Matthew T. O’Hare

A multi-scale hierarchical framework for developing understanding of river behaviour to support river management

This paper introduces this special issue of Aquatic Sciences. It outlines a multi-scale, hierarchical framework for developing process-based understanding of catchment to reach hydromorphology that can aid design and delivery of sustainable river management solutions. The framework was developed within the REFORM (REstoring rivers FOR effective catchment Management) project, funded by the European Union’s FP7 Programme. Specific aspects of this ‘REFORM framework’ and some applications are presented in other papers in this special issue. The REFORM framework is founded on previous hierarchical frameworks, sixteen examples of which are reviewed. However, the REFORM framework has some particular properties that reflect the European context for which it was developed. The framework delineates regional landscapes into nested spatial units at catchment, landscape unit, segment, reach, geomorphic unit and finer scales. Reaches, regardless of their ‘naturalness’, are assigned to a river type based on valley confinement, planform and bed material. Indicators are quantified at each spatial scale to feed three groups of assessments. First, contemporary indicators at reach and geomorphic unit scales investigate present processes, forms and human pressures within each reach. These feed assessments of present reach hydromorphological function/alteration, including whether the reach is functioning appropriately for its type; riparian corridor function and alteration; and hydromorphological adjustment. Second, indicators at catchment to segment scales investigate water and sediment production and delivery to reaches and how these are affected by human pressures now and in the past. These are used to construct an inventory of changes over space and time. Third, historical reach and geomorphic unit scale indicators are used to construct the trajectory of reach-scale changes. Contemporary reach-scale assessments, space–time inventory, and trajectory of changes are then combined to establish how river reaches of different type, subject to different human pressures, and located in different environmental contexts behave in response to changes at all considered spatial scales. These support forecasts of the likely responses of reaches to future scenarios (e.g., changes in climate, land cover, channel interventions).

Changes in rotifer species composition and abundance along a trophic gradient in Loch Lomond, Scotland, UK

Loch Lomond is the second largest body of freshwater in Great Britain. It is a long, narrow lake (36.4 km long, 8.8 km wide). The northern basin is fjord-like and surrounded by a mountainous, base-poor, rocky catchment. In contrast, the southern basin is much broader and shallower with a mainly lowland, calcareous, agricultural catchment. This causes a trophic gradient along the length of the loch that runs from the oligotrophic northern basin to the more mesotrophic southern basin. Rotifer samples were collected at monthly intervals between May and October 2002 at three locations along the length of the loch. More than 12 species were found, the commonest of which were Keratella cochlearis (Gosse) and Trichocerca stylata (Gosse). Although the species composition of the rotifer community varied very little among the sites, rotifer abundance increased markedly from north to south, apparently reflecting the trophic gradient along the length of the loch. The results suggest that rotifer abundance may be a more sensitive indicator of trophic state, and changes in trophic state, than species composition.

Understanding plant community responses to combinations of biotic and abiotic factors in different phases of the plant growth cycle.

Understanding plant community responses to combinations of biotic and abiotic factors is critical for predicting ecosystem response to environmental change. However, studies of plant community regulation have seldom considered how responses to such factors vary with the different phases of the plant growth cycle. To address this deficit we studied an aquatic plant community in an ecosystem subject to gradients in mute swan (Cygnus olor) herbivory, riparian shading, water temperature and distance downstream of the river source. We quantified abundance, species richness, evenness, flowering and dominance in relation to biotic and abiotic factors during the growth-, peak-, and recession-phases of the plant growth cycle. We show that the relative importance of biotic and abiotic factors varied between plant community properties and between different phases of the plant growth cycle. Herbivory became more important during the later phases of peak abundance and recession due to an influx of swans from adjacent pasture fields. Shading by riparian vegetation also had a greater depressing effect on biomass in later seasons, probably due to increased leaf abundance reducing light intensity reaching the aquatic plants. The effect of temperature on community diversity varied between upstream and downstream sites by altering the relative competitiveness of species at these sites. These results highlight the importance of seasonal patterns in the regulation of plant community structure and function by multiple factors.

