Bottom-up regulation of a pelagic community through spatial aggregations

The importance of spatial pattern in ecosystems has long been recognized. However, incorporating patchiness into our understanding of forces regulating ecosystems has proved challenging. We used a combination of continuously sampling moored sensors, complemented by shipboard sampling, to measure the temporal variation, abundance and vertical distribution of four trophic levels in Hawaii's near shore pelagic ecosystem. Using an analysis approach from trophic dynamics, we found that the frequency and intensity of spatial aggregations-rather than total biomass-in each step of a food chain involving phytoplankton, copepods, mesopelagic micronekton and spinner dolphins (Stenella longirostris) were the most significant predictors of variation in adjacent trophic levels. Patches of organisms had impacts disproportionate to the biomass of organisms within them. Our results are in accordance with resource limitation-mediated by patch dynamics-regulating structure at each trophic step in this ecosystem, as well as the foraging behaviour of the top predator. Because of their high degree of heterogeneity, ecosystem-level effects of patchiness such as this may be common in many pelagic marine systems.     

Pelagic food webs: Responses to environmental processes and effects on the environment

Changes in both the environment and environmental research have led to the development of new protocols and approaches. These new approaches consider both the effects of changes in the global environment on living organisms (i.e. the responses of ecosystems to environmental processes) and the feedback responses of these organisms and ecosystems (i.e. the effects of living organisms on the environment). The present paper focuses on pelagic food webs in aquatic ecosystems. We examine three major effects of global environmental changes on aquatic organisms: (i) the release of pollutants and biological agents in lakes and nearshore marine waters; (ii) the loss of biodiversity and the collapse of commercially exploited resources that were heretofore renewable. We develop detailed examples of the effects of human activities on marine organisms (i.e. the effects of nutrient supply on the structure of pelagic food webs in marine systems. Finally, we examine (iii) the food-web-controlled exchanges of CO₂ between the atmosphere and the ocean, as a feedback effect of pelagic ecosystems on the global environment with respect to the ongoing climate change.     

水生态系统中宇宙发展现状

水生态系统中宇宙指人工构建的河流、湖泊、湿地、海洋等水生态系统近自然模拟装置,可以控制一定的物理、化学要素和生境条件,开展生物群落响应研究。中宇宙实验可以弥补传统实验室小型装置模拟要素过于简单,而野外现实系统复杂不可控制和难以开展对比实验的局限。本文列举了当前国际上多个科研机构长期运行的水生态系统中宇宙,将其归纳为流水型中宇宙、围隔装置、陆基模拟池、移动式水槽4类,各类中宇宙一般包括物理-生物模拟系统、自动调控系统、监测分析系统、中央控制系统。近年来,我国水生态系统的研究逐渐从单一环境要素向生态系统整体转变,对中宇宙模拟实验的需求愈加迫切。建议我国水生态系统中宇宙建设应提高结构复杂性,并与野外现场监测相结合,共同支撑更大尺度系统的整体模拟。     

Predictive models in spatially and temporally variable freshwater systems

Abstract This paper discusses the relationships between scaling and predictability in ecosystems. The logical basis of ecosystem modelling is explored using ideas first developed in complexity theory and analogies with the behaviour of complex adaptive systems. Any ecological model is a scale-dependent entity and both empirical and dynamic models of freshwater systems have their strengths and weaknesses. The logical basis of modelling using functional groups is explored. I conclude that such an approach can be justified and that such models have predictive power. Any predictive model of freshwater systems must take the major scales of external (atmospheric and catchment) forcing into account as well as the scales of key processes in the ecosystem itself. The importance of so-called ‘pink noise’ spectra, which arise both from external forcing and the internal dynamics of dynamic systems, is noted. The key scales of pattern and process in freshwater ecosystems are discussed in relation to the properties of the major functional groups. In order to have predictive power, I conclude that models of freshwater systems must include sediment exchanges and the properties of aquatic macrophytes as well as water column interactions and the pelagic components. When viewed at the scale of functional groups and the major biogeochemical processes, freshwater ecosystems may not be as complex as is often assumed.     

