Trade-offs in the response of mayflies to low oxygen and fish predation

Summary We examined how mayfly larvae (Ephemeroptera,Callibaetis montanus) balance the conflicting demands of avoiding both benthic hypoxia and fish predators. Using vertical oxygen and temperature gradients typical of ice-covered lakes, we observed the behavior of mayflies in the presence and absence of fish. In the absence of fish and with adequate oxygen, mayflies spent most of the time on the bottom substrate. As benthic oxygen concentration declined, mayflies increased their activity and moved up in the water column. In the presence of fish and with adequate oxygen, mayflies spent even more time associated with the bottom substrate and reduced their activity levels. As benthic oxygen concentrations declined, mayflies increased their activity and moved up in the water column, but to a lesser extent than when fish were absent. Because of this depression in activity and reluctance to leave the bottom substrate, mayflies endured lower oxygen concentrations in the face of predation threat relative to when fish were absent. Despite this trade-off, benthic hypoxia resulted in increased mortality due to fish predation. Because benthic invertebrates vary in their ability to tolerate hypoxia and in their vulnerability to fish predators, periods of benthic hypoxia could lead to selective predation on some taxa and be an important force structuring benthic invertebrate assemblages.     

The effect of Echinorhynchus borealis (Acanthocephala) infection on the anti-predator behavior of a benthic amphipod

In benthic habitats, predators can generally not be detected visually, so olfaction may be particularly important for inducing anti-predation behaviors in prey organisms. Manipulative parasites infecting benthic hosts could suppress these responses so as to increase the probability of predation and thus trophic transmission. We studied how infection with the acanthocephalan Echinorhynchus borealis affects the response of the benthic amphipod Pallasea quadrispinosa to water conditioned by burbot (Lota lota), the parasite's definitive host. In normal lake water, refuge use by infected and uninfected amphipods was similar, but when exposed to burbot-conditioned water, uninfected amphipods spent much more time hiding than infected amphipods. Thus, rather than affecting ambient hiding behavior, E. borealis infection seems to alter host response to a predator. A group of amphipods sampled from a postglacial spring that is devoid of fish predators exhibited only a weak response to burbot-conditioned water, perhaps suggesting these anti-predator behaviors are costly to maintain. The hiding behavior of spring and infected amphipods was very similar. If the reduced refuge use by the spring amphipods reflects adaptation to a predator-free environment, this indicates that E. borealis severely weakens its host's anti-predator behavior. Presumably this increases the likelihood of parasite transmission.     

Impacts of introduced brown and rainbow trout on benthic invertebrate communities in shallow New Zealand lakes

1. Brown and rainbow trout have been introduced to many inland waters in New Zealand, but research on the impacts on native communities has focused mainly on streams. The purpose of this study was to compare the benthic communities of trout and troutless lakes. Based on previous studies in North America and Europe, we predicted that the benthic biomass, and especially the abundance of large invertebrates, would be lower in lakes with trout as compared to those without. We surveyed the invertebrate fauna of 43 shallow, high‐elevation lakes (26 with and 17 without trout) in four geographic clusters on the central South Island and then conducted a detailed quantitative study of invertebrate biomass and community structure in 12 of these lakes. 2. Benthic community composition and diversity of lakes with and without trout were nearly identical and biomass was as high or higher in the lakes with as without trout. There was no evidence that trout have caused local extinctions of benthic invertebrates. Although the proportional abundance of large‐bodied aquatic was slightly lower in lakes with than without trout, the abundance of several groups of large‐bodied benthic taxa (dragonflies, caddisflies and water bugs) did not differ. 3. Our findings are in contrast to those in North American and Europe where trout introductions into previously troutless lakes have led to declines in the abundance of benthic invertebrates, especially large‐bodied taxa. We propose that the modest effects of trout in New Zealand could be explained by (i) the high areal extent of submergent vegetation that acts as a benthic refuge, (ii) low intensity of trout predation on benthic communities and/or (iii) characteristics of the benthic invertebrates that make them relatively invulnerable to fish predation. 4. Regardless of the relative importance of these hypotheses, our results emphasise that the same invertebrates occurred in all of the lakes, regardless of size, elevation and presence of trout, suggesting habitat generalists dominate the benthic fauna in shallow New Zealand lakes.     

