Climate-induced changes in bottom-up and top-down processes independently alter a marine ecosystem

Climate change has complex structural impacts on coastal ecosystems. Global warming is linked to a widespread decline in body size, whereas increased flood frequency can amplify nutrient enrichment through enhanced run-off. Altered population body-size structure represents a disruption in top-down control, whereas eutrophication embodies a change in bottom-up forcing. These processes are typically studied in isolation and little is known about their potential interactive effects. Here, we present the results of an in situ experiment examining the combined effects of top-down and bottom-up forces on the structure of a coastal marine community. Reduced average body mass of the top predator (the shore crab, Carcinus maenas) and nutrient enrichment combined additively to alter mean community body mass. Nutrient enrichment increased species richness and overall density of organisms. Reduced top-predator body mass increased community biomass. Additionally, we found evidence for an allometrically induced trophic cascade. Here, the reduction in top-predator body mass enabled greater biomass of intermediate fish predators within the mesocosms. This, in turn, suppressed key micrograzers, which led to an overall increase in microalgal biomass. This response highlights the possibility for climate-induced trophic cascades, driven by altered size structure of populations, rather than species extinction.     

Climate change and commercial fishing practices codetermine survival of a long-lived seabird

Understanding the environmental mechanisms that govern population change is a fundamental objective in ecology. Although the determination of how top-down and bottom-up drivers affect demography is important, it is often equally critical to understand the extent to which, environmental conditions that underpin these drivers fluctuate across time. For example, associations between climate and both food availability and predation risk may suggest the presence of trophic interactions that may influence inferences made from patterns in ecological data. Analytical tools have been developed to account for these correlations, while providing opportunities to ask novel questions regarding how populations change across space and time. Here, we combine two modeling disciplines—path analysis and mark-recapture-recovery models—to explore whether shifts in sea-surface temperatures (SSTs) influenced top-down (entanglement in fishing equipment) or bottom-up (forage fish production) population constraints over 60 years, and the extent to which these covarying processes shaped the survival of a long-lived seabird, the Royal tern. We found that hemispheric trends in SST were associated with variation in the amount of fish harvested along the Atlantic coast of North America and in the Caribbean, whereas reductions in forage fish production were mostly driven by shifts in the amount of fish harvested by commercial fisheries throughout the North Atlantic the year prior. Although the indirect (i.e., stock depletion) and direct (i.e., entanglement) impacts of commercial fishing on Royal tern mortality has declined over the last 60 years, increased SSTs during this time period has resulted in a comparable increase in mortality risk, which disproportionately impacted the survival of the youngest age-classes of Royal terns. Given climate projections for the North Atlantic, our results indicate that threats to Royal tern population persistence in the Mid-Atlantic will most likely be driven by failures to recruit juveniles into the breeding population.     

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.     

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.     

The ups and downs of trophic control in continental shelf ecosystems

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.     

It is the economy,stupid! Projecting the fate of fish populations using ecological–economic modeling

Four marine fish species are among the most important on the world market: cod, salmon, tuna, and sea bass. While the supply of North American and European markets for two of these species – Atlantic salmon and European sea bass – mainly comes from fish farming, Atlantic cod and tunas are mainly caught from wild stocks. We address the question what will be the status of these wild stocks in the midterm future, in the year 2048, to be specific. Whereas the effects of climate change and ecological driving forces on fish stocks have already gained much attention, our prime interest is in studying the effects of changing economic drivers, as well as the impact of variable management effectiveness. Using a process‐based ecological–economic multispecies optimization model, we assess the future stock status under different scenarios of change. We simulate (i) technological progress in fishing, (ii) increasing demand for fish, and (iii) increasing supply of farmed fish, as well as the interplay of these driving forces under different scenarios of (limited) fishery management effectiveness. We find that economic change has a substantial effect on fish populations. Increasing aquaculture production can dampen the fishing pressure on wild stocks, but this effect is likely to be overwhelmed by increasing demand and technological progress, both increasing fishing pressure. The only solution to avoid collapse of the majority of stocks is institutional change to improve management effectiveness significantly above the current state. We conclude that full recognition of economic drivers of change will be needed to successfully develop an integrated ecosystem management and to sustain the wild fish stocks until 2048 and beyond.     

Wasp-waist interactions in the North Sea ecosystem

Background

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Methodology/Principal Findings

We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill.

Conclusion/Significance

Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that “wasp-waist” interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.     

