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.     

Impact of Non-Native Terrestrial Mammals on the Structure of the Terrestrial Mammal Food Web of Newfoundland,Canada

The island of Newfoundland is unique because it has as many non-native terrestrial mammals as native ones. The impacts of non-native species on native flora and fauna can be profound and invasive species have been identified as one of the primary drivers of species extinction. Few studies, however, have investigated the effects of a non-native species assemblage on community and ecosystem properties. We reviewed the literature to build the first terrestrial mammal food web for the island of Newfoundland and then used network analyses to investigate how the timing of introductions and trophic position of non-native species has affected the structure of the terrestrial mammal food web in Newfoundland. The first non-native mammals (house mouse and brown rat) became established in Newfoundland with human settlement in the late 15^th and early 16^th centuries. Coyotes and southern red-backed voles are the most recent mammals to establish themselves on the island in 1985 and 1998, respectively. The fraction of intermediate species increased with the addition of non-native mammals over time whereas the fraction of basal and top species declined over time. This increase in intermediate species mediated by non-native species arrivals led to an overall increase in the terrestrial mammal food web connectance and generality (i.e. mean number of prey per predator). This diverse prey base and sources of carrion may have facilitated the natural establishment of coyotes on the island. Also, there is some evidence that the introduction of non-native prey species such as the southern red-backed vole has contributed to the recovery of the threatened American marten. Long-term monitoring of the food web is required to understand and predict the impacts of the diverse novel interactions that are developing in the terrestrial mammal food web of Newfoundland.     

地上枯落物的累积、分解及其在陆地生态系统中的作用

了解陆地生态系统地上枯落物的累积和分解过程对认识它的生态作用、通过管理地上枯落物调控陆地生态系统功能和服务有重要意义。综述了陆地生态系统地上枯落物的积累和分解过程及其影响因素,然后概括了通过这些过程地上枯落物所发挥的生态作用,最后,在全球变化背景下,基于当前研究进展提出陆地生态系统地上枯落物研究的前景。地上枯落物累积在时间尺度上一般遵循植物的生命周期,同时也受环境因子的调控。大的空间尺度上,枯落物累积主要受水热因子控制,伴随植被类型的变化,表现随纬度升高而减少的趋势。然而,在局域尺度内,枯落物累积除受水、热因子限制,还被群落结构、土壤条件、植食动物等因素影响,表现较大变异性。当前,人类干扰作为一个不可忽视的因素,正在强烈甚至不可逆转的改变地表植被覆盖和枯落物累积。地上枯落物的分解过程包括淋溶、光降解、土壤动物和微生物分解,这些过程同时进行并相互影响。尽管目前还不清楚,但区分这些分解过程和分解产物的去向对了解陆地生态系统物质循环有重要意义。枯落物分解首先被自身类型、化学组成、物种多样性决定,同时也受分解者群体、非生物环境影响。其中,枯落物分解与其化学特性、物种多样性及土壤养分状况的关系是研究的热点,也是广泛争议的焦点。通过累积和分解,地上枯落物对陆地生态系统有物理、化学、生物作用。目前,枯落物的物理和化学作用研究较为透彻,而由于受枯落物数量、环境条件、响应植物特征或一些有待挖掘的未知因素的共同限制,地上枯落物的生物作用,尤其对植物的作用在不同研究中仍没有达成普遍的共识。全球变化可能影响地上枯落物累积、分解和生态作用。在全球变化的背景,研究地上枯落物产量和性状变化、阐明枯落物分解的分室模型、继续分析枯落物性状和分解关系、深入揭示枯落物的生态作用及其制约因素,理解和预测地上枯落物数量和质量变化对陆地生态系统功能和服务的影响是必要的。     

Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems

Carrion use by terrestrial vertebrates is much more prevalent than conventional theory implies, and, rather than a curiosity of animal behavior, is a key ecological process that must be accounted for. Human aversion to rotted substances and difficulties associated with identifying scavenged material in studies of food habits have contributed to the relative lack of information concerning scavenging behavior in vertebrates. Several lines of evidence, however, suggest that carrion resources are more extensively used by vertebrates than has been widely assumed: 1) a substantial number of animals die from causes other than predation and become available to scavengers, 2) a wide variety of vertebrate scavengers, rather than microbes or arthropods, consume most available carcasses, and 3) intense competition exists between vertebrate scavengers and decomposers, especially in warm climates. Although vultures are best adapted to use carrion, nearly all vertebrate predators are also scavengers to some extent. The costs and benefits associated with carrion use influences the evolution of scavenging behavior in vertebrates, resulting in a continuum of facultative scavengers that use carrion to varying degrees. The realized usage of carrion by a vertebrate species is influenced by the speed and efficiency with which it forages, its visual and olfactory abilities, and its capacity for detoxifying products of decomposition. A deeper understanding of carrion use by facultative scavengers will improve our knowledge of community and ecosystem processes, especially the flow of energy through food webs.     

