Life-history strategies in freshwater macroinvertebrates

• 1 Explaining spatial and temporal differences in species assemblages is a central aim of ecology. It requires a sound understanding of the causal mechanisms underlying the relationship of species with their environment. A species trait is widely acknowledged to be the key that links pattern and process, although the enormous variety of traits hampers generalization about which combination of traits are adaptive in a particular environment.
• 2 In three steps, we used species traits to match species and environment, and chose lentic freshwater ecosystems to illustrate our approach. We first identified key environmental factors and selected the species traits that enable the organism to deal with them. Secondly, we investigated how investments in these traits are related (e.g. through trade‐offs). Thirdly, we outlined 13 life‐history strategies, based on biological species traits, their interrelations known from life‐history theory and their functional implications.
• 3 Species traits and environmental conditions are connected through life‐history strategies, with different strategies representing different solutions to particular ecological problems. In addition, strategies may present an integrated response to the environment as they are based on many different traits and their interrelationships. The presence and abundance of (species exhibiting) different life‐history strategies in a location may therefore give direct information about how a particular environment is experienced by the species present.
• 4 Life‐history strategies can be used to (i) explain differences in species assemblages either between locations or in different periods; (ii) compare waterbodies separated by large geographical distances, which may comprise different regional species pools or span species distribution areas and (iii) reduce often very complex, biodiverse assemblages into a few meaningful, easily interpretable relationships.

     

Biological and ecological traits of benthic freshwater macroinvertebrates: relationships and definition of groups with similar traits

• 1 Relating species traits to habitat characteristics can provide important insights into the structure and functioning of stream communities. However, trade‐offs among species traits make it difficult to predict accurately the functional diversity of freshwater communities. Many authors have pointed to the value of working with groups of organisms as similar as possible in terms of relationships among traits and have called for definition of groups of organisms with similar suites of attributes.
• 2 We used multivariate analyses to examine separately the relationships among 11 biological traits and among 11 ecological traits of 472 benthic macroinvertebrate taxa (mainly genera). The main objective was to demonstrate (1) potential trade‐offs among traits; (2) the importance of the different traits to separate systematic units or functional groupings; and (3) uniform functional groups of taxa that should allow a more effective use of macroinvertebrate biological and ecological traits.
• 3 We defined eight groups and 15 subgroups according to a biological trait ordination which highlighted size (large to small), reproductive traits (K to r strategists), food (animal to plant material) and feeding habits (predator to scraper and/or deposit feeder) as ‘significant’ factors determining the ordination of taxa. This ordination partly preserved phylogenetic relationships among groups.
• 4 Seven ecological groups and 13 ecological subgroups included organisms with combinations of traits which should be successively more adequate in habitats from the main channel to temporary waters, and from the crenon to the potamic sections of rivers, and to systems situated outside the river floodplain. These gradients corresponded to a gradual shift from (1) rheophilic organisms that lived in the main channel of cold oligotrophic mountain streams to (2) animals that preferred eutrophic habitats of still or temporary waters in lowlands. The groups with similar ecological traits had a more diverse systematic structure than those with similar biological traits.
• 5 Monitoring and assessment tools for the management of water resources are generally more effective if they are based on a clear understanding of the mechanisms that lead to the presence or absence of species groups in the environment. We believe that groups with similar relationships among their species traits may be useful in developing tools that measure the functional diversity of communities.

     

Contrasting responses of oligochaete communities to the abatement of eutrophication in Lake Geneva

Tubificid and lumbriculid worms were used to monitor, at depths of 150 m, the recovery of Lake Geneva (Switzerland) from eutrophication. As predicted from the decrease of phosphorus concentrations, relative abundance of oligotrophic species was higher from 1988 to 1993 than in 1983, i.e. before the abatement of eutrophication. However, this trend towards oligotrophication can be reversed, as indicated by a decrease of oligotrophic species, recorded in 1993. But this change corresponded to the effects of an increase of water temperature on the abundance of the mesotrophic species Potamothrix vejdovskyi rather than to a deterioration of the profundal. In addition to this short-term setback, oligochaete communities located at a depth of 150 m responded more slowly and less clearly to the decrease of phosphorus concentrations than those located at a depth of 40 m. However, the zoobenthos indicated more clearly the recovery of Lake Geneva than the phytoplankton.     

