Modeling the trophic effects of marine protected area zoning policies: A case study

Marine protected areas (MPAs) are increasingly being recognized as an alternative management tool for conserving marine resources and ecosystems. By integrating organism dispersal rates, ecosystem interactions and fishing effort dynamics, ECOSPACE, a spatially explicit ecosystem-based modeling tool, allowed us to compare the ecological consequences of alternative MPA zoning policies within the proposed Gwaii Haanas National Marine Conservation Area, located off the west coast of British Columbia, Canada. The desired effects of MPAs include higher fishery yields, the conservation of biodiversity, and/or the preservation of intact ecosystems. However, ECOSPACE predicts that when MPAs are small, species interactions and movements may make these objectives difficult to achieve. ECOSPACE suggests that the effects of MPAs are reduced at their boundaries where fishing effort is predicted to concentrate. Furthermore, top predators may become more abundant within MPAs, which could lead to a depression of their prey species and a subsequent increase of species at even lower trophic levels. Trophic cascade patterns and density gradients across boundaries are nontrivial departures from our simple expectations of how MPAs protect areas and will force us to reconsider what constitutes effective conservation. Our ECOSPACE model indicates that the establishment of multi-use buffer zones may help alleviate these realistic but worrisome ecological predictions. When coupled with an overall reduction in harvest pressure, ECOSPACE suggests that a MPA with a large core `no-take' zone and large buffer will result in the greatest increase in organism biomass. The use of marine zoning may be an effective management tactic to reduce social conflict and conserve marine ecosystems.     

Predator-induced demographic shifts in coral reef fish assemblages

In recent years, it has become apparent that human impacts have altered community structure in coastal and marine ecosystems worldwide. Of these, fishing is one of the most pervasive, and a growing body of work suggests that fishing can have strong effects on the ecology of target species, especially top predators. However, the effects of removing top predators on lower trophic groups of prey fishes are less clear, particularly in highly diverse and trophically complex coral reef ecosystems. We examined patterns of abundance, size structure, and age-based demography through surveys and collection-based studies of five fish species from a variety of trophic levels at Kiritimati and Palmyra, two nearby atolls in the Northern Line Islands. These islands have similar biogeography and oceanography, and yet Kiritimati has ～10,000 people with extensive local fishing while Palmyra is a US National Wildlife Refuge with no permanent human population, no fishing, and an intact predator fauna. Surveys indicated that top predators were relatively larger and more abundant at unfished Palmyra, while prey functional groups were relatively smaller but showed no clear trends in abundance as would be expected from classic trophic cascades. Through detailed analyses of focal species, we found that size and longevity of a top predator were lower at fished Kiritimati than at unfished Palmyra. Demographic patterns also shifted dramatically for 4 of 5 fish species in lower trophic groups, opposite in direction to the top predator, including decreases in average size and longevity at Palmyra relative to Kiritimati. Overall, these results suggest that fishing may alter community structure in complex and non-intuitive ways, and that indirect demographic effects should be considered more broadly in ecosystem-based management.     

New trophic indicators and target values for an ecosystem-based management of fisheries

In the present study, we tested five trophic indicators and we demonstrated their usefulness to assess the environmental status of marine ecosystems and to implement an ecosystem approach to fisheries management (EAFM). The tested indicators include the slope of the biomass spectrum, the mean trophic level (MTL), the marine trophic index (MTI) and two newly developed indicators, the high trophic level indicator (HTI) and the apex predator indicator (API). Indicators are compared between current state and potential reference situations, using as case studies: the Celtic Sea/Bay of Biscay, North Sea and English Channel ecosystems. Trophic spectra are obtained from Ecopath models while reference situations are estimated, simulating with EcoTroph and Ecosim different fishing pressures including three candidate scenarios for an EAFM. Inter-ecosystems assessments are done using Ecopath models, simulations outputs and scientific surveys data to assess the current states of the studied ecosystems, contrast the reference situations and analyze the responses of all indicators. Sensitivity analyses are also conducted on the main simulation parameters to test the robustness of the chosen indicators. Ecosystems specific targets for EAFM are proposed for the five trophic indicators estimated from whole-ecosystem models, while in the Celtic Sea/Bay of Biscay ecosystem targets are proposed for the MTL (=3.85) and HTI (48%) estimated from standard bottom-trawl surveys. The HTI is proposed to be relevant for survey data and the API is recommended using whole-ecosystem models. We conclude that HTI and API show trends in ecosystems health better than MTI.     

