Bioturbation as Driver of Zooplankton Recruitment,Biodiversity and Community Composition in Aquatic Ecosystems

In an experimental study we assessed if benthic bioturbating invertebrates affect the recruitment (hatching) of zooplankton from the sediment, and if this effect persists as differences in the zooplankton community in the water column, that is, if bioturbation quantitatively stimulates benthic–pelagic coupling. We investigated the effects of four different benthic invertebrates (Asellus aquaticus, Chironomus plumosus, Tubifex tubifex in the presence or absence of the predator Sialis lutaria). In total, 45 zooplankton taxa hatched from the sediment and the hatching success of some of these was dependent on the species identity of the bioturbating invertebrate. The predator Sialis reduced the abundance of all three invertebrate species, but tended to positively influence the zooplankton recruitment rates, possibly through increasing the activity of the bioturbating invertebrates. The most striking effect of bioturbation on the hatching and pelagic zooplankton community properties was that, on average, 11% more species hatched in the Asellus treatment than in any other treatment. This was also mirrored in the zooplankton water column community where, on average, 7% more species established a viable population in treatments with Asellus as bioturbator. In a complementary field survey, Asellus was more common in littoral than in profundal sediments. Because Asellus strongly affected recruitment of zooplankton in our experiment, we argue that bioturbation may partly explain why recruitment of resting stages of both phyto- and zooplankton is generally higher in littoral than in profundal areas.     

Bioturbating space enhances the effects of non‐additive interactions among benthic ecosystem engineers on cross‐habitat nutrient regeneration

Understanding the mechanisms that drive complementary interactions among species is key to the progress of the debate over the role of biodiversity in ecosystem functioning. In addition, interspecific interactions among physical ecosystem engineers have rarely been framed in the context of biodiversity experiments. Here, we provide an empirical test of how the physical niche space of species influences the effects of the biodiversity of bioturbators on cross‐habitat nutrient fluxes in benthic sediments. In the laboratory, we orthogonally manipulated the number and composition of three benthic invertebrate bioturbator species that differ in the dimensions of their bioturbating space niche; i.e. their vertical distribution in the sediment over a gradient of sediment depth and volume. The ammonium (NH₄‐N) flux from the sediment to the water was positively related to bioturbator species richness only in the sediments with the deepest depth and greatest volume. The non‐additive effects of bioturbator species richness on the benthic–pelagic NH₄‐N flux increased linearly with sediment depth and volume, but only in the three‐species mixtures. Furthermore, no individual species dominated the rates of H₄‐N fluxes, indicating that biodiversity effects were mainly driven by complementarity. These results suggest that sediment bioturbating space mediates the magnitude of non‐additive effects among the three invertebrate species and sheds light on the importance of physical niche space in modulating the positive effects of biodiversity on ecosystem functioning.     

Bioturbation by a dominant detritivore in a headwater stream: litter excavation and effects on community structure

Bioturbation can affect community structure by influencing resource distribution and habitat heterogeneity. Bioturbation by detritivores in small headwater streams could affect community structure by reintroducing buried detrital resources into the food web and could also affect the distribution of various taxa on detritus. We evaluated the ability of the caddisfly Pycnopsyche gentilis to uncover experimentally buried leaves in a headwater stream. Packs of leaves were placed in enclosures and covered with a known volume of sediment. We added 0, 3 or 6 large Pycnopsyche to the enclosures which were permeable to most other invertebrate taxa. Leaf packs were sampled after 23 days and leaf pack mass, the amount of sediment covering the leaf packs, and macro‐ and microinvertebrate densities on leaf packs were quantified. There was a significant negative relationship between Pycnopsyche density and leaf pack mass. Pycnopsyche also reduced the volume of sediment covering leaf packs. Pycnopsyche had complex effects on the abundance of invertebrate taxa associated with the leaves. Some taxa exhibited their highest abundance in the 3 Pycnopsyche treatment while others exhibited non‐significant increases as Pycnopsyche density increased. These results suggest that the beneficial effects of Pycnopsyche (e.g. uncovering leaves which increases the availability of habitat and food) outweigh any negative effects (e.g. disturbance, encounter competition) of the caddisfly when it is present at lower densities. However, the negative impacts of Pycnopsyche appear to outweigh the positive effects via sediment removal at higher caddisfly densities for some taxa. Our results suggest that bioturbating organisms in streams have the potential to reintroduce organic matter to detrital food webs and affect the distribution and abundance of benthic taxa associated with organic matter.     

