Value of long‐term ecological studies

Long‐term ecological studies are critical for providing key insights in ecology, environmental change, natural resource management and biodiversity conservation. In this paper, we briefly discuss five key values of such studies. These are: (1) quantifying ecological responses to drivers of ecosystem change; (2) understanding complex ecosystem processes that occur over prolonged periods; (3) providing core ecological data that may be used to develop theoretical ecological models and to parameterize and validate simulation models; (4) acting as platforms for collaborative studies, thus promoting multidisciplinary research; and (5) providing data and understanding at scales relevant to management, and hence critically supporting evidence‐based policy, decision making and the management of ecosystems. We suggest that the ecological research community needs to put higher priority on communicating the benefits of long‐term ecological studies to resource managers, policy makers and the general public. Long‐term research will be especially important for tackling large‐scale emerging problems confronting humanity such as resource management for a rapidly increasing human population, mass species extinction, and climate change detection, mitigation and adaptation. While some ecologically relevant, long‐term data sets are now becoming more generally available, these are exceptions. This deficiency occurs because ecological studies can be difficult to maintain for long periods as they exceed the length of government administrations and funding cycles. We argue that the ecological research community will need to coordinate ongoing efforts in an open and collaborative way, to ensure that discoverable long‐term ecological studies do not become a long‐term deficiency. It is important to maintain publishing outlets for empirical field‐based ecology, while simultaneously developing new systems of recognition that reward ecologists for the use and collaborative sharing of their long‐term data sets. Funding schemes must be re‐crafted to emphasize collaborative partnerships between field‐based ecologists, theoreticians and modellers, and to provide financial support that is committed over commensurate time frames.     

Developing a vision for improved monitoring and reporting of riparian restoration projects

Representatives from agencies involved in natural resource management in the Murray‐Darling Basin gathered for a workshop in November 2010 to develop a vision for improved monitoring and reporting of riparian restoration projects. The resounding message from this workshop was that the effectiveness of riparian restoration depends on having sound, documented and agreed evidence on the ecological responses to restoration efforts. Improving our capacity to manage and restore riparian ecosystems is constrained by (i) a lack of ecological evidence on the effects of restoration efforts, and (ii) short‐termism in commitment to restoration efforts, in funding of monitoring and in expected time spans for ecosystem recovery. Restoration at the effective spatial scope will invariably require a long‐term commitment by researchers, funding agencies, management agencies and landholders. To address the knowledge gaps that constrain riparian restoration in the Basin, participants endorsed four major fields for future research: the importance of landscape context to restoration outcomes; spatio‐temporal scaling of restoration outcomes; functional effects of restoration efforts; and developing informative and effective indicators of restoration. To improve the monitoring and restoration of riparian zones throughout the Basin, participants advocated an integrated approach: a hierarchical adaptive management framework that incorporates long‐term ecological research.     

Coordinated approaches to quantify long‐term ecosystem dynamics in response to global change

Many serious ecosystem consequences of climate change will take decades or even centuries to emerge. Long‐term ecological responses to global change are strongly regulated by slow processes, such as changes in species composition, carbon dynamics in soil and by long‐lived plants, and accumulation of nutrient capitals. Understanding and predicting these processes require experiments on decadal time scales. But decadal experiments by themselves may not be adequate because many of the slow processes have characteristic time scales much longer than experiments can be maintained. This article promotes a coordinated approach that combines long‐term, large‐scale global change experiments with process studies and modeling. Long‐term global change manipulative experiments, especially in high‐priority ecosystems such as tropical forests and high‐latitude regions, are essential to maximize information gain concerning future states of the earth system. The long‐term experiments should be conducted in tandem with complementary process studies, such as those using model ecosystems, species replacements, laboratory incubations, isotope tracers, and greenhouse facilities. Models are essential to assimilate data from long‐term experiments and process studies together with information from long‐term observations, surveys, and space‐for‐time studies along environmental and biological gradients. Future research programs with coordinated long‐term experiments, process studies, and modeling have the potential to be the most effective strategy to gain the best information on long‐term ecosystem dynamics in response to global change.     

