How effective are plant macrofossils as a proxy for macrophyte presence? The case of Najas flexilis in Scotland

© 2017 The Author(s) Preventing biodiversity loss is a key aim of modern conservation, and paleolimnology can inform conservation strategies for target species and habitats where other data are unavailable. Care must be taken to fully understand the possibilities and limits of such techniques, particularly where they concern single species. This study uses plant and seed distribution data to inform macrofossil reconstructions of the rare macrophyte Najas flexilis (Slender Naiad) in Scotland, UK. It answers three questions: (a) How does the location of N. flexilis seeds in the surface sediments relate to the distribution of N. flexilis plants? (b) How do the numbers of seeds in surface sediments correlate with % cover of N. flexilis plants across lakes with differing N. flexilis abundances? (c) What are the implications of these findings for paleolimnology? Percentage N. flexilis cover and number of N. flexilis seeds in surface sediments were recorded at ~100 sample points at each of three sites; one where the species was abundant, one where it was occasional and one where it was extinct. At all sites, N. flexilis seeds were present in surface sediments across the entire lake. No correlation between % cover N. flexilis and the number of seeds in surface sediments was found within individual sites. The distribution of seeds in these lakes appeared to be related to multiple environmental and ecological variables including latitude and longitude (proxies for water currents). This is attributed to the ability of seed-bearing N. flexilis plants to fragment and float large distances on water. Between sites, there was a significant difference in the mean seed counts, with higher mean seed counts corresponding to higher abundances of N. flexilis plants. It is concluded that N. flexilis is likely to be well represented in sediment cores taken from any location within a basin, but that care should be taken when inferring changes in N. flexilis abundance from changes in the numbers of seeds in sediment samples. This work demonstrates that the reproductive ecology (number of seeds produced and dispersal mechanisms) is an important factor to consider when attempting reconstructions of single aquatic plant populations from macrofossil records

Mission possible: diatoms can be used to infer past duckweed (lemnoid Araceae) dominance in ponds

Compared to larger lakes, ponds have rarely been the focus of palaeoecological studies. A common feature of ponds, especially those subject to eutrophication, is mass surface coverings of lemnoid Araceae (duckweed) which have severe implications for ecological processes in small waterbodies, in particular lowered oxygen content. To help understand the implications of duckweed dominance for the long-term ecology of ponds, and to determine the potential for palaeoecological studies in ponds more generally, we develop a new diatom-based Lemna-indicator metric. Recent studies of diatom host-plant relationships have shown significant associations between duckweed and the epiphytes Lemnicola hungarica and Sellaphora saugerresii (formally known as Sellaphora seminulum). To determine the potential of these species as palaeo-indicators of long-term duckweed dynamics in ponds, we investigated the diatom composition of surface sediment assemblages in sets of duckweed and non-duckweed-dominated ponds in Norfolk, eastern England. In addition, we undertook diatom analysis of two cores from a small farmland pond (Bodham Rail Pit) subject to a known duckweed dominance event (1999–2005). Both L. hungarica and S. saugerresii were significant predictors of past Lemna dominance in the surface sediments. Further, in the core study, both diatom species accurately and closely tracked the documented “on–off” duckweed cycle. Our study suggests huge potential for using ponds in palaeoecological studies and for diatom-based investigations of floating plant histories

Recent heavy metal contamination of the Thurne Broads

The banning of tributyltin (TBT) from boat antifouling paints in the late 1980s led to its replacement by alternative biocide additives (Voulvoulis et al., 2000; Marcheselli et al., 2010), including Cu (Dahl & Blanck, 1996) and Zn as active ingredients. It has been reported that Cu and Zn compounds associated with these biocides have caused substantial contamination of harbour and marina sediments (Eklund et al., 2010; Parks et al., 2010), with negative toxic consequences for aquatic organisms (Ytreberg et al., 2010). Indeed, it is evident that Cu and Zn compounds present in paint fragments are readily leached into the water column allowing entry into aquatic food webs (Jessop & Turner, 2011). Nevertheless, relatively little is known regarding antifoulant-derived metals contamination in freshwater lakes. The Norfolk and Suffolk Broads (Eastern England, UK) have been contaminated by antifoulant-derived heavy metals, particularly Cu and Zn which have increased since the banning of TBT (post-1987) in parts of the boated system (Boyle et al., in prep.). Further, recent studies suggest that current levels of sediment contamination by Cu may have negative ecological effects for aquatic ecosystems including inhibition of aquatic macrophyte germination and performance (Boyle et al., submitted; S. Lambert, unpublished data). In Hickling Broad (Thurne Broads system), post-TBT increases in Cu and Zn are also evident, with an interesting peak in Cu for the late 1990s in core HICK1 (Figure 1). This coincides with the large-scale loss of aquatic macrophytes (especially Characeae) from the lake in 1999 (Barker et al., 2008). HICK1 was collected in 2003. In the proposed study we sought to gain a fuller understanding of recent metal contamination in Hickling Broad up to the present day, whilst looking to verify and better contextualise the late 1990s Cu peak. Further, we aimed to determine whether the peak in Cu for Hickling Broad was also recorded at Horsey Mere which is used as a control site in this study i.e. is it just a Hickling phenomenon? Or is it a Thurne Broads system-wide effect