Predation of perch on vendace larvae: diet composition in an oligotrophic lake and digestion time of the larvae

The diet of perch Perca fluviatilis was studied to reveal possible predation on vendace Coregonus albula larvae in an oligotrophic lake. Perch diet changed with the size of the fish: small perch ate mainly zooplankton and the diet shifted more to benthic invertebrates and fishes in larger perch. There were also annual and spatial differences in the diet, probably reflecting differences in the availability of prey animals. Perch predation on vendace larvae was only observed in the area with high availability of the larvae. The result suggested strengthened predation when the density of the larvae increases. According to bioenergetics modelling, the perch population increased natural mortality of vendace larvae only marginally. Food intake of spawning female perch was slightly reduced, whereas spawning males fed similarly to non-spawning males. Hence, the spawning period of perch was only a minor refuge for vendace larvae. Laboratory experiments of perch digestion rate demonstrated that, due to rapid digestion of the small fish larvae, diet sampling interval should not be >2 h in the field.     

Community structure of benthic invertebrates in interior Alaskan (USA) streams and rivers

Taxonomic composition of benthic invertebrates in interior Alaskan streams and rivers is summarized from published and unpublished data. Diptera dominate the Alaskan stream fauna and constitute a larger proportion of the benthos in Alaskan streams than in streams of temperate North America. Plecoptera and Ephemeroptera are the next most abundant in Alaskan streams with Trichoptera generally very scarce. Several orders that occur regularly in streams of temperate North America are absent (or in very low abundance) in interior Alaskan streams: Hemiptera, Odonata, Megaloptera, Coleoptera. Net-spinning caddisflies, burrowing mayflies, and several families of stoneflies (Pteronarcyidae, Peltoperlidae and Perlidae) are conspicuous by their absence or extreme scarcity. Taxonomic composition varies significantly among hydrologic regions (major watersheds) and among stream types (springs, headwater streams, small rivers, and large rivers). Only two taxa (Chironomidae and Nemouridae) significantly increase in proportional contribution from south to north while many taxa decrease.     

Contrasting effects of an invasive crayfish (Procambarus clarkii) on two temperate stream communities

1. The effects of omnivorous exotic species on native communities are often difficult to predict because of the broad diets and behavioural flexibility of the omnivore, and the diverse abiotic and biotic characteristics of invaded systems. We investigated experimentally the effects of a gradient of density of the introduced, omnivorous red swamp crayfish Procambarus clarkii (Decapoda: Cambaridae) on two stream communities in southern California, U.S.A. 2. The Ventura River is a clear, flowing stream with a cobble substratum, with abundant algae but low densities of large invertebrates, small herbivores and snails. The Santa Ynez River at the time of the study consisted of a series of drying pools underlain by sand, with abundant charophytes, large predatory invertebrates and herbivores, including snails. 3. In the Ventura River, periphyton biomass and inorganic sediment decreased with increasing crayfish abundance, but in the Santa Ynez River, periphyton and sediment were unrelated to crayfish densities. 4. In the Ventura River, the biomass and density of all benthic invertebrates combined, chironomids, micropredators, the meiofauna (chydorid cladocerans, copepods and ostracods), and specific predatory and herbivorous taxa, as well as taxon richness, were negatively related to crayfish density. In the Santa Ynez River, the biomass and average body size of benthic invertebrates, predatory invertebrates, herbivores and chironomids, but not total invertebrate density or taxon richness, were negatively related to crayfish density. 5. Fewer large predatory invertebrates and snails (Physella gyrina) in both streams, and baetid mayflies in the Ventura River, were visible at night in channels where crayfish were abundant. Snails responded to crayfish by moving above the water line in the Santa Ynez River, but not in the Ventura River. 6. We suggest that the same omnivore had different effects on these neighbouring streams because of crayfish predation on large invertebrates in the Santa Ynez River and the scarcity of such prey in the Ventura River, leading to increased crayfish grazing on periphyton, and reductions in periphyton‐associated invertebrates, in the Ventura River.     