Relative influences of submersed macrophytes and bioturbating fauna on biogeochemical processes and microbial activities in freshwater sediments

1. Invertebrates and aquatic plants often play a key role in biogeochemical processes occurring at the water–sediment interface of aquatic ecosystems. However, few studies have investigated the respective influences of plants and bioturbating animals on ecological processes (nutrient fluxes, benthic oxygen uptake, microbial activities) occurring in freshwater sediments. 2. We developed a laboratory experiment in aquaria to quantify the effects of (i) one invertebrate acting as a bioturbator (Tubifex tubifex); (ii) one submersed plant with a high sediment‐oxygenating potential (Myriophyllum spicatum) and (iii) one submersed plant with a low sediment‐oxygenating potential (Elodea canadensis). 3. The tubificid worms significantly increased the fluxes of nitrogen at the water–sediment interface (influx of nitrate, efflux of ammonium), whereas the two plant species did not have significant influences on these nitrogen fluxes. The differences in nitrogen fluxes between tubificid worms and plants were probably due to the bioirrigation process caused by T. tubifex, which increased water exchanges at the water–sediment interface. Tubifex tubifex and M. spicatum produced comparable reductions of nutrient concentrations in pore water and comparable stimulations of benthic oxygen uptake and microbial communities (percentages of active eubacteria and hydrolytic activity) whereas E. canadensis had a very weak influence on these variables. These differences between the two plants were due to their contrasting abilities to increase oxygen in sediments by radial oxygen losses (release of oxygen from roots). 4. Our study suggests that the bioirrigation process and radial oxygen loss are major functional traits affecting biogeochemical functioning at the water–sediment interface of wetlands.     

Microbial Response of a Freshwater Benthic Community to a Simulated Diatom Sedimentation Event: Interactive Effects of Benthic Fauna

Abstract The response of a sediment microbial assemblage to a pulse of diatoms was studied over 36 days by measuring bacterial activity and biomass, ATP concentration, and overall community respiration in laboratory microcosms. Also, the contribution of macrofaunal chironomids to the decomposition of settling diatoms in benthic communities, and the relative importance of benthic meiofauna in community metabolism, were determined. The addition of diatoms resulted in an immediate response by sediment bacteria, with higher bacterial production recorded after only 2 h, and a more than tenfold increase within one day. The rapid response by sediment bacteria was accompanied by relatively high initial concentrations of dissolved organic carbon. In treatments receiving diatoms, higher bacterial production was sustained throughout the experiment. Surprisingly, neither these elevated production estimates, nor the starvation of controls affected bacterial abundance. Mean bacterial cell volume, however, was markedly affected by the addition of diatoms. Combining community respiration measurements and bacterial production estimates showed that growth efficiencies for sediment bacteria ranged from 14.6 to 34.5%. The contribution of ambient meiozoobenthos to carbon metabolism was less than 1%. Carbon budgets showed that 1.3 mg C was cooxidized along with 4.3 mg added diatom C. Sediment reworking by Chironomus larvae initially enhanced bacterial production, but the presence of Chironomus resulted in lower bacterial production estimates after 16 and 36 days. This was interpreted as a result of faster decomposition of diatoms in treatments with chironomids, which was validated by a faster decline of ATP and chlorophyll a in the sediment. Our results indicate that Chironomus larvae compete with sediment bacteria for available organic substrates. Received: 11 June 1996; Accepted: 13 August 1996     