MICROALGAE AND CYANOBACTERIA: FOOD FOR THOUGHT1

In non‐Western civilizations, cyanobacteria have been part of the human diet for centuries. Today, microalgae and cyanobacteria are either produced in controlled cultivation processes or harvested from the natural habitats and marketed as food supplements around the world. Cyanobacteria produce a vast array of different biologically active compounds, some of which are expected to be used in drug development. The fact that some of the active components from cyanobacteria potentially have anticancer, antimicrobial, antiviral, anti‐inflammatory, and other effects is being used for marketing purposes. However, introduction of these products in the form of whole biomass for alimentary purposes raises concerns regarding the potential toxicity and long‐term effects on human health. Here, we review data on the use of cyanobacteria and microalgae in human nutrition and searched for available information on legislature that regulates the use of these products. We have found that, although the quality control of these products is most often self‐regulated by the manufacturers, different governmental agencies are introducing strict regulations for placing novel products, such as algae and cyanobacteria, on the market. The existing regulations require these products to be tested for the presence of toxins, such as microcystin; however, other, sometimes novel, toxins remain undetected, and their long‐term effects on human health remain unknown.     

Post‐Project Appraisals of River Restoration in Advanced University Instruction

Post‐project appraisals (PPAs) are systematic assessments of built restoration projects, which provide feedback on performance of restoration approaches to improve future restoration efforts. Unfortunately, most restoration projects are not subject to systematic assessment because of lack of institutional arrangements to sustain long‐term evaluation and the orientation of most funding agencies towards project implementation rather than “studies.” As semester‐long courses on river restoration increasingly appear in university curricula at the graduate and advanced undergraduate level, independent student research projects for such courses can provide a mechanism for building a database of PPAs (and components thereof) and providing the students with a powerful learning experience. In two UC Berkeley courses, we require independent student projects involving original field research, peer review of first drafts, instructor (and often outside) review of second drafts, and presentation of results to a public symposium. Since 1995, the revised, final papers have been added to the University of California library, where they constitute one of the largest collections of restoration‐related studies currently available for any region: over 300 restoration‐related studies, of which 80 are PPAs or components thereof. Since 2003, the papers have been posted on‐line, with 40,000 full text downloads through 2010. Some term projects have directly influenced river restoration programs, inducing changes in salmon habitat enhancement project design, documenting failure of projects based on inappropriate restoration approaches, and contributing to systematic assessments of step‐pool and compound channel designs in urban areas. Student evaluations cite the term projects as valuable learning experiences.     

Long-term studies of freshwater macroinvertebrates: a review of the frequency, duration and ecological significance

1. The importance of a long‐term ecological perspective is well documented, yet the availability of long‐term data remains limited. This paper highlights the value of long‐term ecological studies of freshwater macroinvertebrates by reviewing both the availability of long‐term data and recent ecological contributions based on them. 2. A survey of recent literature on stream macroinvertebrates identified 46 papers published between 1987 and 2004 that included long‐term (i.e. ≥5 years) data. Most recently published long‐term studies of stream macroinvertebrates began collecting data in the 1970s and 1980s and their duration (time between first and last year sampled) was relatively brief (median = 9 years, maximum = 96 years). Most studies did not expand their temporal perspective by incorporating older data collected by other researchers. 3. Recent long‐term studies of macroinvertebrates have made major contributions to our understanding of interannual variation and cycles, complex abiotic and biotic interactions, and natural and anthropogenic disturbance and recovery. Without these studies, we would know much less about the magnitude of natural temporal variation, the importance of physical and biological disturbance and interactions, the role of pathogens and introduced species, the overall impact of pollution and the effectiveness of protection and remediation efforts. 4. If we are to encourage long‐term perspectives in our science, we need to facilitate the transfer of individual studies, as well as knowledge and data, among scientists. This includes efforts to archive and annotate data more effectively, so that they can be more easily incorporated into future research.     