Predator defense along a permanence gradient: roles of case structure, behavior, and developmental phenology in caddisflies

Species replacements along freshwater permanence gradients are well documented, but underlying mechanisms are poorly understood for most taxa. In subalpine wetlands in Colorado, the relative abundance of caddisfly larvae shifts from temporary to permanent basins. Predators on caddisflies also shift along this gradient; salamanders (Ambystoma tigrinum nebulosum) in permanent ponds are replaced by predaceous diving beetles (Dytiscus dauricus) in temporary habitats. We conducted laboratory and field experiments to determine the effectiveness of caddisfly cases in reducing vulnerability to these predators. We found that larvae of a temporary-habitat caddisfly (Asynarchus nigriculus) were the most vulnerable to salamanders. Two relatively invulnerable species (Limnephilus externus, L. picturatus) exhibited behaviors that reduced the likelihood of detection and attack, whereas the least vulnerable species (Agrypnia deflata) was frequently detected and attacked, but rarely captured because cases provided an effective refuge. Vulnerability to beetle predation was also affected by cases. The stout cases of L. externus larvae frequently deterred beetle larvae, whereas the tubular cases of the other species were relatively ineffective. Two of these vulnerable species (A. nigriculus and L. picturatus) often co-occur with beetles; thus, case construction alone is insufficient to explain patterns of caddisfly coexistence along the permanence gradient. One explanation for the coexistence of these two species with beetles is that they develop rapidly during early summer and pupate before beetle larvae become abundant. One species (L. picturatus) pupates by burying into soft substrates that serve as a refuge. The other (A. nigriculus) builds stone pupal cases, which in field experiments, more than doubles survival compared to organic pupal cases. The combined results of these experiments suggest that caddisfly distributions along permanence gradients depend on a suite of primary and secondary predator defenses that include larval and pupal case structure, predator-specific escape behaviors, and the phenology of larval development.     

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).     

The impact of predation by burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L.) on the macroinvertebrate community in the littoral zone of a large lake

We studied the effects of predation by juvenile burbot (Lota lota) on the macroinvertebrate community in mesocosm experiments in the stony littoral zone of Lake Constance, a large prealpine lake in Central Europe. Although the growth data of the burbot suggest that the benthivorous fish exerts strong predation pressure on the invertebrate community, the predicted level of consumption is only poorly reflected by changes in biomass and abundance of most invertebrate prey taxa. High exchange rates of the prey between the mesocosms and the ambient littoral environment apparently masked the true predation effects of fish. Also, life-history events such as hatching or synchronised emergence of larvae led to temporal effects that obscured the impact of predation. However, for the dominant prey organism, the freshwater amphipod Gammarus roeseli, direct lethal effects appeared when its migration was limited. When exchange with the ambient littoral zone was possible, abundance and biomass of G. roeseli were unexpectedly high in the mesocosms stocked with burbot, indicating behavioural responses to fish presence. During the experiment, the burbot gained ca. 15% in length and about 60% in body mass. According to stomach content analyses at the end of the experiment, the ingested prey consisted mainly of relatively large and abundant invertebrates. Our study indicates that predation by juvenile burbot should be an important factor in structuring the benthic invertebrate community in terms of qualitative and quantitative effects.     

