Carl D. Sayer

DOES THE FISH-INVERTEBRATE-PERIPHYTON CASCADE PRECIPITATE PLANT LOSS IN SHALLOW LAKES?

Alternative equilibria dominated by either submerged plants or phytoplankton have been described for communities in shallow lakes. A nutrient-mediated increase in periphyton (algae attached to plant surfaces) is often described as being responsible for the loss of plants from shallow lakes, yet this violates the stochastic assumptions of alternative equilibria. To determine if periphyton is capable of forcing a switch between these communities, the factors governing the success of aquatic plants were surveyed in 17 plant-dominated shallow lakes in Norfolk (United Kingdom) that varied in nutrient concentration and fish biomass. In these lakes, plant biomass was negatively related to the density of periphyton. However, the density of periphyton on the plants was correlated with the density of grazing invertebrates, not nutrient concentration. In turn, the biomass of fish determined the density of invertebrates. This cascade from fish to periphyton via invertebrates appeared to be evident even though plant-dominated lakes are heterogeneous and complex. Under conditions of plant dominance, periphyton appeared to have a stronger influence on plant growth than phytoplankton. Our data support a model where, within the range of nutrients where alternative equilibria are possible, fish are the prime determinants of community structure in shallow lakes, through a cascading effect of predation on grazing invertebrates influencing the biomass of periphyton and hence, plants. We suggest that the stochasticity required for alternative equilibria in shallow lake communities is often derived from the vagaries of fish colonization and reproduction.

Zooplankton as indicators in lakes: a scientific-based plea for including zooplankton in the ecological quality assessment of lakes according to the European Water Framework Directive (WFD)

With the implementation of the EU Water Framework Directive (WFD), the member states have to classify the ecological status of surface waters following standardised procedures. It was a matter of some surprise to lake ecologists that zooplankton were not included as a biological quality element (BQE) despite their being considered to be an important and integrated component of the pelagic food web. To the best of our knowledge, the decision of omitting zooplankton is not wise, and it has resulted in the withdrawal of zooplankton from many so-far-solid monitoring programmes. Using examples from particularly Danish, Estonian, and the UK lakes, we show that zooplankton (sampled from the water and the sediment) have a strong indicator value, which cannot be covered by sampling fish and phytoplankton without a very comprehensive and costly effort. When selecting the right metrics, zooplankton are cost-efficient indicators of the trophic state and ecological quality of lakes. Moreover, they are important indicators of the success/failure of measures taken to bring the lakes to at least good ecological status. Therefore, we strongly recommend the EU to include zooplankton as a central BQE in the WFD assessments, and undertake similar regional calibration exercises to obtain relevant and robust metrics also for zooplankton as is being done at present in the cases of fish, phytoplankton, macrophytes and benthic invertebrates.

Looking forward through the past : identification of 50 priority research questions in palaeoecology

Summary 1. Priority question exercises are becoming an increasingly common tool to frame future agendas in conservation and ecological science. They are an effective way to identify research foci that advance the field and that also have high policy and conservation relevance. 2. To date there has been no coherent synthesis of key questions and priority research areas for palaeoecology, which combines biological, geochemical and molecular techniques in order to reconstruct past ecological and environmental systems on timescales from decades to millions of years. 3. We adapted a well-established methodology to identify 50 priority research questions in palaeoecology. Using a set of criteria designed to identify realistic and achievable research goals, we selected questions from a pool submitted by the international palaeoecology research community and relevant policy practitioners. This article is protected by copyright. All rights reserved. Accepted Article 4. The integration of online participation, both before and during the workshop, increased international engagement in question selection. 5. The questions selected are structured around six themes: human–environment interactions in the Anthropocene; biodiversity, conservation, and novel ecosystems; biodiversity over long timescales; ecosystem processes and biogeochemical cycling; comparing, combining and synthesizing information from multiple records; and new developments in palaeoecology. 6. Future opportunities in palaeoecology are related to improved incorporation of uncertainty into reconstructions, an enhanced understanding of ecological and evolutionary dynamics and processes, and the continued application of long-term data for better-informed landscape management. 7. Synthesis Palaeoecology is a vibrant and thriving discipline and these 50 priority questions highlight its potential for addressing both pure (e.g. ecological and evolutionary, methodological) and applied (e.g. environmental and conservation) issues related to ecological science and global change.