Global Patterns in Ecological Indicators of Marine Food Webs: A Modelling Approach

Background

Ecological attributes estimated from food web models have the potential to be indicators of good environmental status given their capabilities to describe redundancy, food web changes, and sensitivity to fishing. They can be used as a baseline to show how they might be modified in the future with human impacts such as climate change, acidification, eutrophication, or overfishing.

Methodology

In this study ecological network analysis indicators of 105 marine food web models were tested for variation with traits such as ecosystem type, latitude, ocean basin, depth, size, time period, and exploitation state, whilst also considering structural properties of the models such as number of linkages, number of living functional groups or total number of functional groups as covariate factors.

Principal findings

Eight indicators were robust to model construction: relative ascendency; relative overhead; redundancy; total systems throughput (TST); primary production/TST; consumption/TST; export/TST; and total biomass of the community. Large-scale differences were seen in the ecosystems of the Atlantic and Pacific Oceans, with the Western Atlantic being more complex with an increased ability to mitigate impacts, while the Eastern Atlantic showed lower internal complexity. In addition, the Eastern Pacific was less organised than the Eastern Atlantic although both of these systems had increased primary production as eastern boundary current systems. Differences by ecosystem type highlighted coral reefs as having the largest energy flow and total biomass per unit of surface, while lagoons, estuaries, and bays had lower transfer efficiencies and higher recycling. These differences prevailed over time, although some traits changed with fishing intensity. Keystone groups were mainly higher trophic level species with mostly top-down effects, while structural/dominant groups were mainly lower trophic level groups (benthic primary producers such as seagrass and macroalgae, and invertebrates). Keystone groups were prevalent in estuarine or small/shallow systems, and in systems with reduced fishing pressure. Changes to the abundance of key functional groups might have significant implications for the functioning of ecosystems and should be avoided through management.

Conclusion/significance

Our results provide additional understanding of patterns of structural and functional indicators in different ecosystems. Ecosystem traits such as type, size, depth, and location need to be accounted for when setting reference levels as these affect absolute values of ecological indicators. Therefore, establishing absolute reference values for ecosystem indicators may not be suitable to the ecosystem-based, precautionary approach. Reference levels for ecosystem indicators should be developed for individual ecosystems or ecosystems with the same typologies (similar location, ecosystem type, etc.) and not benchmarked against all other ecosystems.     

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.     

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.     

Structure and functioning of two pelagic communities in the North Chilean Patagonian coastal system

The size composition of primary producers is important for how energy is channeled through a food web and on to the higher trophic levels and eventually to fisheries. To evaluate this, we studied the productive patterns for large (micro) versus small (nano) phytoplankton in two south marine Patagonian ecosystems: The Inner Sea of Chiloe—ISCh and, Moraleda Channel—MCh. We built Ecopath models (EwE), and evaluated the hypothesis that the overall primary productivity—rather than the ratio of large to small primary producers—constitutes an adequate proxy for predicting the amount of secondary and tertiary production and biomass (up to the fisheries). The EwE model included four small-scale fisheries and 36 functional groups. The functioning of both ecosystems was similar but the ecosystem parameters (biomass, energy transfer efficiencies from primary producers, secondary, and tertiary production) were twice as much in the basin with more microphytoplankton biomass. Overall, the hypothesis was rejected, albeit it was possible to highlight the importance of the quality and size spectrum of plankton on the structure of marine ecosystem, and to demonstrate the key role of the microbial loop over traditional food web in the functioning of the carbon biological pump in Patagonia ecosystems.     

Land use change affects macroinvertebrate community size spectrum in streams: the case of Pinus radiata plantations

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Synergistic Effects of Climate and Fishing in a Marine Ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere. Author Contributions  RRK conceived the project and GB analysed the data. RRK, GB and JAL co-wrote the paper.     