Species traits for future biomonitoring across ecoregions: patterns along a human-impacted river

1. Current budgets for environmental management are high, tend to increase, and are used to support policy and legislation which is standardized for large geographic units. Therefore, the search for tools to monitor the effects of this investment is a major issue in applied ecology. Ideally, such a biomonitoring tool should: (1) be as general as possible with respect to its geographic application; (2) be as specific as possible by separating different types of human impact on a given ecosystem; (3) reliably indicate changes in human impact of a particular type; and (4) be derived from a sound theoretical concept in ecology. 2. We developed an approach to biomonitoring which matches these ‘ideal’ characteristics by focusing on numerous, general biological species traits (e.g. size, number of descendants per reproductive cycle, parental care, mobility) and on the habitat templet concept, which relates trends in these general species traits to disturbance patterns. Using the French Rhône River and benthic macroinvertebrates as an example, we have used the data to demonstrate a general framework and the potential of our approach rather than to produce a ready-made tool. Our data covered a large river and its major tributaries, which has a catchment that crosses ecoregions, and known gradients and discontinuities in human impact. 3. We applied multivariate analyses to evaluate how the distribution of species traits in invertebrate communities could discriminate environmental differences along the Rhône in comparison to traditionally used approaches (e.g. community structure, based on species abundances, or ecological species traits, such as velocity preferences and pollution tolerance). Invertebrate community structure expressed in terms either of the abundance or the traits of species reliably indicated differences in overall human impact. The community structure based on biological traits was less confounded by natural spatial gradients and reliably indicated human impact, while community structure based on ecological traits was the most confounded by natural spatial gradients and was the poorest indicator of human impact. Community structure based on species abundances was an intermediate indicator of human impact. 4. These results indicate that a revision of biomonitoring approaches which have been based on a single aspect of the biological responses may be warranted. The biological traits of species could separate the different types of human impact. Therefore, the use of these traits in biomonitoring could improve existing multi-metric approaches. Future research has to show if the general applicability of species traits allows the development of a unique biomonitoring tool for running waters of the European Union, for running waters in temperate climates on several continents, for freshwater, marine and terrestrial systems, and/or for global biodiversity assessment.     

Development of a phytoplankton indicator system for the ecological assessment of brackish coastal waters (German Baltic Sea coast)

The implementation of the European Water Framework Directive (WFD) requires the development of ecologically-based classification systems for anthropogenically-induced eutrophication in all types of water bodies. Due to the inherent high temporal and spatial variability of hydrological and geochemical parameters of the coastal waters of the southern Baltic Sea, discrimination between anthropogenic impact and natural variability is necessary. The development of statistical methods for this discrimination was the main aim of this study. These methods were used to derive indicative phytoplankton parameters for different stages of eutrophication for the investigation area. For this purpose, a long-term phytoplankton data series was analysed, which covered a broad salinity and eutrophication gradient. In order to detect eutrophication effects, the analysis was restricted to phytoplankton spring bloom events and to the salinity range between 5 and 10 psu, i.e. superimposing seasonal and hydrodynamic effects were eliminated. An artificial abiotic degradation vector was developed based on four typical water quality parameters. A total of 11 potentially indicative phytoplankton parameters on different taxonomical levels arose from a correlation analysis with this degradation vector. These indicators were then tested for their ability to discriminate between three eutrophication levels. Finally, seven phytoplankton indices could be proposed: total phytoplankton biovolume, the percentage of diatoms and the biovolume of different size ranges of diatoms and one indicative species (Woronichinia compacta). Guest editors: A. Razinkovas, Z. R. Gasiūnaitė, J. M. Zaldivar & P. Viaroli European Lagoons and their Watersheds: Function and Biodiversity     