Carrion cycling in food webs: comparisons among terrestrial and marine ecosystems

In light of current global changes to ecosystem function (e.g. climate change, trophic downgrading, and invasive species), there has been a recent surge of interest in exploring differences in nutrient cycling among ecosystem types. In particular, a growing awareness has emerged concerning the importance of scavenging in food web dynamics, although no studies have focused specifically on exploring differences in carrion consumption between aquatic and terrestrial ecosystems. In this forum we synthesize the scavenging literature to elucidate differences in scavenging dynamics between terrestrial and marine ecosystems, and identify areas where future research is needed to more clearly understand the role of carrion consumption in maintaining ecosystem function within each of these environments. Although scavenging plays a similar functional role in terrestrial and aquatic food webs, here we suggest that several fundamental differences exist in scavenging dynamics among these ecosystem types due to the unique selection pressures imposed by the physical properties of water and air. In particular, the movement of carcasses in marine ecosystems (e.g. wave action, upwelling, and sinking) diffuses biological activity associated with scavenging and decomposition across large, three‐dimensional spatial scales, creating a unique spatial disconnect between the processes of production, scavenging, and decomposition, which in contrast are tightly linked in terrestrial ecosystems. Moreover, the limited role of bacteria and temporal stability of environmental conditions on the sea floor appears to have facilitated the evolution of a much more diverse community of macrofauna that relies on carrion for a higher portion of its nutrient consumption than is present in terrestrial ecosystems. Our observations are further discussed as they pertain to the potential impacts of climate change and trophic downgrading (i.e. removal of apex consumers from ecosystems) on scavenging dynamics within marine and terrestrial ecosystems.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

A review of climate-driven mismatches between interdependent phenophases in terrestrial and aquatic ecosystems

Mismatches in phenology between mutually dependent species, resulting from climate change, can have far-reaching consequences throughout an ecosystem at both higher and lower trophic levels. Rising temperatures, due to climate warming, have resulted in advances in development and changes in behaviour of many organisms around the world. However, not all species or phenophases are responding to this increase in temperature at the same rate, thus creating a disruption to previously synchronised interdependent key life-cycle stages. Mismatches have been reported between plants and pollinators, predators and prey, and pests and hosts. Here, we review mismatches between interdependent phenophases at different trophic levels resulting from climate change. We categorized the studies into (1) terrestrial (natural and agricultural) ecosystems, and (2) aquatic (freshwater and marine) ecosystems. As expected, we found reports of 'winners' and 'losers' in each system, such as earlier emergence of prey enabling partial avoidance of predators, potential reductions in crop yield if herbivore pests emerge before their predators and possible declines in marine biodiversity due to disruption in plankton-fish phenologies. Furthermore, in the marine environment rising temperatures have resulted in synchrony in a previously mismatched prey and predator system, resulting in an abrupt population decline in the prey species. The examples reviewed suggest that more research into the complex interactions between species in terrestrial and aquatic ecosystems is necessary to make conclusive predictions of how climate warming may impact the fragile balances within ecosystems in future.     

Complex wasp-waist regulation of pelagic ecosystems in the Pacific Ocean

‘Wasp-waist’ control of marine ecosystems is driven by a combination of top-down and bottom-up forcing by a few abundant short-lived species occupying intermediate trophic levels that form a narrow ‘waist’ through which energy flow from low to high trophic levels is controlled. It has been assumed that wasp-waist control occurs primarily in highly productive and species-poor systems (e.g. upwelling regions). Two large, species-rich, pelagic ecosystems in the relatively oligotrophic eastern and western Pacific Ocean also show wasp-waist-like structure, in that short-lived and fast-growing cephalopods and fishes at intermediate trophic levels comprise the vast majority of the biomass. Possible forcing dynamics of these systems were examined using ecosystem models by altering the biomass of phytoplankton (bottom-up forcing), large pelagic predators (top-down forcing), and intermediate ‘wasp-waist’ functional groups independently and observing how these changes propagated throughout the ecosystem. The largest effects were seen when altering the biomass of mid trophic-level epipelagic and mesopelagic fishes, where dramatic trophic cascades occurred both upward and downward in the system. We conclude that the high productivity and standing biomass of animals at intermediate trophic levels has a strong top-down influence on the abundance of primary producers. Furthermore, their importance as prey for large predators results in bottom-up controls on populations at higher trophic levels. We show that these tropical pelagic ecosystems possess a complex structure whereby several waist groups and alternate trophic pathways from primary producers to apex predators can cause unpredictable effects when the biomasses of particular functional groups are altered. Such models highlight the possible structuring mechanisms in pelagic systems, which have implications for fisheries that exploit these wasp-waist groups, such as squid fisheries, as well as for fisheries of top predators such as tunas and billfishes that prey upon wasp-waist species.     