Production and energy flow in the benthic community of a Texas pond

Energy flow (A = R + P; assimilation = respiration + production) and life-history/density-biomass cycles were followed for 1 year in a mayfly (Brachycercus sp.) population and two chironomid (Procladius sp. and Tendipes decorus) populations in a 0.94 ha pond located in north-central Texas. Temperature (5, 10, 15, 20, 25 and 30°C) effects on R and patterns of metabolic compensation/acclimatization were determined for the mayfly and chironomids. Chironomid and mayfly populations dominated the nine benthic macroinvertebrates identified in the pond, with greatest abundances in winter and winter-spring respectively. Annual energy flow was 51 kcal m^–2 (R = 40 kcal, P = 11 kcal or 1.9 g) in Brachycercus sp., 23 kcal m^–2 (R = 5 kcal, P = 18 kcal or 2.4 g) in Procladius sp. and 74 kcal m^–2 (R = 17 kcal, P = 57 kcal or 6.0 g) in T. decorus. Each species metabolically compensated partially to temperatures en countered during their life histories.     

Bioturbation enhances the aerobic respiration of lake sediments in warming lakes

While lakes occupy less than 2% of the total surface of the Earth, they play a substantial role in global biogeochemical cycles. For instance, shallow lakes are important sites of carbon metabolism. Aerobic respiration is one of the important drivers of the carbon metabolism in lakes. In this context, bioturbation impacts of benthic animals (biological reworking of sediment matrix and ventilation of the sediment) on sediment aerobic respiration have previously been underestimated. Biological activity is likely to change over the course of a year due to seasonal changes of water temperatures. This study uses microcosm experiments to investigate how the impact of bioturbation (by Diptera, Chironomidae larvae) on lake sediment respiration changes when temperatures increase. While at 5°C, respiration in sediments with and without chironomids did not differ, at 30°C sediment respiration in microcosms with 2000 chironomids per m² was 4.9 times higher than in uninhabited sediments. Our results indicate that lake water temperature increases could significantly enhance lake sediment respiration, which allows us to better understand seasonal changes in lake respiration and carbon metabolism as well as the potential impacts of global warming.     

Bioturbation as a mechanism for setting and maintaining levels of diversity in subtidal macrobenthic communities

Over 2 years, experiments were conducted tocompare the effects of sediment disturbance by different bioturbating, macrofaunal organisms on the diversity and structure of the associated infaunal community. The four species investigated were the bivalves Nuculoma tenuis (Montagu, 1808) and Abra alba (Wood, 1802), the heart urchin Brissopsis lyrifera (Forbes, 1841), and the burrowing decapod Calocaris macandreae (Bell, 1846). These organisms were chosen to allow assessment of the effects of contrasting feeding activities and body sizes of the bioturbating species on the diversity of the macrobenthic communities. Bioturbation by the sub-surface deposit feeders N. tenuis and B. lyrifera promoted higher levels of and diversity in treatments exposed to intermediate levels of disturbance. Whilst no such intermediate response was demonstrated for A. alba or C. macandreae, it was evident that changes in the associated fauna were influenced by the feeding type of the bioturbating organism responsible. It was also shown that different elements of the associated community responded differently to biotic disturbance. The results indicate that the variability in density and distribution of such bioturbators are important factors in structuring infaunal communities, and in setting and maintaining levels of diversity in apparently homogeneous areas.     