Long-term trends in Loch Leven invertebrate communities

Detailed studies of the macroinvertebrate benthos and zooplankton communities in Loch Leven, the largest shallow lowland lake in Scotland, UK, were carried out from 1966 to 1973 as part of the International Biological Programme (IBP). The results revealed a reduction in species diversity that was attributed to increasing eutrophication. This work provides a baseline against which the response of the invertebrate communities to subsequent changes in management can be assessed. This article compares macroinvertebrate benthos and zooplankton data from the IBP study with the post-IBP era during which changes at Loch Leven included a 60% reduction in the phosphorus input from external sources and variations in fish stocking rates. Only in recent years has there been evidence of ecological recovery by the invertebrate communities: the number of macroinvertebrate and zooplankton taxa has increased (including taxa considered to be sensitive to nutrient enrichment) and invertebrate abundances have declined. These changes appear to reflect the improvements in water quality and habitat conditions at Loch Leven that have occurred as a result of the recent reduction in nutrient loads, albeit with a substantial delay before any ecological response could be detected. This time lag in recovery has important implications for assessing improvements in the ecological status of other lake systems, as is required by the EU Water Framework Directive.

Invertebrate hydraulic microhabitat and community structure in Callitriche stagnalis Scop. patches

In this paper, we report the structure of the benthic invertebrate community in submerged Callitriche stagnalis Scop. stands in relation to velocity, in a Scottish river, the Blane Water. We compared the community within the macrophyte beds to that of adjacent unvegetated substrate. Callitriche stagnalis stands supported different taxa richness (no. of taxa) and abundances (no. of individuals) of benthic invertebrates than the unvegetated riffle substrate (Wilcoxon Matched-Pairs test: p<0.01, n = 10). To see if different sections of the stands supported different invertebrate communities, samples were taken from the outer, mid and root sections of the stands. The sections were separated by Detrended Correspondence Analysis (DCA) using their macroinvertebrate communities (eigenvalues: axis 1 = 0.6, axis 2 = 0.136). Abundance and taxa richness were different between the outer and mid sections, and between outer and root sections (Wilcoxon Matched-Pairs test: respectively p<0.05, p<0.01, n = 10). Community structure also differed between sections. The outer section community had a structure similar to that of an extreme environment where Simuliidae were dominant. Ephemerella ignita dominated in the mid and root sections. Differing combinations of plant structure and velocity appear to be a major factor influencing habitat structure, creating a range of stability conditions in the stands, which support the observed diversity of invertebrate assemblages present.

Evaluating the effects of population management on a herbivore grazing conflict.

Abundant herbivores can damage plants and so cause conflict with conservation, agricultural, and fisheries interests. Management of herbivore populations is a potential tool to alleviate such conflicts but may raise concerns about the economic and ethical costs of implementation, especially if the herbivores are ‘charismatic’ and popular with the public. Thus it is critical to evaluate the probability of achieving the desired ecological outcomes before proceeding to a field trial. Here we assessed the potential for population control to resolve a conflict of non-breeding swans grazing in river catchments. We used a mathematical model to evaluate the consequences of three population management strategies; (a) reductions in reproductive success, (b) removal of individuals, and (c) reduced reproductive success and removal of individuals combined. This model gave accurate projections of historical changes in population size for the two rivers for which data were available. Our model projected that the River Frome swan population would increase by 54%, from 257 to 397 individuals, over 17 years in the absence of population control. Removal of ≥60% of non-breeding individuals each year was projected to reduce the catchment population below the level for which grazing conflicts have been previously reported. Reducing reproductive success, even to 0 eggs per nest, failed to achieve the population reduction required. High adult and juvenile survival probabilities (>0.7) and immigration from outside of the catchment limited the effects of management on population size. Given the high, sustained effort required, population control does not represent an effective management option for preventing the grazing conflicts in river catchments. Our study highlights the need to evaluate the effects of different management techniques, both alone and in combination, prior to field trials. Population models, such as the one presented here, can provide a cost-effective and ethical means of such evaluations.