东太平洋中部中上层鲨鱼群落营养生态位分化

鲨鱼在大洋生态系统中占据着重要的生态地位,其作为顶级捕食者,通过下行效应直接影响生态系统的稳定.稳定同位素技术是目前研究摄食生态学强有力的手段之一,可利用碳氮稳定同位素在食物网中的特性分别指示鲨鱼的食物来源和营养级.本研究选取8种130尾采集自东太平洋中部的中上层鲨鱼,应用稳定同位素绘制其种群生态位图谱,比较不同种群间的生态地位及资源分配方式上的差异.结果表明:不同鲨鱼种群碳、氮稳定同位素比值存在显著差异;8种鲨鱼在东太平洋生态系统中的营养级为4.3~5.4,大青鲨、尖吻鲭鲨与其他6种鲨鱼存在摄食隔离,表现出独特的营养生态地位.这些结果充分证明大洋性中上层鲨鱼并非生态系统的冗余种,其营养生态位的独特性不会被其他捕食者简单地替代和弥补.     

Identifying the dominant process (physical versus biological) in pelagic marine ecosystems from field estimates of chlorophyll a and phytoplankton production

A new approach is described to identify the dominant process (physical versus biological) in a pelagic marine ecosystem, from simple biological oceanographic field variables. The approach is based on quantification of the matching (M) between phytoplankton production (P) and losses, from field estimates of chlorophyll a(Chl) and P. Coefficient M is estimated for a wide range of oceanic and coastal environments and of trophic characteristics, using dat from the literature. Results show that M characterizes the dominance of physical versus biological processes in pelagic systems. The coefficient may be especially useful as a means for extracting process information on pelagic marine ecosystems from large data sets of Chl and P, e.g. recorded by moored instruments or provided by satellite images of ocean colour.      

Climate effects on fish body size–trophic position relationship depend on ecosystem type

The energetic demand of consumers increases with body size and temperature. This implies that energetic constraints may limit the trophic position of larger consumers, which is expected to be lower in tropical than in temperate regions to compensate for energy limitation. Using a global dataset of 3635 marine and freshwater ray‐finned fish species, we addressed if and how climate affects the fish body size–trophic position relationship in both freshwater and marine ecosystems, while controlling for the effects of taxonomic affiliation. We observed significant fish body size–trophic position relationships for different ecosystems. However, only in freshwater systems larger tropical fish presented a significantly lower trophic position than their temperate counterparts. Climate did not affect the fish body size–trophic position relationship in marine systems. Our results suggest that larger tropical freshwater fish may compensate for higher energetic constraints feeding at lower trophic positions, compared to their temperate counterparts of similar body size. The lower latitudinal temperature range in marine ecosystems and/or their larger ecosystem size may attenuate and/or compensate for the energy limitation of larger marine fish. Based on our results, temperature may determine macroecological patterns of aquatic food webs, but its effect is contingent on ecosystem type. We suggest that freshwater ecosystems may be more sensitive to warming‐induced alterations in food web topology and food chain length than marine ecosystems.     

Assessing the trophic impacts of marine mammals: From metabolism to food web indices

Marine mammals are an important part of ecosystems, and their trophic role and potential impact have been increasingly studied. One key question is how these large animals interact with fisheries or compete for similar resources. Consequently, some models once used only for fisheries management are now including pinnipeds and cetaceans. However, fish and marine mammals do not share the same ecology and bioenergetics, and complex ecosystem models may not be the best way to assess the impact of pinnipeds or cetaceans in food webs. Indeed, simpler methods based on thermodynamics might give us reasonable answers with limited amounts of data. Here, we present an assessment of two different approaches to assess the trophic role of marine mammals in the northern Gulf of St. Lawrence (Canada): mixed trophic impacts (MTI) based on ecosystem modeling and surface index (SI) impact based on bioenergetics. Our results show that while modeling represents a good way of getting a holistic view of the role of marine mammals in ecosystems, trophic impact estimates based on fundamental thermodynamics principles can also give us answers requiring less data. The body surface area approach presented here might provide a practical tool for ecologists, who are not necessarily ecosystem modelers, to study this issue.     

