular traits 2,8 thus depends on the nature of the ecosystem function(s) of interest and trait composition not only across the resident community but also within species. Trait-based approaches have, however, largely focused on assigning traits to species rather than individuals 9 . Hence, while interspecific differences in trait composition and the subsequent functional contribution to ecosystem processes have been acknowledged, differences in intra-specific trait characteristics are rarely addressed 10 (but see e.g. [11][12][13] ).Body mass is a fundamental organism trait that affects metabolic rate, energy demand and uptake rate 9,14,15 , and is an important characteristic of overall population and community structure through density-mass allometric relationships 14 . Even though high numbers of small individuals may dominate specific ecosystem functions through rapid turnover rates, the fewer large individuals may dominate other functions and provide important habitat structuring effects 16 . Particularly where ecosystem functions relate to the generation of biogenic habitat or organisms mediation of the nature and flux of energy and matter, size matters. This is potentially very important in marine sediments where large organisms can be expected to displace more sediment, pump more water an...