Temporary pond ecosystem functioning shifts mediated by the exotic red swamp crayfish (<Emphasis Type="Italic">Procambarus clarkii</Emphasis>): a mesocosm study

Temporary ponds, acknowledged for their conservation value, are colonized by the invasive crayfish Procambarus clarkii. We have tested the consequences of this colonization for the ecosystem under two contrasted scenarios: one single individual arrival or three individuals arrival. We recreated the temporary pond ecosystem in 1 m² tanks to investigate the impact of the two crayfish densities. We studied the macrophyte community composition and abundance, chlorophyll a and total suspended solids concentrations, and the diversity and functional composition of micro-crustacean and macro-invertebrate communities. We observed a reduction of macrophyte biomass in experimental crayfish mesocosms in comparison with control tanks, nearly 80 and 40% less in 3 and 1 crayfish/m² tanks, respectively. The macrophyte community shifted, followed by a filamentous algae development, an increase of bare sediment and turbidity in crayfish tanks. The macro-invertebrate community suffered a richness loss of 28 and 22%, in 3 and 1 crayfish/m² tanks, respectively. Functionally, macro-invertebrate diversity reduction most strongly affected the grazer, detritivore and predator trophic groups. Microcrustaceans seemed not to be affected by the introduction of the crayfish. The introduction of the crayfish greatly altered the ecosystem structure and subsequently the ecosystem functioning.     

Cattle waste reduces plant diversity in vernal pool mesocosms

In California, much of the remaining vernal pool habitat is used for cattle grazing. Some studies suggest that grazing helps promote native plant diversity on grasslands, but the impact of grazing on plants that reside in pool basins is largely unknown. We investigated how one aspect of cattle grazing, the deposition of waste, affects these plant species by adding dung and urine to mesocosms lined with vernal pool soil. As a result of dung input, orthophosphate, conductivity, and turbidity increased in our mesocosms while dissolved oxygen decreased. Such changes in water quality are consistent with a shift toward a eutrophic state. Algal biomass and percent-cover also increased in dung-treated mesocosms. When the mesocosms dried, vascular plant species richness and percent-cover in dung-treated mesocosms were reduced by up to 54% and 87%, respectively. We attribute this to light attenuation by algal mats that flourished in the nutrient-enriched water. We also found that dung input caused significant, but weak, shifts in the composition of the vascular plant community. We conclude that cattle grazing may be detrimental to plant communities in vernal pools via increased nutrient loading, which promotes algal growth. Any beneficial effects of grazing may thus be limited to the surrounding grassland. Studies that examine the regional-scale impacts of grazing on vernal pool grasslands should separately consider the impacts to local-scale (i.e., within-pool) plant diversity, as most of the threatened and endangered plant species of California vernal pools reside primarily in pool basins.     

Effects of different vegetative substrates on algal composition in vernal mesocosms

We investigated the response of an algal assemblage to different vegetative substrates in controlled vernal mesocosms. Litter was collected from four vascular plant communities (Deciduous Forest, Macrophyte, Old Field and Pinus stands) and the litter was used to line the benthos of the vernal mesocosms. The development and response of the algal assemblage in treatment and control mesocosms was tracked biweekly for a period of 56 days. A repeated-measure MANOVA and Bonferroni (Dunn) post-hoc test indicated that the Pinus treatment produced a significantly greater biomass than all other treatments. The Pinus treatment mesocosms had acidic (4.5–4.7) water when sampled on day 42, which continued until the completion of the study. The greatest levels of species richness and diversity were recorded from the Deciduous Forest and Old Field treatments. Algal assemblage analyses indicated that there was much overlap in community structure between various litter treatments and algae alone (no litter). Most of the vernal mesocosm treatments were dominated by the filamentous chlorophytes Mougeotia, Oedogonium and Ulothrix. These results suggest that, in this study, the vegetative litter of vernal mesocosms (with the exception of Pinus) exhibits limited independent influence on the developmental trajectories of algal communities.     