Trophic levels of fish species of commercial importance in the Colombian Caribbean

Ecological studies on commercial important fish species are of great value to support resource management issues. This study calculated trophic levels of those Colombian Caribbean fish species whose diet has been locally described. Usable diet data of 119 species resulted in 164 trophic level estimates. An ordinary regression model relating trophic level and fish size was formulated. The regression slope was positive and significantly different from zero (p < 0.05) suggesting a scaling of trophic level with fish size. Both the list of trophic levels and the regression model should be of help in the formulation of trophic indicators and models of neotropical ecosystems.     

Ecological indicators to capture the effects of fishing on biodiversity and conservation status of marine ecosystems

IndiSeas (“Indicators for the Seas”) is a collaborative international working group that was established in 2005 to evaluate the status of exploited marine ecosystems using a suite of indicators in a comparative framework. An initial shortlist of seven ecological indicators was selected to quantify the effects of fishing on the broader ecosystem using several criteria (i.e., ecological meaning, sensitivity to fishing, data availability, management objectives and public awareness). The suite comprised: (i) the inverse coefficient of variation of total biomass of surveyed species, (ii) mean fish length in the surveyed community, (iii) mean maximum life span of surveyed fish species, (iv) proportion of predatory fish in the surveyed community, (v) proportion of under and moderately exploited stocks, (vi) total biomass of surveyed species, and (vii) mean trophic level of the landed catch. In line with the Nagoya Strategic Plan of the Convention on Biological Diversity (2011–2020), we extended this suite to emphasize the broader biodiversity and conservation risks in exploited marine ecosystems. We selected a subset of indicators from a list of empirically based candidate biodiversity indicators initially established based on ecological significance to complement the original IndiSeas indicators. The additional selected indicators were: (viii) mean intrinsic vulnerability index of the fish landed catch, (ix) proportion of non-declining exploited species in the surveyed community, (x) catch-based marine trophic index, and (xi) mean trophic level of the surveyed community. Despite the lack of data in some ecosystems, we also selected (xii) mean trophic level of the modelled community, and (xiii) proportion of discards in the fishery as extra indicators. These additional indicators were examined, along with the initial set of IndiSeas ecological indicators, to evaluate whether adding new biodiversity indicators provided useful additional information to refine our understanding of the status evaluation of 29 exploited marine ecosystems. We used state and trend analyses, and we performed correlation, redundancy and multivariate tests. Existing developments in ecosystem-based fisheries management have largely focused on exploited species. Our study, using mostly fisheries independent survey-based indicators, highlights that biodiversity and conservation-based indicators are complementary to ecological indicators of fishing pressure. Thus, they should be used to provide additional information to evaluate the overall impact of fishing on exploited marine ecosystems.     

The Role of Pre-Existing Disturbances in the Effect of Marine Reserves on Coastal Ecosystems: A Modelling Approach

We have used an end-to-end ecosystem model to explore responses over 30 years to coastal no-take reserves covering up to 6% of the fifty thousand square kilometres of continental shelf and slope off the coast of New South Wales (Australia). The model is based on the Atlantis framework, which includes a deterministic, spatially resolved three-dimensional biophysical model that tracks nutrient flows through key biological groups, as well as extraction by a range of fisheries. The model results support previous empirical studies in finding clear benefits of reserves to top predators such as sharks and rays throughout the region, while also showing how many of their major prey groups (including commercial species) experienced significant declines. It was found that the net impact of marine reserves was dependent on the pre-existing levels of disturbance (i.e. fishing pressure), and to a lesser extent on the size of the marine reserves. The high fishing scenario resulted in a strongly perturbed system, where the introduction of marine reserves had clear and mostly direct effects on biomass and functional biodiversity. However, under the lower fishing pressure scenario, the introduction of marine reserves caused both direct positive effects, mainly on shark groups, and indirect negative effects through trophic cascades. Our study illustrates the need to carefully align the design and implementation of marine reserves with policy and management objectives. Trade-offs may exist not only between fisheries and conservation objectives, but also among conservation objectives.     