Predation,sediment stability and food availability as determinants of the benthic invertebrate fauna in two shallow lakes

The sedimentary benthos of a series of shallow, eutrophicated lakes, the Norfolk Broads is, in general, low not only in number of species but unexpectedly in number of individuals. In two of the lakes, Hudsons Bay and Hoveton Great Broad, chironomids and oligochaetes dominated the fauna. Hudsons Bay has an extensive stand of water lilies (Nuphar lutea); Hoveton Great Broad does not. There were significant relationships between number of chironomids and of Potamothrix hammoniensis with organic content of the sediments, but these were due not to food availability but to the structure imparted to the otherwise fluid sediment by the organic matter. Sediment stabilised in plastic bowls developed much larger populations of oligochaetes than found in the unrestricted sediment. Protection of the community from fish predation resulted in a further major increase in numbers. Sediment stability and predation rather than food supply were the major determinants of these benthic populations.     

Distribution and production of chironomids (Diptera: Chironomidae) in a shallow, alkaline lake (Neusiedler See, Austria)

From July 1990 to July 1991 the benthic community of the open water zone of Neusiedler See, one of the largest shallow lakes in central Europe, was studied with special reference to the chironomids. Only 16 spp. of chironomids inhabited the sediment of the open water zone. The numerically dominant species were Tanypus punctipennis, Procladius cf. choreus, Microchironomus tener and Cladotanytarsus gr. mancus. Most invertebrates showed a distinct horizontal distribution. Species richness and abundance were highest on muddy and organically rich substrates near the reed belt. Chironomid densities in this area reached 54,000 ind m^–2 and biomass was 2.0 g dw m^–2. The two tanypod species accounted for more than 90% of the standing stock of the macrozoobenthos near the reed belt. The sediment of the open lake and of the eastern part of Neusiedler See was composed of compact clay and sand as a result of the erosion of fine material due to strong waves and currents. Individual densities in these areas were much lower. Production of the numerically dominant species T. punctipennis was estimated using the increment-summation method, whereas production of the remaining species was estimated using an empirically derived multiple regression. Mean annual production of chironomids exceeded 6 g dw m^–2 yr^–1 near the reed belt, but it reached only 0.55 g dw m^–2 yr^–1 in the open lake. These values are rather low compared with other lakes and can be explained by unfavourable sediment conditions due to wave action and by physiological stress due to the water chemistry.     

Interactive effects of porewater nutrient enrichment, bioturbation and sediment characteristics on benthic assemblages in sandy sediments

Effective management of eutrophic ecosystems requires an understanding of how nutrient input affects the structure and function of benthic communities. The effects of nutrients in soft sediment habitats can be influenced by a variety of factors including sediment characteristics, hydrodynamic exposure, and the presence of bioturbating macroinvertebrates. We used a large scale exclusion experiment (400 m² areas, n = 6) to test if bioturbating lugworms, Arenicola marina mediate the effects of nutrient enrichment. We incorporated small plots (30 × 30 cm) dosed with household garden fertilizer within the lugworm exclusion and corresponding control areas and predicted that the effects of nutrient enrichment would be greater in the absence of lugworms. We found that the increases in nutrient concentrations were higher in the absence of lugworms, but only in the less permeable sediment in the low intertidal zone compared to the more permeable sediment in the high intertidal. Contrary to expectations, the accumulation of nutrients in the plots did not affect the organic matter and chlorophyll levels in the sediment. Interestingly, there were overall negative effects of nutrient additions on some of the most abundant molluscs, Hydrobia ulvae, Retusa obtusa and juvenile Cerastoderma edule. Possible explanations for these adverse effects such as the changes in the sediment chemistry or the physical presence of the fertilizer in the sediment caused by the nutrient additions are discussed. We conclude that the effects of nutrient enrichment in soft sediment habitats on benthic assemblages are determined by the interplay between the presence of bioturbating macroinvertebrates, tidal height and sediment characteristics.     