Comparing RADseq and microsatellites to infer complex phylogeographic patterns, an empirical perspective in the Crucian carp, Carassius carassius, L

The conservation of threatened species must be underpinned by phylogeographic knowledge. This need is epitomized by the freshwater fish Carassius carassius, which is in decline across much of its European range. Restriction site‐associated DNA sequencing (RADseq) is increasingly used for such applications; however, RADseq is expensive, and limitations on sample number must be weighed against the benefit of large numbers of markers. This trade‐off has previously been examined using simulation studies; however, empirical comparisons between these markers, especially in a phylogeographic context, are lacking. Here, we compare the results from microsatellites and RADseq for the phylogeography of C. carassius to test whether it is more advantageous to genotype fewer markers (microsatellites) in many samples, or many markers (SNPs) in fewer samples. These data sets, along with data from the mitochondrial cytochrome b gene, agree on broad phylogeographic patterns, showing the existence of two previously unidentified C. carassius lineages in Europe: one found throughout northern and central‐eastern European drainages and a second almost exclusively confined to the Danubian catchment. These lineages have been isolated for approximately 2.15 m years and should be considered separate conservation units. RADseq recovered finer population structure and stronger patterns of IBD than microsatellites, despite including only 17.6% of samples (38% of populations and 52% of samples per population). RADseq was also used along with approximate Bayesian computation to show that the postglacial colonization routes of C. carassius differ from the general patterns of freshwater fish in Europe, likely as a result of their distinctive ecology.

Relationships between fish feeding guild and trophic structure in English lowland shallow lakes subject to anthropogenic influence: implications for lake restoration

The shallow lakes of Eastern England have been subject to intense anthropogenic pressures including nutrient enrichment and fish stocking. We sought to determine the relationships between fish community structure and other ecosystem characteristics in 28 of these lakes through classification of fish species into piscivorous, zooplanktivorous and benthivorous feeding guilds according to the literature. Canonical correspondence analysis produced clear associations between fish and ecosystem characteristics that generally agreed with other theoretical (e.g. the alternative stable states hypothesis) and empirical studies, but with some important differences. There was a striking lack of relationships between nutrients and other variables, indicating the importance of top-down rather than bottom-up processes as a structuring force in the generally eutrophic study lakes. The presence of submerged (and shoreline) vegetation was associated with a diverse assemblage of apparently co-existing piscivorous (principally pike Esox lucius) and zooplanktivorous species. Perch Perca fluviatilis, a significant predator in other studies, was unimportant and argued to be limited by water quality in the extremely shallow lakes. In contrast, the benthivorous fish guild (principally carp Cyprinus carpio, bream Abramis brama and tench Tinca tinca) essentially represented the inverse of the potential pelagic associations between piscivores/zooplanktivores and vegetation. The introduction of large benthivores to many study lakes could have precipitated a loss of submerged vegetation through direct uprooting during foraging, with the effect of simplifying the fish community being most acute where littoral vegetation was limited by other anthropogenic factors. It is implied that attempts to promote or restore submerged vegetation in these lakes would best target benthivorous species.

The role of cladocerans in tracking long-term change in shallow lake trophic status