Experimental test of the ecosystem impacts of the keystone predator Triops cancriformis (Branchiopoda: Notostraca) in temporary ponds

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Effect of Vallisneria americana (L.) on community structure and ecosystem function in lake mesocosms

Submerged aquatic vegetation is known as a key structural component and regulator in ecosystems. In this mesocosm study, we examine community- and system-level responses to the presence of Vallisneria americana (L), a deep-rooted macrophyte. Phytoplankton, bacteria and filamentous algal biomasses were significantly lowered in the presence of V. americana. In addition, mesocosms with macrophytes had significantly reduced porewater phosphate and iron, water column dissolved organic carbon and total suspended solids, but elevated sediment redox. All mesocosms were net autotrophic (gross primary production/respiration >1). Compared to the macrophyte treatments, the control mesocosms had lower diel net primary production (NPP) midway through the experiment (d 16), but at the end of the experiment (d 36), the controls had the higher values, presumably due to increased filamentous algae. NPP and NPP/R were constant in the macrophyte treatments, whereas NPP/R increased significantly from middle to end of the experiment in the controls. We show that community and system-level responses to the presence of V. americana have significant consequences on system structure and function.     

Multi‐faceted impacts of native and invasive alien decapod species on freshwater biodiversity and ecosystem functioning

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Microbial community diversity and composition varies with habitat characteristics and biofilm function in macrophyte‐rich streams

Biofilms in streams play an integral role in ecosystem processes and function yet few studies have investigated the broad diversity of these complex prokaryotic and eukaryotic microbial communities. Physical habitat characteristics can affect the composition and abundance of microorganisms in these biofilms by creating microhabitats. Here we describe the prokaryotic and eukaryotic microbial diversity of biofilms in sand and macrophyte habitats (i.e. epipsammon and epiphyton, respectively) in five macrophyte‐rich streams in Jutland, Denmark. The macrophyte species varied in growth morphology, C:N stoichiometry, and preferred stream habitat, providing a range in environmental conditions for the epiphyton. Among all habitats and streams, the prokaryotic communities were dominated by common phyla, including Alphaproteobacteria, Bacteriodetes, and Gammaproteobacteria, while the eukaryotic communities were dominated by Stramenopiles (i.e. diatoms). For both the prokaryotes and eukaryotes, the epipsammon were consistently the most diverse communities and the epiphytic communities were generally similar among the four macrophyte species. However, the communities on the least complex macrophyte, Sparganium emersum, had the lowest richness and evenness and fewest unique OTUs, whereas the macrophyte with the most morphological complexity, Callitriche spp., had the highest number of unique OTUs. In general, the microbial taxa were ubiquitously distributed across the relatively homogeneous Danish landscape as determined by measuring the similarity among communities (i.e. Sørensen similarity index). Furthermore, we found significant correlations between microbial diversity (i.e. Chao1 rarefied richness and Pielou's evenness) and biofilm structure and function (i.e. C:N ratio and ammonium uptake efficiency, respectively); communities with higher richness and evenness had higher C:N ratios and lower uptake efficiency. In addition to describing the prokaryotic and eukaryotic community composition in stream biofilms, our study indicates that 1) physical habitat characteristics influence microbial diversity and 2) the variation in microbial diversity may dictate the structural and functional characteristics of stream biofilm communities.     

Persistence of a rare aquatic species along gradients of disturbance and sediment richness

Abstract. This study examines the capacity of establishment of a rare aquatic macrophyte, Luronium natans, within plant communities and habitat types in which it does not occur spontaneously. The species, generally limited to disturbed or nutrient‐poor habitats, was transplanted into a series of sites situated along natural gradients of disturbance (flush‐floods and intermittent sediment exposure) and sediment nutrient‐richness. The transplanted colonies were given a competition‐free establishment period. Colony dynamics of Luronium as well as size structure of the recolonizing macrophyte communities were monitored over three growing seasons. At the end of this period, transplanted colonies still persisted in five out of 12 transplantation sites. Apparently successful integration into the community occurred at both ends of the nutrient gradient, in periodically disturbed habitats. At intermediate to high nutrient richness Luronium maintained one of the highest cover values within the recolonizing community. The study supports previous presumptions that long‐term persistence of Luronium depends on processes limiting community biomass through occurrence of disturbance. But it also reveals the existence of suitable, yet not occupied habitats in which the species can persist as successfully as species from the local species pool. This finding modulates the presumption that Luronium's rarity is mainly caused by a weak competitive ability in the established phase. It thus rises questions about the species’ performance at other stages of its life cycle and on its dispersability.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