The contribution of induction of temperate phages to the numbers of free somatic coliphages in waters is not significant

Somatic coliphages have been proposed as indicators of water quality. But several factors have been considered a drawback for their use as indicators. We evaluated the contribution of temperate phages to the numbers of somatic coliphages detected in water by ISO (International Standards Organization) standardised methods. Prophage induction from naturally occurring bacteria was assayed with mitomycin C, ciprofloxacin and UV irradiation. Results indicate that the presence of prophages will not influence the determinations of somatic coliphages in water.     

The macrofauna distribution in brackish inland waters in relation to chlorinity and other factors

The macrofauna of small isolated brackish waters on a barrier island in the Dutch Wadden Sea was sampled qualitatively. An attempt was made to correlate species composition and species richness to chlorinity characteristics and to data on the water regime of the localities studied.It is shown that the observed differences in the macrofauna cannot be explained by differences in chlorinity only. Present and former connections to other water bodies seem equally important.     

Elaborate ornaments are costly to maintain: evidence for high maintenance handicaps

Elaborate secondary sexual traits, such as the ornamental plumageof birds, may be favored by female choice because they serveas honest indicators of male quality. Elaborate traits are thoughtto be honest signals because they are expensive to produce andincrease predation risk. Here we investigate another potentialcost of elaborate traits, i.e., the time and energy requiredto maintain them in good condition. We tested the hypothesisthat species of birds with ornamental plumage invest more timein maintenance behavior than do related species without suchplumage. To test the hypothesis we quantified the maintenancebehavior of nine ornamental and nine non-ornamental speciesin aviaries and zoos. To test the validity of using captivebirds, we first collected data on 12 captive species for whichdata from wild individuals were also available. The maintenancetimes of captive and wild individuals were highly correlatedacross species. Maintenance time was also correlated with plumagelength, independent of body size. Ornamental species had longerplumage than non-ornamental species, and they devoted significantlymore time to maintenance. Time spent on maintenance cannot bedevoted to other activities. This temporal trade-off reinforcesthe honesty of ornamental plumage. We suggest that high maintenancehandicaps are present in a variety of animals.     

Tuning in to multiple predators: conflicting demands for shell morphology in a freshwater snail

1. We examined the response to chemical cues from fish and crayfish, two predators with contrasting feeding modes, and their single and combined effect on shell morphology in the freshwater snail Radix balthica. 2. Snails were subjected to four treatments: tench (Tinca tinca), signal crayfish (Pacifastacus leniusculus), a combination of tench and signal crayfish and no predators (control). Shell shape, crushing resistance and shell thickness were quantified. We also analysed whether shape or shell thickness contributes most to crushing resistance. 3. Chemical cues from the fish induced a rounder shell shape in R. balthica, a thicker shell and a higher crushing resistance, whereas crayfish chemical cues had no effect on shell morphology, shell thickness or crushing resistance. Shell shape contributed more to crushing resistance than shell thickness. 4. The combined predator treatment showed an intermediate response between the fish and crayfish treatments. Shell roundness was reduced compared with the fish treatment, but the reduced crushing resistance that comes with a less rounded shell was compensated by an increased investment in extra shell material, exceeding that of the fish treatment. 5. Our study extends previous studies of multipredator effects on phenotypically plastic freshwater snails by showing that the snails are able to fine‐tune different elements of morphology to counter predator‐specific foraging modes.     