Quantification of sediment reworking by the Asiatic clam Corbicula fluminea Müller, 1774

Active organisms modify the substratum in which they dwell. This process, called “bioturbation”, affects the way that biogeochemical fluxes are mediated at the substratum–water interface. In the frame of this work, the bioturbation potential of the Asiatic clam Corbicula fluminea was characterized and quantified. We measured the displacement of fluorescent particles by C. fluminea burying in a size-based experimental design in order to explore the effects of body-size on sediment reworking. Our results stress that C. fluminea belongs to the functional group of biodiffusors, and that C. fluminea can be considered as an intermediate sediment reworker. We suggest that bioturbation was mainly induced by the pedal-feeding activity of the clams. Results also showed that, though large clams induced displacement of particles deeper into the sediment, small clams showed the highest net sediment reworking activity. This result was in contrast to the initial hypothesis of biovolume as the main driver for particle displacement by bioturbating organisms. Life-history traits and specific features of pedal-feeding could explain the observed pattern.     

Invasive ecosystem engineers on soft sediment change the habitat preferences of native mayflies and their availability to predators

1. Dreissenid mussels (quagga mussels, Dreissena bugensis, and zebra mussels, D. polymorpha) are invasive species that function as ecosystem engineers in the Laurentian Great Lakes. Dreissena are increasingly abundant on silt, sand and other soft substrates; by altering benthic habitat, these mussels can alter benthic community structure. 2. We used laboratory mesocosm experiments to examine the effects of soft‐sediment Dreissena clusters on the habitat preference of Hexagenia, a native burrowing mayfly that is an important food source to fish. We conducted three experiments to test whether Hexagenia: (1) select for bare sediment, soft sediment covered with live Dreissena (added structure and food resources) or soft sediment with clusters made of empty Dreissena shells (added structure only), (2) prefer a specific density of live Dreissena on soft sediment and (3) select for or avoid sediment with an accumulation of empty Dreissena shells. 3. Contrary to initial expectations, we found that Hexagenia selected for sediment covered with live Dreissena clusters, followed by empty Dreissena shells clusters, and lastly what was previously thought to be the preferred habitat, bare sediment. Not only did Hexagenia prefer Dreissena‐covered sediment, but they also preferred high densities of Dreissena. 4. We also experimentally tested the effects of Dreissena‐covered soft sediment on the availability of Hexagenia to fish. We had three treatment levels representing three distinct habitat types: (1) bare sediment (no Dreissena) treatment in which water was turbid because of mayfly activity, (2) Dreissena‐covered sediment treatment in which water was clear because of Dreissena filtration and (3) Dreissena‐covered sediment with added turbidity. We found that in low light conditions, similar to many locations where both organisms are found to co‐occur, both yellow perch and round goby consumption of Hexagenia significantly decreased when Dreissena covered the bottom sediment. 5. These results suggest that by choosing Dreissena‐covered habitat, Hexagenia receive protection from fish predation in turbid/low light systems. However, protection from predation cannot be the only reason Hexagenia select Dreissena‐covered sediments, as Hexagenia selected for live clusters more often than empty clusters and may be a result of additional food resources.     

A bioturbation classification of European marine infaunal invertebrates

Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical processes in marine systems. In situ quantification of bioturbation can be achieved in a myriad of ways, requiring expert knowledge, technology, and resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption of a trait‐based approach to estimate community bioturbation potential (BP_c). This index can be calculated from inventories of species, abundance and biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking mode and mobility of benthic species. Based on information from the literature and expert opinion, we provide a functional classification for 1033 benthic invertebrate species from the northwest European continental shelf, as a tool to enable the standardized calculation of BP_c in the region. Future uses of this classification table will increase the comparability and utility of large‐scale assessments of ecosystem processes and functioning influenced by bioturbation (e.g., to support legislation). The key strengths, assumptions, and limitations of BP_c as a metric are critically reviewed, offering guidelines for its calculation and application.     