Sedimentary context controls the influence of ecosystem engineering by bioturbators on microbial processes in river sediments

By modifying the physical environment, ecosystem engineers can have inordinately large effects on surrounding communities and ecosystem functioning. However, the significance of engineering in ecosystems greatly depends on the physical characteristics of the engineered habitats. Mechanisms underlying such context‐dependent impact of engineers remain poorly understood even though they are crucial to establish general predictions concerning the contribution of engineers to ecosystem structure and function. The present study aimed to decrypt such mechanisms by determining how the environmental context modulates the effects of ecosystem engineers (bioturbators) on microorganisms in river sediments. To test the effects of environmental context on the role of bioturbators in sediments, we used mesocosms and recreated two sedimentary contexts in the laboratory by adding a layer of either fine or coarse sand at the top of a gravel‐sand matrix. For each sediment context, we examined how the sediment reworking activity of a bioturbating tubificid worm (Tubifex tubifex) generated changes in the physical (sediment structure and permeability) and abiotic environments (hydraulic discharge, water chemistry) of microorganisms. Microbial characteristics (abundances, activities) and leaf litter decomposition – a major microbially‐mediated ecological process – were measured to evaluate the impact of bioturbation on biotic compartment. Our results showed that the permeability, the availability of oxygen and the activities of microorganisms were reduced in sediments covered with fine sand, in comparison with sediments covered with coarse sand. Tubifex tubifex significantly increased permeability (by about six‐fold), restored aerobic conditions and ultimately stimulated microbial communities (resulting in a 30% increase in leaf litter breakdown rate) in sediments covered with fine sand. In contrast T. tubifex had low effects in sediments topped by coarse sand, where O₂ was already available for hyporheic microorganisms. Our study supports the idea that context dependency mainly modulates the effects of engineering by controlling the ability of engineers to create changes on abiotic (O₂ in the present study) factors that are limiting for surrounding communities.     

Impact of resuspended sediment on zooplankton feeding in Lake Waihola, New Zealand

1. Wind‐induced sediment resuspension in shallow lakes affects many physical and biological processes, including food gathering by zooplankton. The effects of suspended sediment on clearance rate were determined for a dominant cladoceran, Daphnia carinata, and calanoid copepod, Boeckella hamata, in Lake Waihola, New Zealand. 2. Animals were incubated at multiple densities for 4 days in lake water containing different amounts of suspended lake sediment. Rates of harvest of major food organisms were determined for each sediment level (turbidity) from changes in net growth rate with grazer density. 3. Daphnia cleared all food organisms 7–40 μm in length at similar rates, but was less efficient in its removal of free bacteria, phytoplankton <7 μm, and large cyanobacterial filaments. Elevation of sediment turbidity from 2 to 10 nephelometric turbidity units (NTU) (63 mg DW L^?1 added sediment) reduced Daphnia clearance of phytoplankton, heterotrophic flagellates and ciliates by 72–100%, and of amoebae and attached bacteria by 21–44%. Further inhibition occurred at higher turbidity. 4. Boeckella hamata removed microzooplankton primarily, rather than phytoplankton. The rate at which it cleared rotifers was reduced by 56% when turbidity was increased from 2.5 to 100 NTU. 5. In the absence of macrozooplankton, algal growth increased with sediment turbidity, suggesting that sediment also inhibits rotifer grazing. 6. As mid‐day turbidity in Lake Waihola is ≥10 NTU about 40% of the time, sediment resuspension may play a major role in moderating energy flow and structuring pelagic communities in this lake.     

Livestock grazing affects vernal pool specialists more than habitat generalists in montane vernal pools

Questions

Do livestock grazing and seasonal precipitation structure species composition in montane vernal pools? Which grazing and precipitation variables best predict cover of vernal pool specialists and species with broader habitat requirements? Is vernal pool species diversity correlated with livestock exclosure, and at what spatial scales?

Location

Montane vernal pools, northeast California, USA.

Methods

Vegetation was sampled in 20 vernal pools, including pools where livestock had been excluded for up to 20 years We compared plant species composition, functional group composition and species diversity among sites that varied in grazing history and seasonal precipitation using CCA and LMM.