Decomposition rate of carrion is dependent on composition not abundance of the assemblages of insect scavengers

Environmental factors and biodiversity affect ecosystem processes. As environmental change modifies also biodiversity it is unclear whether direct effects of environmental factors on ecosystem processes are more important than indirect effects mediated by changes in biodiversity. High-quality resources like carrion occur as heterogeneous pulses of energy and nutrients. Consequently, the distribution of scavenging insects is related to resource availability. Therefore, carrion decomposition represents a suitable process from which to unravel direct effects of environmental change from indirect biodiversity-related effects on ecosystem processes. During three field seasons in 2010 we exposed traps baited with small-mammal carrion at 21 sites along a temperature gradient to explore the insect carrion fauna and decomposition rate in the Bohemian Forest, Germany. The abundance component of beetle and fly assemblages decreased with decreasing temperature. Independently, the composition component of both taxa changed with temperature and season. The change in the composition component of beetles depicted a loss of larger species at higher temperatures. Decomposition rate did not change directly along the temperature gradient but was directly influenced by season. The composition component of beetles, and to a small extent of flies, but not their abundance component, directly affected carrion decomposition. Consequently, lower decomposition rates at lower temperatures can be explained by the absence of larger beetle species. Thus, we predict that future environmental change will modify carrion fauna composition and thereby indirectly decomposition rate. Moreover, reorganizations of the insect carrion composition will directly translate into modified decomposition rates, with potential consequences for nutrient availability and carbon storage.     

农田土壤食物网管理的原理与方法

土壤食物网在维持生态系统生产力和健康等方面起着重要作用,但现代农业中,化肥农药等外部投入已经改变或部分替代了土壤食物网的功能,由此也造成一系列的环境问题。为了协调作物高产与环境保护的利益,需要对土壤食物网进行管理,使土壤食物网符合作物生长的需要,即建立健康土壤食物网。管理土壤食物网有两种方式:(1)直接方式,即通过调节食物网各个功能群的组成来管理土壤食物网;(2)间接方式,即根据农田土壤食物网以自下而上调控方式为主、强调低营养阶层的资源限制的原理,通过调节碎屑的数量和质量来管理食物网。在这两种调控方式中,都需要对被管理的食物网进行监测,监测的方式也分两种,一种是直接测定食物网各功能群的数量和生物量,另外一种方式即以线虫为工具来反应土壤食物网的结构。     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Terrestrial food web of the Malpelo Fauna and Flora Sanctuary,Colombia: An analysis from a topological approach

Malpelo Island is the largest marine protected area (MPA) in the Colombian Pacific; however, the lack of information regarding its ecological dynamics suggests that management and conservation strategies are developed from an individual approach (species or groups of species) and not from an ecosystem one. This study analyzes the terrestrial food web of Malpelo Island, Colombia, based on topological analysis (e.g., centrality). The food web was constructed from 27 nodes representing the main trophogroups, which consist of species or functional groups. Malpelo Island had a food web of four steps with a maximum separation among all trophogroups and trophic pathways, with two steps in average length. Furthermore, the food web was divided into three food web compartments, with a recurrence of connectivity patterns dominated by apparent and exploitative competition, followed by tri-trophic chains and omnivory. Five key trophogroups control the energy flow throughout the food web (detritus, the land crab Johngarthia malpilensis, the lizard Anolis agassizi, the Malpelo dotted galliwasp Diploglossus millepunctatus, and the Nazca booby Sula granti). The high importance of detritus suggests that bottom-up processes act as a control and regulation mechanism of trophic flows. The low number of food web compartments and a high recurrence of specific connectivity patterns in the Malpelo Island terrestrial ecosystem evidence different ecological processes centered on five trophogroups, allowing stability against disturbances. In addition, the simulation of trophogroup removal (randomly or directed) suggests that food web can be vulnerable to structural alterations in their properties, which may have consequences on the resilience of this ecosystem. This study contributes to the knowledge of the trophic dynamics of Malpelo Island, providing a potential tool for management and conservation measures from an ecosystemic approach.     