Shallow lakes have been affected by a variety of human activities profoundly altering their ecological structure and function. Cladocerans have been used to track change resulting from a variety of drivers at a number of time scales. Aquatic macrophytes are well recognised as reflecting the ecological condition of a lake. Here, we compare the plant macrofossils with the sub-fossil cladoceran assemblages from 20 dated sediment cores. Co-correspondence analysis was used to determine the degree of commonality of change in community composition of the two biological groups through time. This analysis revealed very high levels of agreement in the nature and timing of change at all the sites examined with very high correlation coefficients between the axis 1 scores for macrofossils and cladocerans. Furthermore, at all sites a high proportion of the variance (min 20%, max 54%) in the macrofossil data was explained by the change in the cladoceran assemblage. Sub-fossil macrofossil and cladoceran assemblages, from at least from 1700 AD onwards, were examined in more detail at three sites: Ormesby Great Broad, Felbrigg Lake and Lake Søbygaard. There was very good accord in the main shifts of the cladoceran and macrofossil assemblages at all three sites. This may reflect the long-term shift in the principal focus of primary production from the benthic to the pelagic habitat. We suggest that the combination of their central position in the food-web and the presence of both pelagic and benthic taxa make cladocerans a strong candidate as the single best indicator of (palaeo) ecological condition related to changing trophic status and alteration in food-web structure in shallow lakes.

Diatoms as indicators of environmental change in shallow lakes.

© Cambridge University Press 1999, 2010. Introduction Historically, limnological and paleolimnological research has focused on large and typically deep lakes but in the last two decades there has been a growing interest in smaller and shallower water bodies. Shallow lakes are justifiably considered as a separate lake type, distinguished physically from deeper waters by the fact that they are permanently mixed (polymictic) with a consequent lack of stratification of temperature or oxygen and with increased potential for nutrient recycling and redistribution of seston by physical water circulation patterns (Carrick et al., 1994). Whilst this is a useful distinction, there is no single definition of a shallow lake (Padisak & Reynolds, 2003). Scheffer (1998), in his classic text book, acknowledged a fundamental difference in the behavior, ecological functioning, and biotic communities of shallow waters and arbitrarily selected a mean depth of less than 3 m to define shallowness. For the purposes of this chapter we have chosen to adopt this definition and thereby to focus on lakes where, under a favorable light climate, benthic algae and/or rooted submerged macrophytes may occupy the majority of the lakebed (see also Jeppesen et al., 1997). Under enriched conditions, however, the mechanisms that stabilize the macrophyte communities of shallow lakes may often break down and a transition to pelagic production with phytoplankton dominance occurs (Scheffer et al., 1993; Vadeboncoeur et al., 2003). Importantly, because of these characteristics, shallow lakes are, for the most part, more vulnerable to a given pollutant load than large lakes.

Establishing realistic restoration targets for nutrient-enriched shallow lakes: linking diatom ecology and palaeoecology at the Attenborough Ponds, U.K.

Eutrophication is a major problem for shallow lakes in the U.K. lowlands. Over the last few decades issues relating to the management and restoration of these lakes have come to the fore, including the need to develop methods for establishing realistic restoration targets. One group of lakes for which restoration is a key concern are the Attenborough Ponds in the English Midlands, an interconnected series of shallow, nutrient-enriched gravel-pit lakes. In November 1972, the highly polluted River Erewash was diverted into the Attenborough Ponds. To determine the ecological effects of this event and to establish restoration goals for this lake system, two complimentary studies were undertaken here; (i) monitoring of the ecology and sedimentary representation of diatom communities in a disturbed lake connected to the R. Erewash (Tween Pond: high nutrient concentrations, no submerged macrophytes) and a relatively undisturbed lake isolated from the R. Erewash (Clifton Pond: lower nutrient concentrations, abundant submerged macrophytes), and; (ii) examination of the sedimentary diatom record in a short sediment core collected from the chosen disturbed lake (Tween Pond). The species composition, seasonality, relative productivity and sedimentary representation of planktonic and periphytic diatom communities were very different in Tween and Clifton Ponds. In Tween Pond two major phases of planktonic diatom production/sedimentation were observed during March–May and July–September. By contrast, planktonic diatoms were restricted to a single spring peak in Clifton Pond and after May there was a switch to the dominance of periphytic diatoms associated with the development of submerged macrophytes. The 1972 diversion of the River Erewash was clearly reflected in the sedimentary diatom record from Tween Pond, by; (i) the abrupt shift to planktonic diatom dominance, and; (ii) increasing percentages of late summer–autumn associated planktonic diatom species. These changes suggest both significant nutrient enrichment and the switch from submerged macrophyte to phytoplankton dominance. Numerical matching of pre-1972 diatom assemblages with surface sediment assemblages in the gravel-pits using Principal Components Analysis (PCA) and a squared chord distance measure revealed no close analogues. Nevertheless, similarities between; (i) percentages of different diatom habitat and seasonality groups, and; (ii) pre-1972 and contemporary macrophyte survey data suggested that Clifton Pond is probably a good pre-diversion analogue for habitat structure (e.g. macrophyte biomass, composition and architecture) and phenology (e.g. diatom and macrophyte seasonality) in the Ponds. The practical value of combining space-for-time substitution and palaeoecological approaches in restoration ecology studies of shallow lakes is discussed.