Results

Although vernal pool specialists were dominant in montane vernal pools, over a third of plant cover was comprised of species that occur over a broad range of wetland or upland environments. The species composition of vernal pool plant communities was influenced by both livestock grazing and precipitation patterns, however the relative effects of these environmental variables differed by functional group. Livestock exclosures favoured perennial vernal pool specialists over annual vernal pool specialists. In contrast, the cover of habitat generalists was more strongly influenced by seasonal precipitation than livestock grazing. At small spatial scales, species richness and diversity decreased as the number of years a pool had been fenced increased, but this relationship was not significant at a larger spatial scale.

Conclusions

Both livestock grazing and seasonal precipitation structure the montane vernal pool plant community. We found that livestock grazing promotes the cover of annual vernal pool specialists, but at the expense of perennial vernal pool specialists. Wetter vernal pools, however, support higher cover of wetland generalist species regardless of whether pools are grazed.     

Fish induced macrophyte loss in shallow lakes: top–down and bottom–up processes in mesocosm experiments

SUMMARY 1. Macrophyte loss from Sites of Special Scientific Interest in England has become widespread over the last 20 years. One reason for this may be changing trends in angling, a multimillion pound industry that has an enormous impact on aquatic ecosystems. Stocking with cyprinid fish is a common angling management practice but the particular fish species and distribution of their biomass may be crucial to the ecosystem. 2. Carp (Cyprinus carpio), roach (Rutilus rutilus), bream (Abramis brama) and tench (Tinca tinca) at biomasses ranging from 0 to 800 kg ha^?1 and at various sizes were placed into experimental mesocosms in Little Mere, a shallow, fertile lake in Cheshire, U.K. The effects these treatments had on the aquatic ecosystem were studied over two summers. Specifically the effects of the treatments on macrophyte growth, benthic and macrophytic macro‐invertebrate populations, water chemistry, epiphyton production and plankton survival were investigated. 3. Carp had a greater detrimental effect on the macrophytes than bream, tench and in particular roach. A biomass of fish > 200 kg ha^?1 adversely affected the extent of macrophyte growth. 4. The decline in macrophyte growth was most likely as a result of increased epiphyton growth that probably reduced the amount of light and carbon dioxide available to the plant. There were no observed direct fish impacts on macrophytes. 5. The chemical data suggested that inorganic nitrogen levels were low and it is possible that release of nitrogen, from fish excreta, followed by immediate uptake, could have been a major factor stimulating epiphyton growth and subsequently macrophyte loss. Phosphorus concentrations increased even in the controls and substantial amounts were available. Phosphorus stimulation can therefore be discounted. Macrophyte‐associated macro‐invertebrates were positively correlated with epiphyton load but had no impact on the extent of epiphytic growth. Shading from disturbed sediment or phytoplankton was also unimportant.     

Long‐term allelopathic control of phytoplankton by the submerged macrophyte Elodea nuttallii

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Does ambient substrate composition influence consumer diversity effects on algal removal?

Benthic substrates constitute an important habitat template for aquatic communities and may affect the contributions of benthic organisms to ecological processes. To test the effects of ambient substrate composition on the process of algae accrual and removal, we conducted an experiment to examine how substrate type influenced consumer richness effects. We hypothesized that algal removal from focal substrates (ceramic tiles) would be influenced by the surrounding ambient substrate through its effect on nutrient cycling and subsequent algal growth. We manipulated consumer richness in mesocosms at one or three species while holding consumer biomass constant. Aquatic consumers were an amphipod, a snail, and a water boatman, and ambient substrates were either sand or gravel. After 21 days, ambient substrate influenced epilithic algal accrual on tiles, affected physio-chemical parameters within mesocosms, and modified consumer behavior. Chlorophyll a was approximately 2× greater on control tiles surrounded by sand, and FPOM and turbidity were greater on sand than gravel when consumers were present. Substrate modified consumer behavior such that consumers congregated around focal substrates in sand, but dispersed around them in gravel. Consumers also had substrate-specific influences on epilithic chlorophyll, causing a decrease in sand and an increase in gravel. Algal assemblages on focal tiles were dominated by diatoms, and their composition responded to consumer richness and identity, but not substrate. Our data suggest that direct effects (e.g., consumptive removal of epilithon from focal tiles) were more pronounced in sand, whereas indirect effects (e.g., bioturbation and enhanced mixing) promoted algal accrual in gravel. These results show that algae production on exposed surfaces may change as underlying substrate composition changes, and that substrate type can alter consumer diversity effects on algal removal.     