The effect of large macroinvertebrate herbivores on sessile epibenthos in a mountain stream

Studies of benthic invertebrates in lakes and streams suggest thatlarge-bodied herbivores are more efficient grazers than smallerones. In order to assess the effect of larger herbivores on smallergrazing invertebrates, the presence of dominant grazer taxa wasmanipulated in streamside troughs in a first order temperaterainforest stream in British Columbia. The presence of mayflies(Ameletus sp.) and tailed frog tadpoles (Ascaphustruei) reduced both algal biomass and the abundance of herbivorouschironomids (Orthocladiinae) on ceramic tiles. This confirms thatlarge mobile grazers in streams have a negative effect on smallersessile invertebrate grazers either through resource competition ordirect consumption (predation).     

Invertebrate responses to short-term water abstraction in small New Zealand streams

1. Small permanent streams are coming under increasing pressure for water abstraction. Although these abstractions might only be required on a short‐term basis (e.g. summer time irrigation), the highest demand for water often coincides with seasonal low flows. 2. We constructed weirs and diversions that reduced discharge in three small streams (<4 m width) to test the hypotheses that short‐term water abstractions would decrease habitat availability and suitability for invertebrates, resulting in increased invertebrate drift, reduced taxonomic richness and decreased benthic invertebrate densities. 3. We sampled benthic invertebrates, invertebrate drift and periphyton at control (upstream) and impact (downstream) sites on each stream before and during 1 month of discharge reduction. 4. Discharge decreased by an average of 89–98% at impact sites and wetted width decreased by 24–30%. Water depth decreased by 28–64% while velocity decreased by 50–62%. Water conductivity, temperature and dissolved oxygen showed varying responses to flow reduction among the three streams, whereas algal biomass and pH were unaffected in all streams. 5. The densities of invertebrate taxa tended to increase in the impact reaches of these streams, even though invertebrate drift increased at impact sites in the first few days following discharge reduction. There were a higher proportion of mayflies, stoneflies and caddisflies at the impact site on one stream after flow reduction. There were no changes to the number of taxa or species evenness at impact sites. 6. Our results suggest that for these small streams, the response of invertebrates to short‐term discharge reduction was to accumulate in the decreased available area, increasing local invertebrate density.     

Overview of the chemical ecology of benthic marine invertebrates along the western Antarctic peninsula

Thirteen years ago in a review that appeared in the American Zoologist, we presented the first survey of the chemical and ecological bioactivity of Antarctic shallow-water marine invertebrates. In essence, we reported that despite theoretical predictions to the contrary the incidence of chemical defenses among sessile and sluggish Antarctic marine invertebrates was widespread. Since that time we and others have significantly expanded upon the base of knowledge of Antarctic marine invertebrates' chemical ecology, both from the perspective of examining marine invertebrates in new, distinct geographic provinces, as well as broadening the evaluation of the ecological significance of secondary metabolites. Importantly, many of these studies have been framed within established theoretical constructs, particularly the Optimal Defense Theory. In the present article, we review the current knowledge of chemical ecology of benthic marine invertebrates comprising communities along the Western Antarctic Peninsula (WAP), a region of Antarctica that is both physically and biologically distinct from the rest of the continent. Our overview indicates that, similar to other regions of Antarctica, anti-predator chemical defenses are widespread among species occurring along the WAP. In some groups, such as the sponges, the incidence of chemical defenses against predation is comparable to, or even slightly higher than, that found in tropical marine systems. While there is substantial knowledge of the chemical defenses of benthic marine invertebrates against predators, much less is known about chemical anti-foulants. The sole survey conducted to date suggests that secondary metabolites in benthic sponges are likely to be important in the prevention of fouling by benthic diatoms, yet generally lack activity against marine bacteria. Our understanding of the sensory ecology of Antarctic benthic marine invertebrates, despite its great potential, remains in its infancy. For example, along the WAP, community-level non-consumptive effects occur when amphipods chemically sense fish predators and respond by seeking refuge in chemically-defended macroalgae. Such interactions may be important in releasing amphipods from predation pressure and facilitating their unusually high abundances along the WAP. Moreover, recent studies on the sensory biology of the Antarctic keystone sea star Odontaster validus indicate that chemotactile-mediated interactions between conspecifics and other sympatric predatory sea stars may have significant ramifications in structuring community dynamics. Finally, from a global environmental perspective, understanding how chemical ecology structures marine benthic communities along the WAP must increasingly be viewed in the context of the dramatic impacts of rapid climatic change now occurring in this biogeographic region.     