Unravelling the complex structure of forest soil food webs: higher omnivory and more trophic levels

Food web topologies depict the community structure as distributions of feeding interactions across populations. Although the soil ecosystem provides important functions for aboveground ecosystems, data on complex soil food webs is notoriously scarce, most likely due to the difficulty of sampling and characterizing the system. To fill this gap we assembled the complex food webs of 48 forest soil communities. The food webs comprise 89 to 168 taxa and 729 to 3344 feeding interactions. The feeding links were established by combining several molecular methods (stable isotope, fatty acid and molecular gut content analyses) with feeding trials and literature data. First, we addressed whether soil food webs (n = 48) differ significantly from those of other ecosystem types (aquatic and terrestrial aboveground, n = 77) by comparing 22 food web parameters. We found that our soil food webs are characterized by many omnivorous and cannibalistic species, more trophic chains and intraguild‐predation motifs than other food webs and high average and maximum trophic levels. Despite this, we also found that soil food webs have a similar connectance as other ecosystems, but interestingly a higher link density and clustering coefficient. These differences in network structure to other ecosystem types may be a result of ecosystem specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food‐web level. In a second analysis of land‐use effects, we found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. Overall, our study has unravelled some systematic structures of soil food‐webs, which extends our mechanistic understanding how environmental characteristics of the soil ecosystem determine patterns at the community level.     

Inter‐specific interactions linking predation and scavenging in terrestrial vertebrate assemblages

Predation and scavenging have been classically understood as independent processes, with predator–prey interactions and scavenger–carrion relationships occurring separately. However, the mere recognition that most predators also scavenge at variable rates, which has been traditionally ignored in food‐web and community ecology, leads to a number of emergent interaction routes linking predation and scavenging. The general goal of this review is to draw attention to the main inter‐specific interactions connecting predators (particularly, large mammalian carnivores), their live prey (mainly ungulates), vultures and carrion production in terrestrial assemblages of vertebrates. Overall, we report an intricate network of both direct (competition, facilitation) and indirect (hyperpredation, hypopredation) processes, and provide a conceptual framework for the future development of this promising topic in ecological, evolutionary and biodiversity conservation research. The classic view that scavenging does not affect the population dynamics of consumed organisms is questioned, as multiple indirect top‐down effects emerge when considering carrion and its facultative consumption by predators as fundamental and dynamic components of food webs. Stimulating although challenging research opportunities arise from the study of the interactions among living and detrital or non‐living resource pools in food webs.     

Integrated research of plant functional traits is important for the understanding of ecosystem processes

Current understanding of carbon cycling in terrestrial ecosystem views “quantity” of litter input as a parameter determining the size of soil C pools and soil respired CO₂: quantity of litter input is not considered a driving factor affecting the patterns of terrestrial ecosystem processes. Emma J. Sayer and collaborators demonstrated that this may not be the case! With a neat and elegant, for its simplicity, manipulation experiment, Sayer et al. (this issue) showed how increasing input of leaf litter affects the patterns of root distribution along the soil profile, in a relatively nutrient rich tropical soil. In their study, roots responded rapidly to changes in fresh leaf litter input and appeared to closely follow the patterns of litter decomposition. Until this study, root exploitation of standing litter was seen solely as an adaptation to nutrient shortage in the mineral soil. Feedback processes between leaf litterfall, decomposition and root dynamics are envisaged. With global climate change likely to alter plant productivity and litterfall, similar feedbacks, if confirmed, will need to be included in terrestrial ecosystem C modelling.     

Direct and indirect food web regulation of microbial decomposers in headwater streams

The direct and indirect regulation of primary productivity has been well established in autotrophic‐based ecosystems; however, less is known about the processes affecting decomposers in detrital‐based ecosystems. Because, small headwater, woodland streams are a dominate feature in most ecosystems and are tightly linked to terrestrial detritus, understanding decomposer‐mediated functions in these systems is critical for understanding carbon processes across the landscape. In this light, we conducted a microcosm and mesocosm experiment to test the direct and indirect food web effects on decomposers in small stream ecosystems. The results from the microcosm experiment supported an existing literature, demonstrating that nutrients directly stimulate decomposers and that microbivores directly reduce decomposers. Based on well‐founded food web theory in autotrophic systems, we predicted that fishes from different trophic‐functional guilds would indirectly stimulate decomposers by enhancing dissolved nutrients and by reducing microbivore densities. Our mesocosm experiment partially supported these predictions. Specifically, we found that fishes that consumed mostly terrestrial foods increased decomposers from the bottom–up by enhancing allochthonous nutrient loading into the stream ecosystems. Contrary to our predictions, however, predatory fishes that consume microbivores did not increase decomposers from the top–down. Rather, in streams with the predatory fish species, microbivores increased (rather than decreased) on leaf litter. This may have resulted from an experimental artifact associated with refuge provided by leaf packs. In conclusion, our data demonstrate that decomposers are regulated by similar direct and indirect processes important in autotrophic‐based ecosystems. This provides further evidence that food web processes can regulate leaf decomposition and flux of detrital carbon through ecosystems.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