Biodiversity effects on ecosystem functioning: insights from aquatic systems

Unprecedented rates of species extinctions have prompted extensive research into the consequences of biodiversity losses on ecosystem functioning. While aquatic species are most threatened, research with freshwater and marine model systems has lagged behind progress made in terrestrial environments. This editorial to a special feature summarizes the main outcomes of a conference aimed at setting the stage for exploring the potential of aquatic systems to assess the role of biodiversity in ecosystem functioning. This series of papers proposes fresh approaches to the study of biodiversity effects on ecosystem functioning, outlines a new way of analyzing experimental data, presents a model that considers scale as an important factor determining outcomes, explores the effects of multiple stressors on species richness and ecosystem processes, and develops a food-web perspective that relates ecosystem properties to biodiversity. An insightful synthesis of lessons learned from aquatic systems is premature at present, but the papers clearly demonstrate the role that marine and freshwater systems can play in resolving open questions. The implications go well beyond the biodiversity in, and functioning of, ecosystems shaped by free-flowing or standing water.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

Death, detritus, and energy flow in aquatic ecosystems

1. Pelagic trophic structure and energy fluxes are evaluated predominantly on the basis of ingestion of particulate organic matter by living organisms and the effects of consumption on the population dynamics of trophic levels. 2. Population fluxes are not representative of the material and energy fluxes of either the composite pelagic region or the lake ecosystem. Metabolism of particulate and especially dissolved organic detritus from many pelagic and non-pelagic autochthonous and from allochthonous sources dominates both material and energy fluxes. Because of the very large magnitudes and relative chemical recalcitrance of these detrital sources, the large but slow metabolism of detritus provides an inherent ecosystem stability that energetically dampens the ephemeral, volatile fluctuations of higher trophic levels. 3. The annual time period is the only meaningful interval in comparative quantitative analyses of material and energy fluxes at population, community, and ecosystem levels. 4. Non-predatory death and metabolism by prokaryotic and protistian heterotrophs dominate. Continued application of animal-orientated relationships to the integrated, process-driven couplings of the aquatic ecosystems impedes understanding of quantitative ecosystem pathways and control mechanisms.     

Overfishing and the Replacement of Demersal Finfish by Shellfish: An Example from the English Channel

The worldwide depletion of major fish stocks through intensive industrial fishing is thought to have profoundly altered the trophic structure of marine ecosystems. Here we assess changes in the trophic structure of the English Channel marine ecosystem using a 90-year time-series (1920–2010) of commercial fishery landings. Our analysis was based on estimates of the mean trophic level (mTL) of annual landings and the Fishing-in-Balance index (FiB). Food webs of the Channel ecosystem have been altered, as shown by a significant decline in the mTL of fishery landings whilst increases in the FiB index suggest increased fishing effort and fishery expansion. Large, high trophic level species (e.g. spurdog, cod, ling) have been increasingly replaced by smaller, low trophic level fish (e.g. small spotted catsharks) and invertebrates (e.g. scallops, crabs and lobster). Declining trophic levels in fisheries catches have occurred worldwide, with fish catches progressively being replaced by invertebrates. We argue that a network of fisheries closures would help rebalance the trophic status of the Channel and allow regeneration of marine ecosystems.     

Nutrient reduction and climate change cause a potential shift from pelagic to benthic pathways in a eutrophic marine ecosystem

The degree to which marine ecosystems may support the pelagic or benthic food chain has been shown to vary across natural and anthropogenic gradients for e.g., in temperature and nutrient availability. Moreover, such external forcing may not only affect the flux of organic matter but could trigger large and abrupt changes, i.e., trophic cascades and ecological regime shifts, which once having occurred may prove potentially irreversible. In this study, we investigate the state and regulatory pathways of the Kattegat; a eutrophied and heavily exploited marine ecosystem, specifically testing for the occurrence of regime shifts and the relative importance of multiple drivers, e.g., climate change, eutrophication and commercial fishing on ecosystem dynamics and trophic pathways. Using multivariate statistics and nonlinear regression on a comprehensive data set, covering abiotic factors and biotic variables across all trophic levels, we here propose a potential regime shift from pelagic to benthic regulatory pathways; a possible first sign of recovery from eutrophication likely triggered by drastic nutrient reductions (involving both nitrogen and phosphorus), in combination with climate‐driven changes in local environmental conditions (e.g., temperature and oxygen concentrations).     