Factors controlling planktonic size spectral responses to autumnal circulation in a Mediterranean lake

1. Studies of planktonic size spectra have been common in recent years, but few concerning the effects of autumnal lake processes on those distributions have been reported. We carried out such a study for 93 days during early circulation in a small, mesotrophic, seepage lake with only benthivorous fish. Two distinct mixing periods occurred before full circulation in Las Madres lake (Spain). As a proxy of overall planktonic‐ and phytoplanktonic size distributions, the shape of a Pareto I power function was traced over time. The phytoplanktonic size spectrum was also used to test the hypothesis that phytoplankton might show a trophic cascade during early circulation. 2. Our results demonstrated that the overall size spectrum changed smoothly during the autumnal mixing, with sequential changes controlled by a combination of biotic and abiotic variables, albeit experiencing different temporal delays, always shorter than a week. A first combined, autogenic process, driven both by algal competition and a crustacean trophic effect, and a second physical‐forcing process, driven by the combined effect of convective cooling and decreasing irradiance on mixed layer dynamics, may be related to the overall planktonic spectrum in the first and second periods of mixing, respectively. 3. The phytoplanktonic size spectrum did not appear to be dictated by a trophic cascade in Las Madres lake, their dynamics being mostly controlled by physical forcing, along with some effect of non‐edible primary producers and cladocerans in the first and second periods of mixing, respectively. 4. Our results and others covering early circulation with longer sampling periods suggest that planktonic size spectral dynamics during lake circulation is context‐dependent (i.e. varying from one lake to another), thus preventing generalisation. However, when further studies with finer temporal resolution have been carried out, it is likely that clear‐cut patterns in the planktonic size spectrum will emerge, arising from the interplay of autogenic plankton dynamics, implying some resistance to community change because of external physical forcing, and the velocity of autumnal mixing.     

Using successional theory to measure marine ecosystem health

Marine ecosystems are diverse and complex, providing significant challenges to the development of generalizable metrics of ecosystem health. Of particular concern is the varied form of change caused by multiple human activities, which limits the capacity to generate a single measure to encapsulate the overall condition of the ecosystem. Here we consider how successional theory can help to simplify our understanding of marine community structure, especially when viewed in context of human disturbance. During succession, the emergent properties of communities change in predictable ways. As communities mature, there is an increase in total production and biomass, the mean size of organisms, the level of internal recycling of food and nutrients, and the mean trophic level. Using a set of multi-species trophic models, we explore the changes in community structure that are likely to occur during succession. These changes include increases in biomass within trophic levels due to decreased rates of energy and food loss through trophic and production inefficiencies, and potential shifts from top-down control early in succession to bottom-up later. Because human activities disproportionately favor early-successional species, we can gain insights by considering community degradation in the context of succession being played in reverse. Indicators of health based on ecological succession thus provide a mechanistic view to measure the impact of human activities (both positive and negative) on marine ecosystems.     

Combined Fishing and Climate Forcing in the Southern Benguela Upwelling Ecosystem: An End-to-End Modelling Approach Reveals Dampened Effects

The effects of climate and fishing on marine ecosystems have usually been studied separately, but their interactions make ecosystem dynamics difficult to understand and predict. Of particular interest to management, the potential synergism or antagonism between fishing pressure and climate forcing is analysed in this paper, using an end-to-end ecosystem model of the southern Benguela ecosystem, built from coupling hydrodynamic, biogeochemical and multispecies fish models (ROMS-N₂P₂Z₂D₂-OSMOSE). Scenarios of different intensities of upwelling-favourable wind stress combined with scenarios of fishing top-predator fish were tested. Analyses of isolated drivers show that the bottom-up effect of the climate forcing propagates up the food chain whereas the top-down effect of fishing cascades down to zooplankton in unfavourable environmental conditions but dampens before it reaches phytoplankton. When considering both climate and fishing drivers together, it appears that top-down control dominates the link between top-predator fish and forage fish, whereas interactions between the lower trophic levels are dominated by bottom-up control. The forage fish functional group appears to be a central component of this ecosystem, being the meeting point of two opposite trophic controls. The set of combined scenarios shows that fishing pressure and upwelling-favourable wind stress have mostly dampened effects on fish populations, compared to predictions from the separate effects of the stressors. Dampened effects result in biomass accumulation at the top predator fish level but a depletion of biomass at the forage fish level. This should draw our attention to the evolution of this functional group, which appears as both structurally important in the trophic functioning of the ecosystem, and very sensitive to climate and fishing pressures. In particular, diagnoses considering fishing pressure only might be more optimistic than those that consider combined effects of fishing and environmental variability.     