A living environment. Objectives and measures in the Swedish Government Bill on environmental policy 1990/91, pertaining to waters and the seas

Increased international cooperation to limit transboundary and global environmental problems, increased sectoral responsibility and decentralization, and improved information and statistics on the state of the environment are the main lines for the development of future environmental policy. For lakes, watercourses, and the seas the overall objective is to maintain viable, balanced populations of naturally occurring species. Pollution is not to limit the use of water for fishing, recreation, or water supply. Foreign species or organisms subjected to genetic mutation are only to be introduced with great restriction. Measures are suggested that will reduce environmental disturbances from agriculture and forestry, traffic, energy production, industry, goods, chemical products, and waste.Paper presented at Seminar on Ecosystems Approach to To Water Management (Oslo, Norway, 27–31 May 1991).     

Solving the problem of how to eat something as big as yourself: Diverse bacterial strategies for degrading polysaccharides

Polysaccharide digestion by bacteria is an important activity in many ecosystems, and a number of bacterial genera can perform this function. Although many papers have been published about the properties of isolated polysaccharide-degrading enzymes, relatively little is known about how intact bacteria degrade polysaccharides. This review summarizes recent findings suggesting that there are at least three different strategies. the most familiar one is the excretion of extracellular polysaccharidases, which diffuse to and degrade nearby polysaccharides. An example of this type of strategy is provided by the plant pathogen,Erwinia spp. A second strategy is to have the enzyme exposed to the extracellular medium but attached to the surface of the cell. Examples of this strategy are provided by the pullulanase system ofKlebsiella oxytoca and the cellulosomes ofClostridium thermocellum. A strategy that could be seen as a combination of the extracellular enzyme strategy and the surface organelle strategy is provided byVibrio harveyi, which attaches to its substrate, chitin, via proteins that appear to be specialized for attachment and produces extracellular enzymes that attack the chitin. A third strategy is to import the polysaccharide, as appears to be done byBacteroides spp. In this instance, the polysaccharide is bound to an outer membrane receptor, then passes into the periplasm where the degradative enzymes are located. The ecological advantages and disadvantages of these systems are discussed, and areas where further research is needed are defined.     

Leaf and shoot water content and leaf dry matter content of Mediterranean woody species with different post-fire regenerative strategies

BACKGROUND AND AIMS: Post-fire regeneration is a key process in Mediterranean shrubland dynamics, strongly determining the functional properties of the community. In this study, a test is carried out to determine whether there is co-variation between species regenerative types and functional attributes related to water use. METHODS: An analysis was made of the seasonal variations in leaf relative water content (RWC), leaf dry matter content (LDMC), leaf moisture (LM) and live fine fuel moisture (LFFM) in 30 woody species of a coastal shrubland, with different post-fire regenerative strategies (seeding, resprouting or both). KEY RESULTS: RWC results suggest that the studied resprouters have more efficient mechanisms to reduce water losses and maintain water supply between seasons. In contrast, seeders are more drought tolerant. LDMC is higher in resprouters over the course of the year, suggesting a more efficient conservation of nutrients. The weight of the phylogenetic constraint to understand differences between regenerative strategies tends to be important for LDMC, while it is not the case for variables such as RWC. CONCLUSIONS: Groups of species with different post-fire regenerative strategies (seeders and resprouters) have different functional traits related to water use. In addition to the role of phylogenetical constraints, these differences are also likely to be related to the respective life history characteristics. Therefore, the presence and abundance of species with different post-fire regenerative responses influence the functional properties of the communities.     

Diatom and micro-invertebrate communities and environmental determinants in the western Australian wheatbelt: a response to salinization