Trophic and functional cascades in tropical versus temperate aquatic microcosms

Inverse trophic cascades are a well explored and common consequence of the local depletion or extinction of top predators in natural ecosystems. Despite a large body of research, the cascading effects of predator removal on ecosystem functions are not as well understood. Developing microcosm experiments, we explored food web changes in trophic structure and ecosystem functioning following biomass removal of top predators in representative temperate and tropical rock pool communities that contained similar assemblages of zooplankton and benthic invertebrates. We observed changes in species abundances following predator removal in both temperate and tropical communities, in line with expected inverse effects of a trophic cascade, where predation release benefits the predator’s preys and competitors and impacts the preys of the latter. We also observed several changes at the community and ecosystem levels including a decrease in total abundance and mean trophic level of the community, and changes in chlorophyll-a and total dissolved particles. Our results also showed an increase in variability of both community and ecosystem processes following the removal of predators. These results illustrate how predator removal can lead to inverse trophic cascades both in structural and functioning properties, and can increase variability of ecosystem processes. Although observed patterns were consistent between tropical and temperate communities following an inverse cascade pattern, changes were more pronounced in the temperate community. Therefore, aquatic food webs may have inherent traits that condition ecosystem responses to changes in top-down trophic control and render some aquatic ecosystems especially sensitive to the removals of top predators.     

Evaluating trophic cascades as drivers of regime shifts in different ocean ecosystems

In ecosystems that are strongly structured by predation, reducing top predator abundance can alter several lower trophic levels—a process known as a trophic cascade. A persistent trophic cascade also fits the definition of a regime shift. Such ‘trophic cascade regime shifts'' have been reported in a few pelagic marine systems—notably the Black Sea, Baltic Sea and eastern Scotian Shelf—raising the question of how common this phenomenon is in the marine environment. We provide a general methodology for distinguishing top-down and bottom-up effects and apply this methodology to time series from these three ecosystems. We found evidence for top-down forcing in the Black Sea due primarily to gelatinous zooplankton. Changes in the Baltic Sea are primarily bottom-up, strongly structured by salinity, but top-down forcing related to changes in cod abundance also shapes the ecosystem. Changes in the eastern Scotian Shelf that were originally attributed to declines in groundfish are better explained by changes in stratification. Our review suggests that trophic cascade regime shifts are rare in open ocean ecosystems and that their likelihood increases as the residence time of water in the system increases. Our work challenges the assumption that negative correlation between consecutive trophic levels implies top-down forcing.     

Exploring the dynamics of ecological indicators using food web models fitted to time series of abundance and catch data

Previously, standardized snap-shot models of the Southern Benguela (1980–1989), Southern Humboldt (1992) and Southern Catalan Sea (1994) ecosystems were examined and found to facilitate assessment of ecosystem characteristics related to the gradient in exploitation status of the ecosystems; highest level of exploitation in the South Catalan Sea (North-western Mediterranean), high in the Southern Humboldt and lower in the Southern Benguela. Subsequently, these models were calibrated and fitted using available catch, fishing effort/mortality and abundance data series and incorporated environmental and internal drivers. This study furthers the previous comparative analyses by comparing changes in ecosystem structure using a selection of ecosystem indicators from the calibrated models and assessing how these indicators change over time in these three contrasting ecosystems. Indicators examined include community turnover rates (production/biomass), trophic level of landings and the community, biodiversity indicators, ratios of predatory/forage fish and pelagic/demersal fish biomass, catch ratios, and network analysis indicators. Using the set of model-derived indicators, the three ecosystems were ranked in terms of exploitation level. This ranking was performed using the values of these indicators in recent years (ecosystem state) as well as their trends over time (ecosystem trend). The non-parametric Kruskal–Wallis and Median tests were used to test for significance of the difference between indicators from the three ecosystems in the last 5 years of the simulation to compare present ecosystem states. We compared the slope of the lineal trend and its significance between ecosystems using the generalized least-squares regression taking auto-correlation into consideration to analyse ecosystem trends. The indicators that capture better the high impacts of fishing prevalent in the Mediterranean and Humboldt ecosystems, and the more conservative exploitation of the Southern Benguela, are the fish/invertebrates biomass and catch ratio, the demersal/pelagic fish biomass and catch ratio (depending on the ecosystem and the fishery being developed), flows to detritus, and the mean trophic level of the community (when large, poorly quantified groups such as zooplankton and detritus are excluded). This study suggests that the best option for classifying ecosystems according to the impact of fishing is to consider a broad range of indicators to understand how and why an ecosystem is responding to particular environmental or fishing drivers (or more likely a combination of these). Our results highlight the importance of including indicators capturing trends over time as well as recent ecosystem states. We also identified 23 pairs of indicators that correlated similarly in the three ecosystems (they showed a significant correlation with same sign). Further comparisons may contribute towards generalization of this list, progressing towards a better understanding of the behaviour of ecological indicators.