The effects of nymphaeid (Nuphar lutea) density and predation by perch (Perca fluviatilis) on the zooplankton communities in a shallow lake

1. The effects of addition of juvenile perch (Perca fluviatilis) on the microcrustacean and rotifer communities associated with nymphaeid beds were studied, at three different plant densities [high (normal), medium (reduced by a half) and low (reduced to a third of normal)], in eighteen 2 m × 1 m enclosures in a shallow lake. 2. At the low and medium densities of lilies, Daphnia densities were high in the absence of perch but very low in the presence of perch. They increased, even in the presence of perch, to high densities (comparable with those in the absence of perch) at the highest plant density. Body sizes of Daphnia hyalina were consistent with high predation by perch at low and medium plant densities but reduced predation at high plant densities. Patterns of chlorophyll a concentration, in the presence of perch, inversely reflected those of D. hyalina density. 3. At naturally high densities but not at reduced densities, the plants appeared to act as refuges against predation for the Daphnia. Reductions in oxygen concentrations in the plant beds were not responsible for the refuge effect, nor could there be avoidance of the beds by the fish. The mechanism of the refuge effect must therefore lie in frustration of the process of capture of the Daphnia by the fish. 4. Numbers of other small Crustacea and rotifers were mostly unaffected by fish predation. Numbers of Asplanchna sp., Chydorus sphaericus and copepodites were higher in the presence of fish and, although there was no main effect of fish on numbers of Diaphanosoma brachyurum, there was a plant–fish interaction, with this species being less abundant in the presence of fish at low plant densities but more abundant in the presence of fish at medium and high plant densities. Main effects of plants were few, with only Asplanchna sp. and Keratella sp. decreasing in numbers with increasing plant density. Most taxa changed in numbers with time but interaction effects between time and plants, and fish and plants, were few.     

Mesofauna associated with the marine sponge Amphimedon viridis. Do its physical or chemical attributes provide a prospective refuge from fish predation?

Marine sponges often harbor an abundance of associated organisms. We characterized mesofauna associated with the common tropical sponge Amphimedon viridis, and then tested whether physical (spicules) or chemical (lipophilic or hydrophilic extracts) properties of this sponge provide a prospective refuge for mesofauna from fish predation. Sponge analyses revealed a moderately diverse and numerically rich community of sponge-associated mesofauna comprised primarily of mesocrustaceans (82% of total fauna). Eighty-nine percent of these were amphipods, but smaller numbers of tanaids, decapods, and isopods also occurred. Quantitative sampling of outer surfaces and interstices of fifteen A. viridis yielded a total mean ± 1 SD density of sponge–associated mesofauna of 53 ± 9.3 individuals per 100 cm³ wet sponge tissue. Among the numerically dominant amphipods, 65% occurred on outer sponge surfaces where they are most vulnerable to fish predators. We evaluated whether A. viridis provides a prospective refuge from predation by assessing the palatability of this sponge to the sympatric generalist pinfish Lagodon rhomboides. When presented small (2 mm) bite-size pieces of whole sponge tissue, similar in size to what fish might incidentally ingest should they attempt to consume sponge-mesofaunal associates, pinfish displayed strong feeding deterrence. Alginate food pellets containing tissue-level concentrations of sponge spicules caused a weak but significant deterrent response. In contrast, alginate pellets containing tissue-level concentrations of either lipophilic or hydrophilic extracts of A. viridis were highly deterrent to pinfish. Thus, chemical, and to a considerably lesser degree, physical defenses (spicules) may contribute to this sponge serving as a protective refuge for associated mesofauana.     