Ecological forecasting under climate change: the case of Baltic cod

Good decision making for fisheries and marine ecosystems requires a capacity to anticipate the consequences of management under different scenarios of climate change. The necessary ecological forecasting calls for ecosystem-based models capable of integrating multiple drivers across trophic levels and properly including uncertainty. The methodology presented here assesses the combined impacts of climate and fishing on marine food-web dynamics and provides estimates of the confidence envelope of the forecasts. It is applied to cod (Gadus morhua) in the Baltic Sea, which is vulnerable to climate-related decline in salinity owing to both direct and indirect effects (i.e. through species interactions) on early-life survival. A stochastic food web-model driven by regional climate scenarios is used to produce quantitative forecasts of cod dynamics in the twenty-first century. The forecasts show how exploitation would have to be adjusted in order to achieve sustainable management under different climate scenarios.     

A test of three alternative pathways for consumer regulation of primary productivity

The pathways linking consumer effects to primary productivity (PPR) are likely to vary among taxa because of species-specific trophic and functional differences. Thus, it is necessary to understand the dynamics of consumer–PPR interactions so that effects of species loss on ecosystem function can be addressed from a mechanistic approach. In this study, I used three fish taxa (orangethroat darter, Etheostoma spectabile; western mosquitofish, Gambusia affinis; and bullhead minnow, Pimephales vigilax) as model consumers with different trophic and functional characteristics to test alternative mechanisms for consumer regulation of PPR (i.e., trophic cascade, terrestrial nutrient translocation, and sedimentary nutrient translocation). Experiments were conducted in stream mesocosms fitted with a combination of fish and terrestrial insect barriers to address relative importance of consumer-driven top-down and bottom-up control of PPR. A predatory invertivore, orangethroat darter, increased PPR through an apparent trophic cascade by localized reduction of benthic grazing invertebrate densities (i.e., top-down). A surface feeding insectivore, western mosquitofish, consumed terrestrial insects on the stream surface, increasing PPR by enhancing allochthonous nutrients in the mesocosms (i.e., bottom-up). A benthic omnivore, bullhead minnow, consumed benthic food items, resulting in increased PPR by enhancing availability of autochthonous nutrients via translocation of sedimentary nutrients (i.e., bottom-up). However, under specific environmental contexts, this species also consumed terrestrial invertebrates, potentially affecting PPR through terrestrial nutrient translocation as well. In this study, the trophic and functional characteristics of different species resulted in alternative pathways that increased PPR, suggesting that in natural ecosystems multiple consumer-driven pathways may be influencing PPR simultaneously and could potentially be important for temporal persistence of ecosystem function in changing environments.     

Metrics of predation: perils of predator-prey ratios

We developed an original modeling approach using program Stella® to investigate the usefulness of predator–prey ratios (PPRs) for interpreting top-down and bottom-up forcing on moose Alces alces. We included density-dependent feedbacks for the moose population, allowed K to vary based on amount and quality of available forage for moose, integrated effects of compensatory mortality, and added time lags in wolves Canis lupus tracking the moose population. Modeling scenarios we developed included bottom-up and top-down regulation as predetermined outcomes. We then evaluated whether PPRs would reflect the various combinations of trajectories of predator and prey populations under top-down versus bottom-up regulation. The resulting patterns of PPRs were impossible to disentangle from one another, and did not provide reliable insights into whether top-down or bottom-forcing occurred, especially over short time spans where critical decisions related to management of moose and wolves might be necessary. Only under top-down regulation did PPRs reflect the degree of predation experienced by moose, but in that instance, knowledge of top-down regulation must be known a priori to correctly interpret PPRs. Potential problems with interpreting PPRs include their double-variable nature, which resulted in the failure to reflect patterns of increase and decrease for predators and prey. We suggest that confidence intervals for PPRs be calculated from a binomial, similar to that proposed for sex and age ratios, which should help discourage the inappropriate use of this metric. We caution that the temptation to use PPRs often is irresistible, but their reliability is highly questionable. We provide an alternative method to using PPRs or other predation metrics for determining whether top-down or bottom-up forcing is occurring by adopting an approach based on the physical condition and life-history characteristics of prey.     

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).     

Is a healthy ecosystem one that is rich in parasites?

Historically, the role of parasites in ecosystem functioning has been considered trivial because a cursory examination reveals that their relative biomass is low compared with that of other trophic groups. However there is increasing evidence that parasite-mediated effects could be significant: they shape host population dynamics, alter interspecific competition, influence energy flow and appear to be important drivers of biodiversity. Indeed they influence a range of ecosystem functions and have a major effect on the structure of some food webs. Here, we consider the bottom-up and top-down processes of how parasitism influences ecosystem functioning and show that there is evidence that parasites are important for biodiversity and production; thus, we consider a healthy system to be one that is rich in parasite species.     