基于土壤食物网的生态系统复杂性-稳定性关系研究进展

复杂性-稳定性关系是生态学核心问题之一。作为模式食物网,土壤食物网在探索生态系统复杂性-稳定性关系中起了极大的作用。总结了以Moore、de Ruiter、Neutel等为代表的理论生态学家以土壤食物网为工具研究生态系统复杂性-稳定性关系的方法、结论及不足之处,并展望了未来的发展方向。Moore、de Ruiter、Neutel等将土壤食物网功能群生物量数据、土壤食物网Lotka-Volterra模型和面向过程模型三者结合起来,描述相互作用强度大小格局、分室、能流组织形式等复杂性特征;将土壤食物网Lotka-Volterra模型与群落矩阵结合起来分析局域稳定性,进而探讨生态系统复杂性-稳定性关系的一般规律。在此基础上,Moore、de Ruiter、Neutel等证明了与随机食物网相比,真实食物网的相互作用强度格局、分室等复杂性特征提高了生态系统稳定性,生产力与稳定性共同决定了食物链的长度,并指出建立在平衡态基础上的静态土壤食物网模型在探索生态系统复杂性-稳定性关系方面具有较大的不足,动态土壤食物网是未来以土壤食物网为工具研究生态系统复杂性-稳定性关系的发展方向。     

The role of carrion in maintaining biodiversity and ecological processes in terrestrial ecosystems

Carrion provides a resource for a subset of animal species that deliver a critical ecosystem service by consuming dead animal matter and recycling its nutrients. A growing number of studies have also shown various effects of carrion on different plant and microbial communities. However, there has been no review of these studies to bring this information together and identify priority areas for future research. We review carrion ecology studies from the last two decades and summarise the range of spatial and temporal effects of carrion on soil nutrients, microbes, plants, arthropods, and vertebrates. We identify key knowledge gaps in carrion ecology, and discuss how closing these gaps can be achieved by focusing future research on the (1) different kinds of carrion resources, (2) interactions between different components of the carrion community, (3) the ways that ecosystem context can moderate carrion effects, and (4) considerations for carrion management. To guide this research, we outline a framework that builds on the ‘ephemeral resource patch’ concept, and helps to structure research questions that link localised effects of carrion with their consequences at landscape scales. This will enable improved characterisation of carrion as a unique resource pool, provide answers for land managers in a position to influence carrion availability, and establish the ways that carrion affects the dynamics of species diversity and ecological processes within landscapes.     

Warming,eutrophication, and predator loss amplify subsidies between aquatic and terrestrial ecosystems

The exchange of organisms and energy among ecosystems has major impacts on food web structure and dynamics, yet little is known about how climate warming combines with other pervasive anthropogenic perturbations to affect such exchanges. We used an outdoor freshwater mesocosm experiment to investigate the interactive effects of warming, eutrophication, and changes in top predators on the flux of biomass between aquatic and terrestrial ecosystems. We demonstrated that predatory fish decoupled aquatic and terrestrial ecosystems by reducing the emergence of aquatic organisms and suppressing the decomposition of terrestrial plant detritus. In contrast, warming and nutrients enhanced cross‐ecosystem exchanges by increasing emergence and decomposition, and these effects were strongest in the absence of predators. Furthermore, we found that warming advanced while predators delayed the phenology of insect emergence. Our results demonstrate that anthropogenic perturbations may extend well beyond ecosystem boundaries by influencing cross‐ecosystem subsidies. We find that these changes are sufficient to substantially impact recipient communities and potentially alter the carbon balance between aquatic and terrestrial ecosystems and the atmosphere.     