Top-down and bottom-up impacts of juvenile fish in a littoral reed stand

1. Research has often focused on pelagic food chains and processes of lakes; less is known about the contribution of benthic energy flows to whole‐lake ecosystem energetics. This stems from the fact that the shoreline and littoral habitats, which provide a key linkage between sediment and water column, have only recently become a significant focus for study. 2. This study aimed to quantify the feeding and phosphorus allocation of a juvenile fish community in a littoral zone of a shallow lake in response to the biomass succession of the invertebrate prey community. Habitats comprising reed and adjacent open water were sampled over two consecutive years during day and night. 3. Although there were substantial year‐to‐year differences in the biomass of invertebrates, the fish community composition, diet consumption rates and phosphorus allocations were very similar in both study years. Biomasses and predation impacts by juvenile fish on prey groups were substantially higher within the reeds than in the adjacent open water habitat. This may be explained by the refuge‐seeking behaviour of the fish. 4. In general, invertebrates were negligibly influenced by fish feeding, with the exception for a strong top‐down control of large cladocerans. In response to the resulting low Daphnia biomass, fish were forced to switch to a higher degree of benthivory. Consequently, juvenile fish in littoral reed stands may shift benthic‐derived energy and phosphorus via the excretion of soluble reactive phosphorus into the open water.     

Seasonal dynamics of long-chain polyunsaturated fatty acids in littoral benthos in the upper Yenisei river

We studied composition and concentrations of fatty acids (FAs) in benthos from pebbly littoral region of the Yenisei River in a sampling site near Krasnoyarsk city (Siberia, Russia) for 1 year from March 2003 to February 2004. Special attention was paid to major long-chain polyunsaturated fatty acids (PUFAs) of the ω3 family: eicosapentaenoic acid (EPA, 20:5ω3) and docosahexaenoic acids (DHA, 22:6ω3). In phytobenthos, which was dominated by diatoms, the annual maxima of EPA and DHA pool occurred in spring and early summer. In zoobenthos, EPA and DHA pool peaked in autumn, due mainly to an increase of the biomass of dominant taxa (gammarids) and to a moderate increase of the PUFA content per body weight. Seasonal peaks of EPA in overwintering insect larvae (chironomids and caddisflies) generally coincided with those of biomass of these larvae, while there was no such trend for amphipods and oligochaetes. In spring and early summer, the main part of ω3 PUFA, 40–97% of total amount, in the littoral region was contained in biomass of producers, i.e., benthic microalgae, and in autumn it was transferred to primary consumers—benthic invertebrates, which contained ∼76–93% of total ω3 PUFAs.     

Comparison of seasonal dynamics of the essential PUFA contents in benthic invertebrates and grayling Thymallus arcticus in the Yenisei river

Seasonal dynamics of contents of essential polyunsaturated fatty acids (PUFA) in dominant groups of benthic invertebrates: gammarids (Gammaridae, Amphipoda), oligochaetes (Oligochaeta), chironomid larvae (Chironomidae, Diptera) and caddisfly larvae (Trichoptera), and dominant benthivorous fish, Siberian grayling Thymallus arcticus, have been studied in ecosystem of the large Siberian river. During the year of the study most benthic invertebrate taxa showed significant variations in the contents of both C-18 and long chain C-20–22 PUFAs. In contrast, the fish, which consumed the zoobenthos, had no significant seasonal variations in long chain PUFAs' contents. Thereby, the fish, as organisms of relatively higher evolution level than invertebrates, was supposed to have more strict metabolic control of long chain PUFAs' contents. Evidence was obtained that the studied fish species, grayling, may be capable to convert dietary EPA into DHA.     