Diatom and micro-invertebrate communities were analyzed in 56 lentic wetlands from the wheatbelt of Western Australia. Sixteen water quality parameters were measured at each habitat and were tested as determinants of diatom and micro-invertebrate distribution. Nearly all waters were dominated by Na¹⁺ and Cl^1– and over half had low buffering capacity (<2 meq/L as TCO). Salinities ranged from <1 to > 240 mS/cm; some hypersaline habitats had pH values as low as 2.1, but most sites were alkaline.Diatoms (165 taxa) were present in all habitats, with indications that assemblages are correlated with land-use practices, i.e., historical clearing of native vegetation for dryland agriculture with resultant hydrological problems of salinization and water-logging. A non-metric multidimensional ordination showed pH, salinity, TCO, and total phosphorus to be significantly correlated with diatom community structure. Principal components analysis showed similar patterns. A salinity index was also developed for numerically important diatom taxa. Principal components analysis showed similar patterns. Certain tolerant Bacillariales, including Hantzschia amphioxys, Nitzschia hybrida, Tryblionella hungarica, and N.recta, were numerically important taxa in wetlands with visible signs of salinization.Micro-invertebrates (143 taxa) were present in habitats with salinities up to 200 mS/cm. Multidimensional ordinations showed salinity to be significantly correlated with micro-invertebrate community structure. The cladoceran Daphniopsis queenslandensis, ostracods Australocypris insularis, Mytilocypris tasmanica chapmani, and Platycypris baueri, and copepods Apocyclops dengizicus and Mesochra nr. flava were numerically important micro-invertebrates in moderate to high salinity environments, including those with signs of salinization. The diatoms, Navicula minuscula var. muralis and Pinnularia divergentissa var.subrostrata, and the copepod, Calamoecia trilobata, were common inhabitants of the extreme hypersaline (75 mS/cm), acidic (<4) habitats. Diatom communities were similar to those previously reported for other regions in Australia subjected to salinization; there were some differences in micro-invertebrate communities.     

A generic aggregation approach to account for statistical uncertainty when combining multiple assessment results

The one-out-all-out approach (OOAO) for aggregating the assessments of single elements (e.g. species or ecosystem components) has found application in environmental polices such as the European Water Framework Directive or the Marine Strategy Framework Directive (MSFD). However, the OOAO has been challenged as being too pessimistic by making positive assessment results virtually impossible along the increasing number of aggregated elements. This study presents a generic approach, the probabilistic One-out-all-out approach, (pOOAO), to account for this issue and thereby reconciling the OOAO with probabilistic aggregation methods The pOOAO allows to determine the minimum number of elements (K_GES), which should meet their assessment benchmarks and thus should achieve a good status. By pre-setting a generic confidence level for each single assessment (e.g. 0.95) the binomial distribution can be used to obtain K_GES for any number of assessed elements. The pOOAO can also accommodate for the integration of assessments from multiple indicators within an element by adjusting the confidence level in relation to the number of integrated indicators. Depending on the generic confidence level as well as on the number of and integrated indicators and aggregated elements, the pOOAO is either consistent with the OOAO or allows for a certain number of negative assessment results, which are attributed to statistical uncertainty and error propagation. The pOOAO is consistent with the OOAO if the desired confidence level in the single assessment results is very high (>0.99) and/or the number of aggregated elements and integrated indicators is low. Through this flexibility the pOOAO could find wide application within integrated ecosystem assessments frameworks such as the MSFD, but would require to estimate the confidence level for each single assessment.     

The development of policy approaches for reducing nitrogen pollution to coastal waters of the USA

Two-thirds of the coastal rivers and bays in the United States are degraded from nutrient pollution, and nitrogen inputs these waters continue to increase. The nitrogen comes from a variety of sources, including runoff from agricultural fields, concentrated animal feeding operations, atmospheric deposition from fossil fuel combustion, and sewage and septic wastes.Technical solutions for nitrogen pollution exist at reasonable cost. That most of these solutions have not yet been implemented to any significant extent across the United States suggests that new policy approaches are necessary. The best solution may involve a combination of voluntary and mandatory approaches, applying different approaches to different sources of nitrogen poilution. A watershed-based approach that relies heavily on voluntary mechanisms (such as crop-yield insurance to reduce over-fertilization) is likely to be the most effective for some sources of nitrogen (such as runoff from agricultural fields), while a uniform national regulatory approach may be better for others (such as NOx emissions from fossil fuel combustion). Implementation of management strategies should be carefully coupled to monitoring programs to assess the effectiveness of these strategies. While both nitrogen and phosphorus are important to control, the focus should be on nitrogen management, in part because nitrogen is more generally the causal agent of coastal eutrophication. Also, while nitrogen-control practices tend to also reduce phosphorus pollution, phosphorus-control practices often have little effect on nitrogen.Although current scientific and technical knowledge is sufficient to begin to make substantial progress toward solving coastal nitrogen pollution, progress will be made more quickly and more cost effectively with increased investment in appropriate scientific research.     