Bottom-Up Effects on Biomass Versus Top-Down Effects on Identity: A Multiple-Lake Fish Community Manipulation Experiment

The extent to which ecosystems are regulated by top-down relative to bottom-up control has been a dominant paradigm in ecology for many decades. For lakes, it has been shown that predation by fish is an important determinant of variation in zooplankton and phytoplankton community characteristics. Effects of fish are expected to not only be a function of total fish biomass, but also of functional composition of the fish community. Previous research on the importance of trophic cascades in lakes has largely focused on the role of zooplanktivorous and piscivorous fish. We conducted a large-scale multiple-lake fish community manipulation experiment to test for the effect of differences in fish functional community composition on the trophic structure of lakes. We examine the effect of top-down and bottom-up factors on phytoplankton and zooplankton biomass as well as on their community composition. We put our data in a broader perspective by comparing our results to data of a survey that also included ponds with low fish densities as well as ponds with very high densities of fish. Our results indicate that the overall food web structure under relative high fish densities is primarily structured by bottom-up factors, whereas community characteristics seem to be primarily regulated by top-down factors. Our results suggest a subtle interplay between bottom-up and top-down factors, in which bottom-up factors dominate in determining quantities while top-down effects are important in determining identities of the communities.     

When does fishing lead to more fish? Community consequences of bottom trawl fisheries in demersal food webs

Bottom trawls are a globally used fishing gear that physically disturb the seabed and kill non-target organisms, including those that are food for the targeted fish species. There are indications that ensuing changes to the benthic invertebrate community may increase the availability of food and promote growth and even fisheries yield of target fish species. If and how this occurs is the subject of ongoing debate, with evidence both in favour and against. We model the effects of trawling on a simple ecosystem of benthivorous fish and two food populations (benthos), susceptible and resistant to trawling. We show that the ecosystem response to trawling depends on whether the abundance of benthos is top-down or bottom-up controlled. Fishing may result in higher fish abundance, higher (maximum sustainable) yield and increased persistence of fish when the benthos which is the best-quality fish food is also more resistant to trawling. These positive effects occur in bottom-up controlled systems and systems with limited impact of fish feeding on benthos, resembling bottom-up control. Fishing leads to lower yields and fish persistence in all configurations where susceptible benthos are more profitable prey. Our results highlight the importance of mechanistic ecosystem knowledge as a requirement for successful management.     

Using an Ecosystem Modeling Approach to Explore Possible Ecosystem Impacts of Fishing in the Beibu Gulf,Northern South China Sea

Using the Ecopath with Ecosim software, a trophic structure model of the Beibu Gulf was constructed to explore the energy flows and provide a snapshot of the ecosystem operations. Input data were mainly from the trawl survey data collected from October 1998 to September 1999 and related literatures. The impacts of various fishing pressure on the biomass were examined by simulation at different fishing mortality rates. The model consists of 20 functional groups (boxes), each representing organisms with a similar role in the food web, and only covers the major trophic flows in the Beibu Gulf ecosystem. It was found that the food web of the Beibu Gulf was dominated by the primary producers path, and phytoplankton was the primary producer mostly used as a food source. The fractional trophic levels ranged from 1.0 to 4.02, and the marine mammals occupied the highest trophic level. Using network analysis, the ecosystem network was mapped into a linear food chain, and six discrete trophic levels were found with a mean transfer efficiency of 11.2%. The Finn cycling index was 9.73%. The path length was 1.821. The omnivory index was 0.197. The ecosystem had some degree of instability due to exploitation and other human activities, according to Odum’s theory of ecosystem development. A 10-year simulation was performed for each fishery scenario. The fishing mortality rate was found to have a strong impact on the biomass. By keeping the fishing mortality rate at the current level for all fishing sectors, scenario 1 had a drastic decrease in the large fish groups. The biomass of the small and medium pelagic fish would increase to some extent. The biomass of the small and low trophic level species, jellyfish, prawns and benthic crustaceans would be stable. The total biomass of the fishery resources would have a 10% decrease from the current biomass after 10 years. In contrast, the reduced fishing mortality rate induced the recovery of biomass (scenarios 2–4). In scenario 2, the biomass of the large demersal fish and the large pelagic fish would increase to over 16 times and 10 times, respectively, of their current level. In scenario 4, the biomass of the large pelagic fish would increase to over 3 times of its current level. The total biomass of the fish groups, especially the high trophic level groups, would become significantly higher after 10 years, which illustrates the contribution on biomass recovery by relaxing the fishing pressure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author contributions: Xiaoping Jia designed research; Zuozhi Chen and Yongsong Qiu performed research; Zuozhi Chen, Yongsong Qiu, and Shannan Xu analyzed data; and Zuozhi Chen and Shannan Xu wrote the article.     