Regulation of nitrous oxide emission associated with benthic invertebrates

1. A number of freshwater invertebrate species emit N₂O, a greenhouse gas that is produced in their gut by denitrifying bacteria (direct N₂O emission). Additionally, benthic invertebrate species may contribute to N₂O emission from sediments by stimulating denitrification because of their bioirrigation behaviour (indirect N₂O emission). 2. Two benthic invertebrate species were studied to determine (i) the dependence of direct N₂O emission on the preferred diet of the animals, (ii) the regulation of direct N₂O emission by seasonally changing factors, such as body size, temperature and availability and (iii) the quantitative relationship between direct and indirect N₂O emission. 3. Larvae of the mayfly Ephemera danica, which prefer a bacteria‐rich detritus diet, emitted N₂O at rates of up to 90 pmol Ind.^?1 h^?1 under in situ conditions and 550 pmol Ind.^?1 h^?1 under laboratory conditions. In contrast, larvae of the alderfly Sialis lutaria, which prefer a bacteria‐poor carnivorous diet, emitted N₂O at invariably low rates of 0–20 pmol Ind.^?1 h^?1. The N₂O emission rate of E. danica larvae was positively correlated with seasonally changing factors (body size, temperature and availability). Direct N₂O emission by E. danica larvae was limited by low temperature in winter, larval development in spring and low availability in summer. 4. Both E. danica and the non‐emitting S. lutaria increased the total N₂O and N₂ emission from sediment in a density‐dependent manner. While N₂O directly emitted by benthic invertebrates can be partially consumed in the sediment (E. danica), non‐emitting species can still indirectly contribute to total N₂O emission from sediment (S. lutaria).     

Rare but large bivalves alter benthic respiration and nutrient recycling in riverine sediments

Bioturbation studies have generally analyzed small and abundant organisms while the contribution to the benthic metabolism by rare, large macrofauna has received little attention. We hypothesize that large, sporadic bivalves may represent a hot spot for benthic processes due to a combination of direct and indirect effects as their metabolic and bioturbation activities. Intact riverine sediments with and without individuals of the bivalve Sinanodonta woodiana were collected in a reach with transparent water, where the occurrence of the mollusk was clearly visible. The bivalve metabolism and its effects on sedimentary fluxes of dissolved gas and nutrients were measured via laboratory incubations of intact cores under controlled conditions. S. woodiana contributed significantly to O₂ and TCO₂ benthic fluxes through its respiration and to \({\text{NH}}_{4}^{ + }\), SRP and SiO₂ regeneration via its excretion. The bivalve significantly stimulated also microbial denitrification and determined a large efflux of CH₄, likely due a combination of bioturbation and biodeposition activities or to anaerobic metabolism within the mollusk gut. This study demonstrates that a few, large individuals of this bivalve produce significant effects on aerobic and anaerobic benthic metabolism and nutrient mobilization. Random sediment sampling in turbid waters seldom catches these important effects due to low densities of large fauna.     

Short Term Bioturbation Activity in the Lagoonal Sediments of Tikehau Atoll (Tuamotu Archipelago,French Polynesia)

To quantify bioturbation activity in Tikehau lagoon, a tracer made of black basaltic sand was poured over the natural white calcareous sediment surface. Three stations respectively located on the inner flat (-3m), the inner slope (-9m), and the lagoon floor (-19m), were studied for short periods of time (48 hours). Bioturbation by macrofauna was quantified by volume of sediment ejected onto the experimental surface and by volume of tracer incorporated into sediment. The results showed a rapid incorporation of sedimented particles at the interface by way of the funnels and burrows of surface deposit feeders and carnivores. Expelled quantities varied with respect to site location: 213 cm³ · m⁻² · 24 h⁻¹ in the inner flat; 98.9 cm³ · m⁻² · 24 h⁻¹ in the inner slope; 7.9 m³ · m⁻² · 24 h⁻¹ in the lagoon floor. Bioturbation by decapod megafauna appeared to be important in the dynamics of the sediments in the deepest areas of the lagoon. In these areas, with almost no hydrodynamical impacts on sediments, bioturbating events were responsible for sediment mixing (despite lower absolute rates than in shallow area). Hydrodynamics controlled the spatial distribution of macroinvertebrate trophic groups by its effects on sedimentation.     