Filling the “data gap”: Using paleoecology to investigate the decline of Najas flexilis (a rare aquatic plant)

In the absence of long‐term monitoring records, paleoecology can be used to extend knowledge of species and community ecology into the past. The rare and declining aquatic plant Najas flexilis is a priority species for conservation across Europe, and is an ideal candidate for paleoecological study; not only are historical records of the plant sparse, but its seeds are commonly found and well preserved in lake sediment cores. In this study, we investigate the timing and causes of decline at two UK sites at which N. flexilis has recently become extinct: Esthwaite Water (England) and Loch of Craiglush (Scotland). For both sites, multiple paleoecological indicators and available historical biological records and monitoring data are compared to numbers of N. flexilis seeds enumerated in dated sediment cores representing the last 150 years. At Esthwaite Water, N. flexilis seeds were found in abundance in association with indicators of a clear, oligo‐mesotrophic, mildly alkaline lake. Eutrophication led to the disappearance of N. flexilis in the 1980s. By contrast, far fewer N. flexilis seeds were found in a core from Loch of Craiglush, and the current period of N. flexilis absence was found to be one of several over the last 100 years. Species represented in cores taken from Loch of Craiglush were indicative of slightly more acidic conditions than Esthwaite Water. Given that N. flexilis favours circumneutral to alkaline conditions, it is possible that Loch of Craiglush has not always been favourable for the plant. These findings have important implications for future conservation efforts, particularly at Esthwaite Water where they suggest that recent failed attempts to reintroduce the species may have been premature. More generally, this study demonstrates the value of paleoecological techniques as a means to provide the long‐term context that is often missing from conservation planning and management

Habitat heterogeneity enables spatial and temporal coexistence of native and invasive macrophytes in shallow lake landscapes

Macrophyte invasive alien species (IAS) fitness is often hypothesised to be associated with beneficial environmental conditions (environmental matching) or species-poor communities. However, positive correlations between macrophyte IAS abundance and native plant richness can also arise, due to habitat heterogeneity (defined here as variation in abiotic and native biotic conditions over space and time). We analysed survey and palaeoecological data for macrophytes in satellite lakes along the Upper Lough Erne (ULE) system (Northern Ireland, UK), covering a gradient of eutrophication and connectivity to partition how environmental conditions, macrophyte diversity and habitat heterogeneity explained the abundance of Elodea canadensis, a widely distributed non-native macrophyte in Europe. E. canadensis abundance positively correlated with macrophyte richness at both the within- and between-lake scales indicating coexistence of native and invasive species over time. E. canadensis was also more prolific in highly connected and macrophyte-rich lakes, but sparser in the more eutrophic-isolated ones. Partial boosted regression trees revealed that in eutrophic-isolated lakes, E. canadensis abundances correlated with water clarity (negatively), plant diversity (positively), and plant cover (negatively) whereas in diverse-connected lakes, beta diversity (both positively and negatively) related to most greatly E. canadensis abundance. Dense macrophyte cover and unfavourable environmental conditions thus appear to confer invasibility resistance and sufficient habitat heterogeneity to mask any single effect of native biodiversity or environmental matching in controlling E. canadensis abundance. Therefore, in shallow lake landscapes, habitat heterogeneity variously enables the coexistence of native macrophytes and E. canadensis, reducing the often-described homogenisation effects of invasive macrophytes

Recovery of the crucian carp Carassius carassius (L.): Approach and early results of an English conservation project

Aquatic Conservation: Marine and Freshwater Ecosystems published by John Wiley & Sons Ltd The crucian carp Carassius carassius, a cyprinid fish characteristic of small ponds, is in decline throughout most of its European range, including in England where it is currently thought to be non-native. The present study, undertaken by the Norfolk Crucian Project, reports on reductions in pond populations of crucian carp in Norfolk, eastern England as well as the success of recent introduction/re-introduction efforts in terms of crucian survival, recruitment and growth over the last 10 years. A 72% decline in crucian carp distribution was observed between the 1950s–1980s and the 2010s. Of 18 crucian carp introductions/re-introductions to restored and suitable existing ponds, 17 were successful in terms of survival, increasing the number of current crucian sites in Norfolk by 37%. Recruitment of young crucian carp was demonstrated for 12 of the 18 stocked ponds, with apparent elevated juvenile growth relative to other English and European populations. Delays in, or a lack of, crucian recruitment in some ponds appeared to result from the presence of other fish species (especially threespine stickleback Gasterosteus aculeatus) with predation and interspecific competition possible contributory factors. This study shows that, through combinations of pond rehabilitation and stocking, it has been possible to achieve a substantial recovery of crucian carp populations in the study region. Although the crucian carp is currently presumed to be non-native within England, given other scientific studies that show a lack of adverse impacts of this species on native biota, and because it is greatly threatened in its native range, the call is sounded for more crucian carp conservation projects in other parts of England as well as in Europe more generally