Predicting the likely response of data‐poor ecosystems to climate change using space‐for‐time substitution across domains

Predicting ecological response to climate change is often limited by a lack of relevant local data from which directly applicable mechanistic models can be developed. This limits predictions to qualitative assessments or simplistic rules of thumb in data‐poor regions, making management of the relevant systems difficult. We demonstrate a method for developing quantitative predictions of ecological response in data‐poor ecosystems based on a space‐for‐time substitution, using distant, well‐studied systems across an inherent climatic gradient to predict ecological response. Changes in biophysical data across the spatial gradient are used to generate quantitative hypotheses of temporal ecological responses that are then tested in a target region. Transferability of predictions among distant locations, the novel outcome of this method, is demonstrated via simple quantitative relationships that identify direct and indirect impacts of climate change on physical, chemical and ecological variables using commonly available data sources. Based on a limited subset of data, these relationships were demonstrably plausible in similar yet distant (>2000 km) ecosystems. Quantitative forecasts of ecological change based on climate‐ecosystem relationships from distant regions provides a basis for research planning and informed management decisions, especially in the many ecosystems for which there are few data. This application of gradient studies across domains – to investigate ecological response to climate change – allows for the quantification of effects on potentially numerous, interacting and complex ecosystem components and how they may vary, especially over long time periods (e.g. decades). These quantitative and integrated long‐term predictions will be of significant value to natural resource practitioners attempting to manage data‐poor ecosystems to prevent or limit the loss of ecological value. The method is likely to be applicable to many ecosystem types, providing a robust scientific basis for estimating likely impacts of future climate change in ecosystems where no such method currently exists.     

Ecosystem engineering in the Quaternary of the West Coast of South Africa

Despite advances in our understanding of the geographic and temporal scope of the Paleolithic record, we know remarkably little about the evolutionary and ecological consequences of changes in human behavior. Recent inquiries suggest that human evolution reflects a long history of interconnections between the behavior of humans and their surrounding ecosystems (e.g., niche construction). Developing expectations to identify such phenomena is remarkably difficult because it requires understanding the multi‐generational impacts of changes in behavior. These long‐term dynamics require insights into the emergent phenomena that alter selective pressures over longer time periods which are not possible to observe, and are also not intuitive based on observations derived from ethnographic time scales. Generative models show promise for probing these potentially unexpected consequences of human‐environment interaction. Changes in the uses of landscapes may have long term implications for the environments that hominins occupied. We explore other potential proxies of behavior and examine how modeling may provide expectations for a variety of phenomena.     

Predicting restoration outcomes based on organizational and ecological factors

For restoration to be an effective strategy to reverse large‐scale habitat loss and land degradation, funding programs need policies that promote selection of and commitment to projects that can reasonably be expected to succeed. Programmatic project selection practices have received minimal formal evaluation, despite their importance. In this study, we considered the extent to which a program needs to consider both ecological and organizational factors during project selection in order to minimize the incidence of project failure. Our assessment of a long‐term program that funds ecological restoration efforts across Minnesota (U.S.A.), based on project records, manager surveys, and field surveys, yielded several broadly relevant insights. First, factors well understood to confer ecological resilience (level of landscape alteration and starting condition) were clearly associated with restoration outcomes, regardless of time‐since‐initiation of restoration. Second, restoration of low‐resilience ecosystems is typically a labor‐ and skill‐intensive enterprise for organizations that undertake them. Our analysis revealed four organizational limitations, in addition to insufficient funds, that hindered capacity to keep projects on‐track: lack of planning and goal‐setting, inadequate staffing, leadership change, and incomplete records. Third, to reduce risk, programs do not necessarily need to avoid challenging projects, but do need to consider whether organizations proposing restorations have adequate internal capacity to competently plan and to sustain actions for a duration sufficient to restore ecological resilience. If a restoration is degraded enough to require human intervention to recover, the outcome of a project is as likely to reflect its organizational reality as much as its ecological circumstances.     

How to infer population trends in sparse data: examples with opportunistic sighting records for great white sharks

Aim Biodiversity is declining at accelerating rates. Understanding past and ongoing changes to biodiversity is paramount in prioritizing conservation action and restoring functional ecosystems. Yet long‐term, systematic data on the distribution and abundance of species are sparse. For many organisms, specimen collections and anecdotal accounts of chance sightings or captures constitute the only source of information. Such data have the potential to provide valuable insights on long‐term ecosystem changes, but are often neglected because they are difficult to analyse quantitatively. Here we review available methods and introduce a new approach. Location Historic data on sightings and captures of great white sharks in the eastern Adriatic and off eastern Canada serve to illustrate the utility of both the existing methods and the new approach. Method Unlike existing methods, the new approach focuses on estimating population trends rather than verifying extinction and explicitly addresses uncertainty over observation effort via two tiers of sensitivity analysis. It fits a series of generalized linear models that provide multiple estimates of declines under alternate scenarios regarding the appropriate reference period and observer trends. Programming code to implement the approach in freely available software is provided as supplementary material. Results Example analyses of great white shark sightings suggest that local populations of this species have suffered dramatic declines, both in the eastern Adriatic and along Canada’s eastern coast. Although not yet extinct, this top predator may therefore no longer be able to fulfil its former ecological role. Main conclusions Careful quantitative analyses of imperfect historical data can provide valuable insights into past ecological changes. Such insights are crucial to improved management and restoration of individual species and their ecosystems. We therefore hope that our review of available methods will facilitate quantitative evaluations of species for which analysis was previously impeded by a lack of systematically collected data.     