A Phytoplankton Bloom in Shallow Divor Reservoir (Portugal)—The Importance of Internal Nutrient Loading

The development of a heavy phytoplankton bloom (chl. a = 360 mg/m³), which occurred in the summer 1983 in a shallow reservoir, Divor, is described. The study shows that remobilization of phosphate from the sediment was initiating the phytoplankton bloom. This was confirmed not only by calculations of the change in iron-phosphate pool, but also supported by sorption experiments carried out with the sediment. It is discussed that turbidity of the water due to suspended matter caused the reduction in standing stock of phytoplankton to approx. 50 mg chl. a/m³ in late summer.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

Macrofaunal community bioturbation along an estuarine gradient

A multidisciplinary study of the impact of bioturbation on sediment dynamics has been underway for some time at the Plymouth Marine Laboratory. The current programme has been founded upon the careful selection of six sites in the River Tamar, representative of important combinations of physical and biological variables. The paper presents preliminary results illustrating the contrasting importance of the speciesNereis diversicolor andNephtys hombergi as agents of bioturbation, given their different distributions across the six sites. Thus bioirrigation byN. diversicolor increases up the estuary and is greatest in the region of high suspended bed-load due to tidal pumping, where the consequences for chemical exchange processes between sediments and the water column may be most important. The sediment mixing effects ofN. hombergi are likely to be greater towards the seaward end of the estuary but ultimately the particular sediment types and the macrofaunal communities they support dictate the level of bioturbation in ways that do not necessarily relate to simple axial gradients along the estuary.     

Ecological influences on larval chironomid communities in shallow lakes: implications for palaeolimnological interpretations

1. To correctly interpret chironomid faunas for palaeoenvironmental reconstruction, it is essential that we improve our understanding of the relative influence of ecosystem variables, biotic as well as physicochemical, on chironomid larvae. To address this, we analysed the surface sediments from 39 shallow lakes (29 Norfolk, U.K., 10 Denmark) for chironomid head capsules, and 70 chironomid taxa (including Chaoborus) were identified. 2. The shallow lakes were selected over large environmental gradients of aquatic macrophytes, total phosphorus (TP) and fish communities. Redundancy analysis (RDA) identified two significant variables that explained chironomid distribution: macrophyte species richness (P < 0.001) and TP (P < 0.005). Generalised linear models (GLM) identified specific taxa that had significant relationships with both these variables. Macrophyte percentage volume infested (PVI) and species richness were significant in classifying the lake types based on chironomid communities under twinspan analysis, although other factors, notably nutrient concentrations and fish communities, were also important, illustrating the complexities of classifying shallow lake ecosystems. Lakes with plant species richness >10 all had relatively diverse (Hill’s N2) chironomid assemblages, and lakes with Hill’s N2 >10 all had TP <250 μg L⁻¹ and total fish densities <2 fish per m². 3. Plant density (PVI), and perhaps more importantly species richness, were primary controls on the distribution of chironomid communities within these lakes. This clearly has implications for palaeoenvironmental reconstructions using zoobenthos remains (i.e. chironomids) and suggests that they could be used to track changes in benthic/pelagic production and could be used as indicators of changing macrophyte habitat. 4. Measuring key biological gradients, in addition to physicochemical gradients, allowed the major controls on chironomid distribution to be assessed more directly, in terms of plant substrate, food availability, competition and predation pressure, rather than implying indirect mechanisms through relationships with nutrients. Many of these variables, notably macrophyte abundance and species richness, are not routinely measured in such studies, despite their importance in determining zoobenthos in temperate shallow lakes. 5. When physical, chemical and ecological gradients are considered, as is often the case with palaeo‐reconstructions rather than training sets chosen to maximise one gradient, complex relationships exist, and attempting to reconstruct a single trophic variable quantitatively may not be appropriate or reliable.