An Integrated Coral Reef Ecosystem Model to Support Resource Management under a Changing Climate

Millions of people rely on the ecosystem services provided by coral reefs, but sustaining these benefits requires an understanding of how reefs and their biotic communities are affected by local human-induced disturbances and global climate change. Ecosystem-based management that explicitly considers the indirect and cumulative effects of multiple disturbances has been recommended and adopted in policies in many places around the globe. Ecosystem models give insight into complex reef dynamics and their responses to multiple disturbances and are useful tools to support planning and implementation of ecosystem-based management. We adapted the Atlantis Ecosystem Model to incorporate key dynamics for a coral reef ecosystem around Guam in the tropical western Pacific. We used this model to quantify the effects of predicted climate and ocean changes and current levels of current land-based sources of pollution (LBSP) and fishing. We used the following six ecosystem metrics as indicators of ecosystem state, resilience and harvest potential: 1) ratio of calcifying to non-calcifying benthic groups, 2) trophic level of the community, 3) biomass of apex predators, 4) biomass of herbivorous fishes, 5) total biomass of living groups and 6) the end-to-start ratio of exploited fish groups. Simulation tests of the effects of each of the three drivers separately suggest that by mid-century climate change will have the largest overall effect on this suite of ecosystem metrics due to substantial negative effects on coral cover. The effects of fishing were also important, negatively influencing five out of the six metrics. Moreover, LBSP exacerbates this effect for all metrics but not quite as badly as would be expected under additive assumptions, although the magnitude of the effects of LBSP are sensitive to uncertainty associated with primary productivity. Over longer time spans (i.e., 65 year simulations), climate change impacts have a slight positive interaction with other drivers, generally meaning that declines in ecosystem metrics are not as steep as the sum of individual effects of the drivers. These analyses offer one way to quantify impacts and interactions of particular stressors in an ecosystem context and so provide guidance to managers. For example, the model showed that improving water quality, rather than prohibiting fishing, extended the timescales over which corals can maintain high abundance by at least 5–8 years. This result, in turn, provides more scope for corals to adapt or for resilient species to become established and for local and global management efforts to reduce or reverse stressors.     

Incorporating anthropogenic effects into trophic ecology: predator–prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species'' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth''s terrestrial surface.     

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.     

互利关系和植物多样性对节肢动物群落不同营养级的影响路径和强度

食物网中的上行效应和下行效应对于群落的动态和生态系统功能有十分重要的影响,旨在探讨互利关系和植物多样性对节肢动物群落中食物网不同营养级之间的影响。通过随机裂区试验方法,分别设置了3种蚂蚁-紫胶虫互利关系处理(有互利关系、无互利关系和自然对照)以及3种植物多样性处理(单一种植、2树种混植和3树种混植),于2016年8月和9月分两次用手捡法、网扫法和震落法采集试验地寄主植物上所有的节肢动物,并按照不同营养级将其分类。利用结构方程模型分析方法对不同营养级之间的相互作用的路径和强度进行了比较,结果显示:1)互利关系对捕食者和消费者均有显著的下行作用,有互利关系处理下蚂蚁对捕食者的路径强度要强于自然对照组,互利关系对捕食者的影响要强于对消费者的影响。2)植物多样性会通过影响植物的生物量而对消费者和捕食者产生显著的上行效应影响,这种影响会随着营养级的升高而显著减小。3)消费者主要受植物多样性的上行效应影响,而捕食者主要受互利关系的下行效应影响。有互利关系的食物网结构更加复杂,营养级之间的相互作用更为显著。探讨了以蚂蚁-紫胶虫互利关系为核心作用的紫胶林生态系统中互利关系和植物多样性对节肢动物食物网中各营养级的影响,揭示了上行效应和下行效应对各营养级的作用途径和强度,其结果有一定的理论参考价值。     

Exploitation and habitat degradation as agents of change within coral reef fish communities

Over‐exploitation and habitat degradation are the two major drivers of global environmental change and are responsible for local extinctions and declining ecosystem services. Here we compare the top‐down effect of exploitation by fishing with the bottom‐up influence of habitat loss on fish communities in the most diverse of ecological systems, coral reefs. Using a combination of multivariate techniques and path analyses, we illustrate that the relative importance of coral cover and fishing in controlling fish abundance on remote Fijian reefs varies between species and functional groups. A decline in branching Acropora coral is strongly associated with a decline in abundance of coral‐feeding species, and a decrease in coral‐associated habitat complexity, which has indirectly contributed to reduced abundance of small‐bodied damselfish. In contrast, reduced fishing pressure, brought about by declining human populations and a shift to alternate livelihoods, is associated with increased abundance of some piscivores and fisheries target species. However, availability of prey is controlled by coral‐associated habitat complexity and appears to be a more important driver of total piscivore abundance compared with fishing pressure. Effects of both fishing and coral loss are stronger on individual species than functional groups, as variation in the relative importance of fishing or coral loss among species within the same functional group attenuated the impact of either of these potential drivers at the functional level. Overall, fishing continues to have an influence on Fijian fish communities; however, habitat loss is currently the overriding agent of change. The importance of coral loss mediated by climate change is expected to have an increasing contribution to fish community dynamics, particularly in remote locations or where the influence of fishing is waning.