The development of policy approaches for reducing nitrogen pollution to coastal waters of the USA

Two-thirds of the coastal rivers and bays in the United States are degraded from nutrient pollution, and nitrogen inputs these waters continue to increase. The nitrogen comes from a variety of sources, including runoff from agricultural fields, concentrated animal feeding operations, atmospheric deposition from fossil fuel combustion, and sewage and septic wastes. Technical solutions for nitrogen pollution exist at reasonable cost. That most of these solutions have not yet been implemented to any significant extent across the United States suggests that new policy approaches are necessary. The best solution may involve a combination of voluntary and mandatory approaches, applying different approaches to different sources of nitrogen pollution. A watershed-based approach that relies heavily on voluntary mechanisms (such as crop-yield insurance to reduce over-fertilization) is likely to be the most effective for some sources of nitrogen (such as runoff from agricultural fields), while a uniform national regulatory approach may be better for others (such as NOx emissions from fossil fuel combustion). Implementation of management strategies should be carefully coupled to monitoring programs to assess the effectiveness of these strategies. While both nitrogen and phosphorus are important to control, the focus should be on nitrogen management, in part because nitrogen is more generally the causal agent of coastal eutrophication. Also, while nitrogen-control practices tend to also reduce phosphorus pollution, phosphorus-control practices often have little effect on nitrogen. Although current scientific and technical knowledge is sufficient to begin to make substantial progress toward solving coastal nitrogen pollution, progress will be made more quickly and more cost effectively with increased investment in appropriate scientific research.     

Land-use influence on stream water quality and diatom communities in Victoria, Australia: a response to secondary salinization

Diatom communities were analyzed in 39 streams located in drainages with varied land-use practices throughout Victoria, Australia. Thirteen water quality parameters were also measured in each stream. Most streams had low HCO₃ ^1- concentrations (low buffering capacity) with >90% of the waters dominated by Na¹⁺ and Cl^1-. Phosphate concentrations ranged from 0.003 to 2.0 mg/L. Diatom communities (245 taxa) were strongly correlated with land-use practices, i.e. historic clear cutting, and secondary salinization. Streams influenced by heavy irrigation practices and dryland farming had reduced species diversity and richness compared to systems with low to moderate land use. A nonmetric multidimensional ordination of diatom communities in the 39 streams was conducted. An ANOSIM on the ordination showed that diatom communities in upland watersheds with native forest canopies and low salinization, lowland streams in watersheds with cleared forest canopies, moderate agricultural utilization and salinization, and lowland streams in areas with high irrigation and salinization were all significantly different (p<0.001) from one another. Community ordination techniques showed that both specific conductance (salinity) and phosphorus interacted to determine stream diatom community structure in drainages with high secondary salinization. Drainages with low to moderate agricultural activity and low nutrients, but with a wide range of salinities showed strong associations with the diatom taxa Amphora coffeaeformis, Cymbella pusilla and Entomoneis paludosa, whereas, streams in regions with heavy agricultural practices and high phosphorus had Bacillaria paradoxa, Nitzschia hungarica, N. frustulum and Aulacoseira granulata as numerically important diatoms. In contrast, Rhizosolenia eriensis, Frustulia rhomboides, Eunotia pectinatus and Tabellaria flocculosa were strongly associated with upland streams with fast current, relatively low O-PO₄ ^3- concentrations, low pH, low salinity, and low temperature. In general, the diatom communities in saline streams (3 mS) were similar to those previously reported in saline lakes in Victoria.