What are the policy priorities for sustaining ecological processes? A case study from Victoria,Australia

Summary Developments in ecological theory indicate that ecological processes have major implications for sustaining biodiversity and the provision of ecosystem services. Consequently, conservation actions that focus solely on particular species, vegetation communities, habitats or sites (‘assets’) are unlikely to be effective over the long term unless the ecological processes that support them continue to function. Efforts to sustain biodiversity must embrace both ‘assets’ and ‘process‐oriented’ approaches. Existing knowledge about ecological processes, incomplete though it is, has not been adequately considered in government decision making. It is, therefore, necessary to consider how to build consideration of ecological processes into legislative and institutional frameworks, policy and planning processes, and on‐ground environmental management. Drawing on insights from interviews, a facilitated workshop, and a literature review, this paper identifies a suite of policy priorities and associated reforms which should assist in ensuring that ecological processes are given more attention in policy‐making processes. It is concluded that a multi‐pronged approach is required, because there are no ‘silver bullets’ for sustaining ecological processes.     

Robust ecological drought projections for drylands in the 21st century

Dryland ecosystems may be especially vulnerable to expected 21st century increases in temperature and aridity because they are tightly controlled by moisture availability. However, climate impact assessments in drylands are difficult because ecological dynamics are dictated by drought conditions that are difficult to define and complex to estimate from climate conditions alone. In addition, precipitation projections vary substantially among climate models, enhancing variation in overall trajectories for aridity. Here, we constrain this uncertainty by utilizing an ecosystem water balance model to quantify drought conditions with recognized ecological importance, and by identifying changes in ecological drought conditions that are robust among climate models, defined here as when >90% of models agree in the direction of change. Despite limited evidence for robust changes in precipitation, changes in ecological drought are robust over large portions of drylands in the United States and Canada. Our results suggest strong regional differences in long‐term drought trajectories, epitomized by chronic drought increases in southern areas, notably the Upper Gila Mountains and South‐Central Semi‐arid Prairies, and decreases in the north, particularly portions of the Temperate and West‐Central Semi‐arid Prairies. However, we also found that exposure to hot‐dry stress is increasing faster than mean annual temperature over most of these drylands, and those increases are greatest in northern areas. Robust shifts in seasonal drought are most apparent during the cool season; when soil water availability is projected to increase in northern regions and decrease in southern regions. The implications of these robust drought trajectories for ecosystems will vary geographically, and these results provide useful insights about the impact of climate change on these dryland ecosystems. More broadly, this approach of identifying robust changes in ecological drought may be useful for other assessments of climate impacts in drylands and provide a more rigorous foundation for making long‐term strategic resource management decisions.     

A proposed process for applying a structured decision‐making framework to restoration planning in the Atchafalaya River Basin,Louisiana, U.S.A.

Environmental decision‐making issues in the Atchafalaya River Basin (ARB), Louisiana require innovative approaches that combine scientific understanding and local stakeholder values. Management of the ARB has evolved from strong federal control to establish the ARB as a primary floodway of the Mississippi River and Tributaries Project to a state and federal collaboration to accommodate fish and wildlife resource promotion, recreational opportunities, and economic development. The management policy has expanded to include a growing number of stakeholders, but the decision‐making process has not kept pace. Current conflicts among many local stakeholder groups, due in part to their lack of involvement in the decision‐making process, impede restoration efforts. The absence of a long‐term collective vision for the ARB by both local stakeholder groups and management agencies further confounds these efforts. This paper proposes a process to apply a structured decision‐making framework, a values‐based approach that explicitly defines objectives, to promote stakeholder‐driven restoration efforts in the ARB and to better prepare for and manage long‐term environmental issues. The goals of this approach are: (1) to create a process founded on stakeholder values and supported by rigorous scientific assessment to meet management agency mandates and (2) to establish a transparent process for restoration planning in the ARB that incorporates current and future non‐governmental stakeholders into the decision‐making process. Similar frameworks have been successful in other river basins; we feel the structure of current restoration efforts in the ARB is well‐suited to adopt a values‐focused management framework.     