The effect of early and late hatching on the escape response of walleye pollock (Theragra chalcogramma) larvae

Hatching of fish eggs fertilized at the same time occurs overa period of several days. Differences in the escape responseof fish larvae during the hatching period have not hithertobeen studied. In this study, the escape response of walleyepollock (Theragra chalcogramma) larvae over the hatching periodwas examined. Escape speed, response to multiple touches witha fine probe, response to water currents generated by a predatorand predation by euphausiids (Thysanoessa inermis) and amphipods(Pleusirus secorrus) were measured in the laboratory. Otolithmeasurements of field-collected larvae support a broad hatchingperiod for walleye pollock eggs in the sea similar to that observedin the laboratory. The escape response of walleye pollock larvaewas affected by rank in the order of hatching, thus with respectto predation, hatching order may affect the survival of larvaein the sea. Early hatching larvae were smaller, less sensitiveto tactile stimulation, had a slower, weaker escape responseand higher laboratory rates of predation mortality than thosethat hatched later.     

Trace Metals in Mesozooplankton of the North Sea: Concentrations in Different Taxa and Preliminary Results on Bioaccumulation in Copepod Collectives (Calanus finmarchicus/C. helgolandicus)

Trace metals (Cd, Cu, Zp, Pb, Ni) were determined in different zooplankton taxa to assess their role in the biogeochemical cycle in the North Sea. Results of semi-static bioaccumulation experiments on board ship using collectives of Calanus finmarchicus/C. helgolandicus were in agreement with a net accumulation strategy, thus meeting an inevitable precondition for their utilization as biomonitors. Cd, Cu and Zn concentrations in field samples of zooplankton displayed a pronounced variability between taxa, with Cd ranging from 0.13 mg kg-1 (d.w.) in fish larvae to 5I mg kg-1 in hyperiid amphipods. Almost no information is available about metal requirements of copepods. However, most of our and worldwide reported Cu concentrations in calanoid copepods (6-22 mg kg⁻¹) coincided with estimates of enzymatic requirements of decapod crustaceans. Cd levels in Calanus collectives increased significantly from 0.68 mg kg-1 in the German Bight to 11 mg kg⁻¹ at some stations around Scotland above 59° N, in line with literature data reported for benthic decapod crustaceans.     

Benthivorous fish reduce stream invertebrate drift in a large-scale field experiment

Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or to escape after encountering a predator. While the control of the diurnal patterns of invertebrate drift activity by fish predators has received considerable interest, it remains unclear whether benthivorous fish reduce or increase drift activity. We performed a large-scale field experiment in a second-order stream to test if invertebrate drift was controlled by two benthivorous fish species (gudgeon Gobio gobio and stone loach Barbatula barbatula). An almost fishless reference reach was compared with a reach stocked with gudgeon and loach, and density and structure of the invertebrate communities in the benthos and in the drift were quantified in both reaches. The presence of gudgeon and stone loach reduced the nocturnal drift of larvae of the mayfly Baetis rhodani significantly, in contrast to the findings of most previous studies that fish predators induced higher night-time drift. Both drift density and relative drift activity of B. rhodani were lower at the fish reach during the study period that spanned 3 years. Total invertebrate drift was not reduced, by contrast, possibly due to differences in vulnerability to predation or mobility between the common invertebrate taxa. For instance, Chironomidae only showed a slight reduction in drift activity at the fish reach, and Oligochaeta showed no reduction at all. Although benthic community composition was similar at both reaches, drift composition differed significantly between reaches, implying that these differences were caused by behavioural changes of the invertebrates rather than by preferential fish consumption. The direction and intensity of changes in the drift activity of stream invertebrates in response to the presence of benthivorous fish may depend on the extent to which invertebrate taxa can control their drifting behaviour (i.e. active versus passive drift). We conclude that invertebrate drift is not always a mechanism of active escape from fish predators in natural streams, especially when benthos-feeding fish are present.