Structuring dynamic models of exploited ecosystems from trophic mass-balance assessments

The linear equations that describe trophic fluxes in mass-balance, equilibrium assessments of ecosystems (such as in the ECOPATH approach) can be re-expressed as differential equations defining trophic interactions as dynamic relationships varying with biomasses and harvest regimes. Time patterns of biomass predicted by these differential equations, and equilibrium system responses under different exploitation regimes, are found by setting the differential equations equal to zero and solving for biomasses at different levels of fishing mortality. Incorporation of our approach as the ECOSIM routine into the well-documented ECOPATH software will enable a wide range of potential users to conduct fisheries policy analyses that explicitly account for ecosystem trophic interactions, without requiring the users to engage in complex modelling or information gathering much beyond that required for ECOPATH. While the ECOSIM predictions can be expected to fail under fishing regimes very different from those leading to the ECOPATH input data, ECOSIM will at least indicate likely directions of biomass change in various trophic groups under incremental experimental policies aimed at improving overall ecosystem management. That is, ECOSIM can be a valuable tool for design of ecosystem-scale adaptive management experiments     

Causes and projections of abrupt climate-driven ecosystem shifts in the North Atlantic

Warming of the global climate is now unequivocal and its impact on Earth' functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.     

Trophic levels of teleost and elasmobranch species in the Persian Gulf and Oman Sea

Knowing the trophic level of marine organisms is essential to understanding their ecological role in the ecosystem and for quantifying the ecosystem effects of fishing to establish effective management of fishing resources. In comparison to other systems, information about the trophic level of marine organisms in the Persian Gulf and Oman Sea is very scarce. Here, the main aim was to estimate trophic level in these areas using all available diet information from different marine species using TrophLab software. The trophic level of 32 fish species was estimated with the available diet data. The trophic level ranged from 2.28 to 4.50. High trophic levels were found for Chorocentrus nudus (TL = 4.7), Saurida tumbil (TL = 4.6), Rhizoprionodon acutus (TL = 4.5), Torpedo sinuspersici (TL=4.5), Gymnura poecilura (TL = 4.5), Sphyraena putnamae (TL = 4.5) and Euthynnus affinis (TL = 4.5). In contrast, lower trophic levels were estimated for Tenualosa ilisha (TL = 2.28) and Sardinella sindensis (TL = 2.92). As expected, a positive correlation was found between the trophic level and body size, indicating changes in the diet due to variations in predatory capacities. The results of this study may be useful in the formulation of trophic indicators and modelling of the ecosystems.     

Multi-level trophic cascades in a heavily exploited open marine ecosystem

Anthropogenic disturbances intertwined with climatic changes can have a large impact on the upper trophic levels of marine ecosystems, which may cascade down the food web. So far it has been difficult to demonstrate multi-level trophic cascades in pelagic marine environments. Using field data collected during a 33-year period, we show for the first time a four-level community-wide trophic cascade in the open Baltic Sea. The dramatic reduction of the cod (Gadus morhua) population directly affected its main prey, the zooplanktivorous sprat (Sprattus sprattus), and indirectly the summer biomass of zooplankton and phytoplankton (top-down processes). Bottom-up processes and climate-hydrological forces had a weaker influence on sprat and zooplankton, whereas phytoplankton variation was explained solely by top-down mechanisms. Our results suggest that in order to dampen the occasionally harmful algal blooms of the Baltic, effort should be addressed not only to control anthropogenic nutrient inputs but also to preserve structure and functioning of higher trophic levels.     

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).