A survey of long‐term terrestrial ecology studies in Australia

Long‐term ecological studies (LTES) are critical for understanding and managing landscapes. To identify important research gaps, facilitate collaborations and communicate results, several countries have established long‐term ecological research networks. A few initiatives to create such a network in Australia have been undertaken, but relatively few published data exist on the current state of LTES in Australia. In this paper, we present the results of an online survey of terrestrial LTES projects sent to academic, government and non‐governmental organization‐based researchers across Australia. We asked questions pertaining to the focus, scope, support and outcomes of LTES spanning 7 years or longer. Based on the information reported from 85 Australian LTES, we: (i) identify the biomes, processes and species that are under‐represented in the current body of research; (ii) discuss important contributing factors to the successful development and survival of these projects; and (iii) make recommendations to help increase the productivity and influence of LTES across research, management and policy sectors.     

Improving inferences from short‐term ecological studies with Bayesian hierarchical modeling: white‐headed woodpeckers in managed forests

Pilot studies are often used to design short‐term research projects and long‐term ecological monitoring programs, but data are sometimes discarded when they do not match the eventual survey design. Bayesian hierarchical modeling provides a convenient framework for integrating multiple data sources while explicitly separating sample variation into observation and ecological state processes. Such an approach can better estimate state uncertainty and improve inferences from short‐term studies in dynamic systems. We used a dynamic multistate occupancy model to estimate the probabilities of occurrence and nesting for white‐headed woodpeckers Picoides albolarvatus in recent harvest units within managed forests of northern California, USA. Our objectives were to examine how occupancy states and state transitions were related to forest management practices, and how the probabilities changed over time. Using Gibbs variable selection, we made inferences using multiple model structures and generated model‐averaged estimates. Probabilities of white‐headed woodpecker occurrence and nesting were high in 2009 and 2010, and the probability that nesting persisted at a site was positively related to the snag density in harvest units. Prior‐year nesting resulted in higher probabilities of subsequent occurrence and nesting. We demonstrate the benefit of forest management practices that increase the density of retained snags in harvest units for providing white‐headed woodpecker nesting habitat. While including an additional year of data from our pilot study did not drastically alter management recommendations, it changed the interpretation of the mechanism behind the observed dynamics. Bayesian hierarchical modeling has the potential to maximize the utility of studies based on small sample sizes while fully accounting for measurement error and both estimation and model uncertainty, thereby improving the ability of observational data to inform conservation and management strategies.     

Neglect of biological riches: the economics of nature tourism in Nepal

Nepal's spectacular parks and reserves have attracted dramatically increasing numbers of foreign visitors. It might be expected that these protected areas would be nurtured as valuable and irreplaceable economic assets. However they are becoming seriously degraded and the financial resources provided for their management have been inadequate. This paper explores why — starting with the hypothesis that so little of the economic value of protected area tourism in Nepal is captured through fees and other charges assessed on foreign visitors that the protected areas are perceived as being of inconsequential value. It is conservatively estimated that $27 million of tourists' total expenditure in Nepal were attributable to the protected area network in 1988, when the costs of managing the parks were less than $5 million but direct fees colleeted from tourists visiting the protected areas amounted to less than $1 million. These figures suggest the parks are a good investment. But it could also be argued that the costs of park management were more than five times the revenues collected by the government from park tourists. Policy measures are identified which could help Nepal increase the economic as well as environmental benefits from nature tourism.Case studies of Nepal's most-visited protected areas emphasize that the lack of funds for protected area management is not the only constraint on effective management. Some important economic and institutional interests have yet to be effectively reconciled with conservation in the protected areas. Most problematic are local people's economic aspirations and the operating practices of the principal government agencies involved — the Department of National Parks and Wildlife Conservation and the Ministry of Tourism. Fortunately there have recently been some encouraging signs of change within both of these agencies.     