Toxic metal enrichment and boating intensity: sediment records of antifoulant copper in shallow lakes of eastern England

Tributyltin (TBT), an aqueous biocide derived from antifouling paint pollution, is known to have impacted coastal marine ecosystems, and has been reported in the sediment of the Norfolk and Suffolk Broads, a network of rivers and shallow lakes in eastern England. In the marine environment, the 1987 TBT ban has resulted in expanded use of alternative biocides, raising the question of whether these products too have impacted the Broads ecosystem and freshwaters in general. Here we examine the lake sediment record in the Norfolk and Suffolk Broads for contamination by copper (Cu) (as an active biocide agent) and zinc (Zn) (as a component of booster biocides), to assess their occurrence and potential for causing environmental harm in freshwater ecosystems. We find that, after the introduction of leisure boating, there is a statistically significant difference in Cu enrichment between heavily and lightly boated sites, while no such difference exists prior to this time. At the heavily boated sites the onset of Cu enrichment coincides with a period of rapid increase in leisure boating. Such enrichment is maintained to the present day, with some evidence of continued increase. We conclude that Cu-based antifouling has measurably contaminated lakes exposed to boating, at concentrations high enough to cause ecological harm. Similar findings can be expected at other boated freshwater ecosystems elsewhere in the world

Catchment-scale distribution, abundance, habitat use, and movements of European eel (Anguilla anguilla L.) in a small UK river: Implications for conservation management

Effective conservation management of the Critically Endangered European eel (Anguilla anguilla) is hindered by incomplete understanding of distribution, abundance, and habitat requirements at the catchment scale. All habitats available to eels within a small, highly regulated river catchment, representative of many used across the species' range, were sampled using several methods (including point abundance sample electric fishing and fyke nets) and supplemented by individual telemetry to investigate movements. Eels were found throughout the catchment (59% of n = 131 sites) from the coastal marshes to the headwaters, although the probability of presence declined with distance from the estuary. The lack of a clear relationship with perceived barriers may illustrate a mismatch with the reality experienced by eels, as telemetry identified connectivity across obstacles between paludal habitat and estuary, and detected escapement of mature silver eels from both lotic and lacustrine habitat. Different size/age classes occurred in different parts of the catchment, partly linked to different habitat associations, with coastal paludal habitat supporting >50% of the catchment population and especially smaller (possibly male dominated) yellow eels. Recently recruited elvers were most abundant in the lower reaches of lotic habitat. The largest (probably female) eels were concentrated in lacustrine sites, especially at the ‘end-of-the-line’ in the headwaters. Experiences here suggest that conservation management for eels in small catchments is best focused on: (i) improving connectivity and assisting migration of elvers across ‘problem’ barriers that cannot be removed or modified, (ii) river restoration and rewilding, especially measures that increase instream woody material to provide refuge habitat and, (iii) enhancement or creation (where necessary) of suitable lacustrine habitat to benefit large females in particular. Such action across numerous small river catchments may ultimately help support the recovery of eel stocks

Ponds as insect chimneys: restoring overgrown farmland ponds benefits birds through elevated productivity of emerging aquatic insects

Farmland bird populations have experienced severe declines across Europe and elsewhere. Agricultural intensification is believed to be a main factor behind these declines, with losses of non-cropped features, such as farmland ponds, identified as a key driver. Since the 1950s, many European farmland ponds have been in-filled or, through lack of management, become terrestrialised. Restoring terrestrialised farmland ponds has been shown to significantly increase the abundance and diversity of local farmland bird communities. It has been hypothesised that farmland birds are specifically attracted to open-canopy ponds due to increased emergent aquatic insect availability, but this link has hitherto been little explored. This study investigates how farmland pond management influences emergent aquatic insects, and how emergent insect abundance and biomass is linked to local bird assemblages. Insect emergences showed an 18-fold higher abundance and a 25-fold higher biomass at managed open-canopy ponds in comparison to their unmanaged overgrown counterparts, with day-to-day fluctuations in pond water temperature a key predictor of insect emergences. Species richness and abundance of birds at farmland ponds were strongly positively linked to the abundance of emergent insects. Furthermore, insect emergence peaks occurred on different days in different restored ponds such that the pond landscape afforded extended feeding opportunities for birds. Our findings suggest that restoring networks of terrestrialised farmland ponds to open-canopy macrophyte-dominated conditions could be a highly effective way of increasing the availability of aquatic insect prey for birds. This study highlights an urgent need to re-evaluate pond restoration and management within agri-environmental schemes in Europe and beyond