蒙古栎叶片及其土壤碳、氮同位素自然丰度对大气CO2浓度升高的响应

大气CO₂浓度升高对土壤氮素转化过程产生重要影响,研究其变化有助于更好地预测陆地生态系统的固碳潜力.氮同位素自然丰度作为生态系统氮素循环过程的综合指标能够有效地指示CO₂浓度升高对土壤氮素转化过程的影响.本研究采用开顶箱CO₂ 熏蒸法研究连续10年的大气CO₂ 浓度升高对我国东北地区蒙古栎及其土壤和微生物生物量碳、氮同位素自然丰度的影响.结果表明: 大气CO₂浓度升高改变了土壤氮循环过程,增加了土壤微生物和植物叶片δ¹⁵N;促进了富¹³C土壤有机碳分解,中和了贫¹³C植物光合碳输入的效果,导致土壤可溶性有机碳和微生物碳δ¹³C在CO₂升高条件下没有发生显著变化.这些结果表明,CO₂浓度升高很可能促进了土壤有机质矿化过程,并加剧了系统氮限制的状态.     

蒙古栎叶片及其土壤碳、氮同位素自然丰度对大气CO2浓度升高的响应

大气CO₂浓度升高对土壤氮素转化过程产生重要影响,研究其变化有助于更好地预测陆地生态系统的固碳潜力.氮同位素自然丰度作为生态系统氮素循环过程的综合指标能够有效地指示CO₂浓度升高对土壤氮素转化过程的影响.本研究采用开顶箱CO₂ 熏蒸法研究连续10年的大气CO₂ 浓度升高对我国东北地区蒙古栎及其土壤和微生物生物量碳、氮同位素自然丰度的影响.结果表明: 大气CO₂浓度升高改变了土壤氮循环过程,增加了土壤微生物和植物叶片δ¹⁵N;促进了富¹³C土壤有机碳分解,中和了贫¹³C植物光合碳输入的效果,导致土壤可溶性有机碳和微生物碳δ¹³C在CO₂升高条件下没有发生显著变化.这些结果表明,CO₂浓度升高很可能促进了土壤有机质矿化过程,并加剧了系统氮限制的状态.     

Above- and below-ground impacts of introduced predators in seabird-dominated island ecosystems

Predators often exert multi-trophic cascading effects in terrestrial ecosystems. However, how such predation may indirectly impact interactions between above- and below-ground biota is poorly understood, despite the functional importance of these interactions. Comparison of rat-free and rat-invaded offshore islands in New Zealand revealed that predation of seabirds by introduced rats reduced forest soil fertility by disrupting sea-to-land nutrient transport by seabirds, and that fertility reduction in turn led to wide-ranging cascading effects on belowground organisms and the ecosystem processes they drive. Our data further suggest that some effects on the belowground food web were attributable to changes in aboveground plant nutrients and biomass, which were themselves related to reduced soil disturbance and fertility on invaded islands. These results demonstrate that, by disrupting across-ecosystem nutrient subsidies, predators can indirectly induce strong shifts in both above- and below-ground biota via multiple pathways, and in doing so, act as major ecosystem drivers.     

Above‐ and belowground insect herbivores mediate the impact of nitrogen eutrophication on the soil food web in a grassland ecosystem

Insect herbivores are important drivers of ecosystem processes in grasslands, and can mediate the grassland's response to environmental change. For example, recent evidence shows that above‐ and belowground herbivory, individually and in combination, can modify how a plant community responds to nitrogen (N) eutrophication, an important driver of global change. However, knowledge about how such effects extend to the associated soil food web is lacking. In a mesocosm experiment, we investigated how communities of soil nematodes – an abundant and functionally important group of soil organisms – responded to above‐ and belowground insect herbivory at contrasting N levels. We found that the strongest influence of above‐ and belowground herbivory on the nematode community appeared at elevated N. The abundance of root‐feeding nematodes increased when either above‐ or belowground insect herbivores were present at elevated N, but when applied together the two herbivore types cancelled out one another's effect. Additionally, at elevated N aboveground herbivory increased the abundance of fungal‐feeders relative to bacterial‐feeders, which indicates changes in decomposition pathways induced by N and herbivory. Belowground herbivory increased the abundance of omnivorous nematodes. The shifts in both the herbivorous and detrital parts of the soil food web demonstrate that above‐ and belowground herbivory does not only mediate the response of the plant community to N eutrophication, but in extension also the soil food web sustained by the plant community. We conclude that feedbacks between effects of above‐ and belowground herbivory mediate the response of the grassland ecosystem to N eutrophication.