Towards the conservation of crucian carp Carassius carassius: understanding the extent and causes of decline within part of its native English range

The extent and causes of crucian carp Carassius carassius decline were assessed during an initial study of c. 25 ponds in north Norfolk, eastern England, U.K., which was then replicated (a validation study) on another c. 25 ponds in an adjacent area. Of these ponds, c. 40 are known to have contained C. carassius during the 1970s-1980s. In the initial and validation studies, C. carassius were found in only 11 of these ponds, yielding declines of 76% (five of 21 ponds) and 68% (six of 19 ponds), respectively (72% decline overall). Non-native cyprinids, including goldfish Carassius auratus and common carp Cyprinus carpio and their hybrids with C. carassius, were observed in 20% of the ponds. Causes of C. carassius local extinction from 21 ponds were confidently determined as desiccation due to drought, terrestrialization and habitat deterioration, hybridization and competition with non-native cyprinids, agricultural land reclamation and predation (after the introduction of pike Esox lucius). This study led to C. carassius being designated as a Biodiversity Action Plan (BAP) species in the county of Norfolk, the first formal conservation designation for the species in the U.K. The C. carassius BAP plan aims to halt the decline of this much overlooked species through reintroductions and selective stocking of suitable ponds within the native range of the species.

Palaeolimnological records of shallow lake biodiversity change: exploring the merits of single versus multi-proxy approaches

Shallow lakes are among the most threatened ecosystems in the world and many contemporary studies have demonstrated declines in biodiversity due to anthropogenic forcing. Mostly, however, these studies have not covered the full period of human-induced diversity change in lakes which is typically over decades-centuries. Here we provide two examples of palaeoecological studies focussed on reconstructing biodiversity changes in contrasting shallow lake environments that demonstrate the efficacy of the approach—a shallow UK lake and a suite of floodplain lakes (called billabongs) in the Murray-Darling basin, Australia. In the Murray-Darling billabongs, complex sedimentary processes operate, sediment chronologies are less certain and replication of sites is needed to confirm patterns. The combination of sediment records from 10 billabongs showed that diatom diversity changes pre- and post-European (>1850) disturbance were inconsistent; however, reductions in diversity were more common and appear to reflect reductions in macrophyte abundance. At Felbrigg Lake, a multi-proxy study with strong chronological control demonstrated divergent responses of macrophyte, diatom, cladoceran and chironomid richness and diversity to a century of eutrophication. Eutrophication of the site was qualitatively inferred from changes in the macrophyte community in turn reconstructed from plant macrofossils. Benthic cladocerans showed a consistent decline in richness through the record, reflecting the gradual reduction in their macrophyte associated habitat over the past century. Diatom richness and diversity responses were complex, with increases in diversity and richness linked to both increases and decreases in macrophyte species richness and abundance. Chironomid richness and diversity patterns were less consistently linked to eutrophication. The loss of the dominant zooplanktivore (perch) in the 1970s was reflected in the richness and diversity profiles for all groups. Our study reveals clear potential for using sediment cores to infer biodiversity change in shallow lakes and shallow lake regions. However, given the contrasting patterns of diversity change for the different biological groups both in Felbrigg Lake and between Felbrigg and the billabongs, caution is required when interpreting whole-ecosystem biodiversity changes (or the absence of change) based on single as opposed to multi-proxy studies.