Standard of Care,Institutional Obligations,and Distributive Justice

The problem of standard of care in clinical research concerns the level of treatment that investigators must provide to subjects in clinical trials. Commentators often formulate answers to this problem by appealing to two distinct types of obligations: professional obligations and natural duties. In this article, I investigate whether investigators also possess institutional obligations that are directly relevant to the problem of standard of care, that is, those obligations a person has because she occupies a particular institutional role. I examine two types of institutional contexts: (1) public research agencies – agencies or departments of states that fund or conduct clinical research in the public interest; and (2) private‐for‐profit corporations. I argue that investigators who are employed or have their research sponsored by the former have a distinctive institutional obligation to conduct their research in a way that is consistent with the state's duty of distributive justice to provide its citizens with access to basic health care, and its duty to aid citizens of lower income countries. By contrast, I argue that investigators who are employed or have their research sponsored by private‐for‐profit corporations do not possess this obligation nor any other institutional obligation that is directly relevant to the ethics of RCTs. My account of the institutional obligations of investigators aims to contribute to the development of a reasonable, distributive justice‐based account of standard of care.     

Retrospective monitoring of rangeland vegetation change: ecohistory from deposits of sheep dung associated with shearing sheds

This paper explores the potential of a new method of reconstructing historical vegetation change in the Australian rangelands. Historical monitoring of rangeland vegetation has been so deficient that it is not possible to determine whether a long‐term trend toward degradation has occurred (as is often assumed) or, indeed, if it is continuing to occur. Because long‐term records are unavailable any attempt to monitor vegetation retrospectively must be based on proxy measures rather than direct observation. Where historical data are lacking an integration of palaeoecological, archaeological and ecological methods is required to reconstruct the past. Our research is based on a detailed analysis of sheep faeces deposited near a shearing shed in the semiarid rangelands of south‐west Queensland between the late 1930s and the mid‐1990s. The faeces in these deposits represent the diet of sheep in the days leading up to the property’s annual shearing and as such are a potentially useful index to changes in vegetation. Results indicate significant changes in the diet of sheep since the late 1940s. The potential of this method, and its limitations, are discussed. Long‐term records are critical in understanding issues of sustainability in land management and it is intended that this paper will stimulate further research into historical vegetation change in rangelands.     

A test of fundamental questions in mimicry theory using long‐term datasets

Since the phenomenon of mimicry was first described by Bates in 1862 it has become one of the foundational examples of adaptive evolution. Numerous subcategories of mimicry and dozens of hypotheses pertaining to its evolution and maintenance have been proposed. Many of these hypotheses, however, are difficult to test in experimental settings, and data from natural observations are often inadequate. Here we use data from a long‐term survey of butterfly presence and abundance to test several hypotheses pertaining to Batesian and female‐limited polymorphic mimicry (FPM; a special case of Batesian mimicry). We found strong evidence that models outnumber mimics in both mimicry systems, but no evidence for an increase in relative abundance of FPM mimics to their Batesian counterparts. Tests of the early‐emergence/model first hypothesis showed strong evidence that the Batesian mimic routinely emerges after the model, while emergence timing in the FPM system was site specific, suggesting that other ecological factors are at play. These results demonstrate the importance of long‐term field observations for testing evolutionary and ecological hypotheses.     

Decoupled trophic responses to long‐term recovery from acidification and associated browning in lakes

Increases in the concentration of dissolved organic matter (DOM) have been documented in many inland waters in recent decades, a process known as “browning”. Previous studies have often used space‐for‐time substitution to examine the direct consequences of increased DOM on lake ecosystems. However, browning often occurs concomitant with other ecologically important water chemistry changes that may interact with or overwhelm any potential ecological response to browning itself. Here we examine a long‐term (~20 year) dataset of 28 lakes in the Adirondack Park, New York, USA, that have undergone strong browning in response to recovery from acidification. With these data, we explored how primary producer and zooplankton consumer populations changed during this time and what physical and chemical changes best predicted these long‐term ecosystem changes. Our results indicate that changes in primary producers are likely driven by reduced water clarity due to browning, independent of changes in nutrients, counter to previously hypothesized primary producer response to browning. In contrast, declines in calcium concomitant with browning play an important role in driving long‐term declines in zooplankton biomass. Our results indicate that responses to browning at different trophic levels are decoupled from one another. Concomitant chemical changes have important implications for our understanding of the response of aquatic ecosystems to browning.