Can Sacrificial Feeding Areas Protect Aquatic Plants from Herbivore Grazing? Using Behavioural Ecology to Inform Wildlife Management

Effective wildlife management is needed for conservation, economic and human well-being objectives. However, traditional population control methods are frequently ineffective, unpopular with stakeholders, may affect non-target species, and can be both expensive and impractical to implement. New methods which address these issues and offer effective wildlife management are required. We used an individual-based model to predict the efficacy of a sacrificial feeding area in preventing grazing damage by mute swans (Cygnus olor) to adjacent river vegetation of high conservation and economic value. The accuracy of model predictions was assessed by a comparison with observed field data, whilst prediction robustness was evaluated using a sensitivity analysis. We used repeated simulations to evaluate how the efficacy of the sacrificial feeding area was regulated by (i) food quantity, (ii) food quality, and (iii) the functional response of the forager. Our model gave accurate predictions of aquatic plant biomass, carrying capacity, swan mortality, swan foraging effort, and river use. Our model predicted that increased sacrificial feeding area food quantity and quality would prevent the depletion of aquatic plant biomass by swans. When the functional response for vegetation in the sacrificial feeding area was increased, the food quantity and quality in the sacrificial feeding area required to protect adjacent aquatic plants were reduced. Our study demonstrates how the insights of behavioural ecology can be used to inform wildlife management. The principles that underpin our model predictions are likely to be valid across a range of different resource-consumer interactions, emphasising the generality of our approach to the evaluation of strategies for resolving wildlife management problems.

Plants in aquatic ecosystems: current trends and future directions

Aquatic plants fulfil a wide range of ecological roles, and make a substantial contribution to the structure, function and service provision of aquatic ecosystems. Given their well-documented importance in aquatic ecosystems, research into aquatic plants continues to blossom. The 14th International Symposium on Aquatic Plants, held in Edinburgh in September 2015, brought together 120 delegates from 28 countries and six continents. This special issue of Hydrobiologia includes a select number of papers on aspects of aquatic plants, covering a wide range of species, systems and issues. In this paper, we present an overview of current trends and future directions in aquatic plant research in the early twenty first century. Our understanding of aquatic plant biology, the range of scientific issues being addressed and the range of techniques available to researchers have all arguably never been greater; however, substantial challenges exist to the conservation and management of both aquatic plants and the ecosystems in which they are found. The range of countries and continents represented by conference delegates and authors of papers in the special issue illustrates the global relevance of aquatic plant research in the early twenty first century but also the many challenges that this burgeoning scientific discipline must address.

Preface: plants in hydrosystems: from functional ecology to weed research

Aquatic plants are important components of aquatic ecosystems (Chambers et al., 2008). They produce consumable materials that form the basis of trophic networking, they influence the hydrological, geomorphological and chemical environments and they interact in different ways with microbial and animal compartments and ecosystem processes. The interplay between humans and aquatic environments is often mediated through aquatic plants to both positive and negative ends. The biological traits of those plants and their communities are an essential aid to understanding the role of aquatic vegetation and their ecological interactions across different spatial and time scales (Capers et al., 2010; Gurnell et al., 2010; Bornette & Puijalon, 2011). For example, combinations of environmental conditions and traits can lead to invasive processes harmful to human activities and ecosystems (Gordon & Gantz, 2011). But plants can also be used to mitigate human impacts, such as the use of constructed macrophyte beds for nutrient removal (Xian et al., 2010). A mechanistic understanding of plant-ecosystem interactions, and how they are perturbed by human disturbances, enables the monitoring of the macrophyte communities and water bodies where they exist, and ultimately provides the clues and tools to protect and rehabilitate aquatic environments and plant populations (Lumbreras et al., 2013; Aguiar et al., 2014). The present volume is a contribution to such understanding of ecological interactions and its applications to freshwater management.

Responses of aquatic plants to eutrophication in rivers: A revised conceptual model

Compared to research on eutrophication in lakes, there has been significantly less work carried out on rivers despite the importance of the topic. However, over the last decade, there has been a surge of interest in the response of aquatic plants to eutrophication in rivers. This is an area of applied research and the work has been driven by the widespread nature of the impacts and the significant opportunities for system remediation. A conceptual model has been put forward to describe how aquatic plants respond to eutrophication. Since the model was created, there have been substantial increases in our understanding of a number of the underlying processes. For example, we now know the threshold nutrient concentrations at which nutrients no longer limit algal growth. We also now know that the physical habitat template of rivers is a primary selector of aquatic plant communities. As such, nutrient enrichment impacts on aquatic plant communities are strongly influenced, both directly and indirectly, by physical habitat. A new conceptual model is proposed that incorporates these findings. The application of the model to management, system remediation, target setting, and our understanding of multi-stressor systems is discussed. We also look to the future and the potential for new numerical models to guide management.