Disturbance, biological legacies and community development in stream mesocosms

Disturbances reduce the biota in stream ecosystems, and leave biological legacies, including remnant species, which potentially influence post-disturbance community development but are poorly understood. We investigated whether three remnant species, the snail Radix peregra, the mayfly Serratella ignita and the freshwater shrimp Gammarus pulex, affected community development in mesocosms that mimicked disturbed habitat patches in streams. Following 21 days of colonisation, we found that the occurrence of legacy effects depended on the identity of the remnant species. Radix had the strongest effect. By bulldozing epilithon, the snails acted as ecological engineers that promoted settlement of filter feeders (Simuliidae) and invertebrate predators (especially Pentaneura and Aphelocheirus) and strongly deterred settlement of non-predatory chironomids (e.g. Heterotrissocladius and Microtendipes). Gammarus increased in density (by 665%) where remnant, probably through rapid reproduction. Baetis and Pentaneura were scarce, and Asellus absent, in remnant Gammarus treatments, as a consequence of interference and/or predation by the amphipods. In contrast, Serratella tolerated the colonisation of immigrant species and did not affect the structure of the developing benthic community. Despite the observed effects on the presence and abundance of benthos, remnant fauna had no significant effect on assemblage taxon richness, or that of any specific trophic group. The contrasting effects of remnant species on immigrant colonisation echoed differences in their life-history traits and foraging behaviours. Our results indicate that biota can generate spatial patchiness of epilithon and benthic invertebrates in stream ecosystems.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Life history and production of the burrowing mayfly Ephoron virgo (Olivier, 1791) (Ephemeroptera: Polymitarcyidae) in the lower Ebro river: a comparison after 18 years

The life history of the burrowing mayfly Ephoron virgo (Olivier, 1791) (Ephemeroptera: Polymitarcyidae) was studied during spring and summer 2005 in the lower Ebro river (Catalonia) and compared to a previous study performed in 1987 (Ibáñez, Escosa, Muñoz and Prat 1991). The results showed an advancement of Ephoron virgo life cycle and an increase of production estimates. In 2005 larval development reached the maximum size one month earlier than in 1987, and adult emergence peak began three weeks earlier. Comparing adult sex ratios (F:M), there was a major presence of females in 2005 (1:4), while the opposite was observed in 1987 (2:1). Secondary production was higher in 2005 than in 1987, obtaining 950 mg dry weight/m²/year with the increment summation method and 1080 mg dry weight/m²/year using the removal summation method. Higher water temperatures were measured for the entire 2005 larval growth period, which were related to higher air temperatures. Therefore, that temperature increment was likely the main cause of changes observed in the Ephoron virgo life cycle.     

Stream drift as a consequence of disturbance by invertebrate predators

Summary We carried out an experimental field study in a Swedish stream in order to determine whether mobile predators enhance the drift of stream insects. We increased the density of nymphs of the predaceous perlid stonefly, Dinocras cephalotes, in an experimental section of a stream up to densities in another more densely populated part of the same stream. The drift of several benthic species increased significantly compared to a control section where D. cephalotes were rare. Experiments carried out in September showed a strongly elevated drift response in nymphs of the mayfly Baetis rhodani only, whereas May experiments resulted in increased drift in B. rhodani as well as the amphipod Gammarus pulex, the stonefly Leuctra fusca, chironomids, and the total number of drifting animals. In September, we found that the drift response of Baetis rhodani to predator disturbance was dependent on the size of mayfly nymphs; small nymphs appeared in greater numbers in the drift nets than did large nymphs. A subsequent laboratory analysis of drift lengths of B. rhodani nymphs supported the hypothesis that small nymphs travel in the drift for longer than do large nymphs, particularly in darkness. We suggest that morphological constraints in vision or swimming performance, or both, cause small nymphs to drift longer. In May, size-dependent drift was less obvious, probably because the size of the nymphs was considerably greater than in September.