Stonefly nymphs use hydrodynamic cues to discriminate between prey

Summary Playback experiments conducted in a Rocky Mountain, USA, stream determined whether predatory stonefly nymphs (Kogotus modestus; Plecoptera: PerlodiMae) used hydrodynamic cues to discriminate prey species from nonprey species. In the laboratory we recorded pressure wave patterns associated with swimming escape behavior of Baetis bicaudatus (Baetidae), the favored mayfly prey species, and those of a nonprey mayfly, Ephemerella infrequens (Ephemerellidae). We video taped the responses of 24-h starved Kogotus to Baetis playbacks, Ephemerella playbacks or no playbacks made by oscillating (or not) live mayflies (Ephemerella) or clear plastic models placed within in situ flow-through observation boxes. The probability of attacks per encounter with Baetis playbacks was highest and independent of the model type used, but Kogotus also showed an unexpected high probability of attacks per encounter when Ephemerella playbacks were made through live Ephemerella. Thus, Kogotus discriminated between Baetis and Ephemerella swimming patterns but only when playbacks were made through the plastic model. Kogotus never attacked motionless mayflies or motionless plastic models. We allowed some Kogotus to successfully capture one small Baetis immediately before playbacks, which resulted in a much higher probability of attacks per encounter with Baetis playbacks on either model and a heightened discrimination of prey versus nonprey playbacks. The probability of attacks per encounter by Kogotus with live Baetis swimming under similar experimental conditions was strikingly similar to its response to Baetis playbacks made by oscillating the plastic model after a successful capture. Order of playback presentation (Baetis first or Ephemerella first) did not influence predatory responses to mayfly swimming patterns. This study is the first to document the use of hydrodynamic cues by stream-dwelling predators for discrimination of prey from nonprey and provides a mechanism to explain selective predation by stoneflies on Baetis in nature.     

Heptageniidae (Insecta,Ephemeroptera) of Thailand

Nine genera and twenty-two species of heptageniid mayflies from Thailand are defined in this present work as well as one suggested further subgenus, Compsoneuria (Siamoneuria) kovaci (species “incertae sedis”) including some particular characters. Taxonomic remarks, diagnoses, line drawings of key characters, distribution, habitat and biological data, and a larval key to the genera and species are provided. The chorionic eggs of eight genera and eight species were observed and shown using a scanning electron microscope.     

Life history studies of anisocentropus kirramus neboiss (trichoptera: Calamoceratidae) in a tropical Australian rainforest stream

Eggs and larvae of Anisocentropus kirramus were collected from leaf packs in riffles and pools in a small upland rainforest stream in tropical Queensland. Adults were collected in floating emergence traps. Egg masses contained 80–100 eggs. None developed in water at 12–15°C but at 22–25°C larvae hatched in 3–10 days. There were five larval instars and complete development appeared to take several months. Instars II‐V were present in all nine months sampled; instar I was present in all but one month. Adults emerged in all ten months sampled, but there was clear seasonality with peak emergence in the summer. The sex ratio of adults was 1:1.     

Life history and larval feeding of some species of Ephemeroptera and Plecoptera (Insecta) in the Sierra Nevada (Southern Iberian Peninsula)

We monitored two high mountain streams from the Sierra Nevada (Southern Spain) monthly throughout an annual cycle, measuring and analyzing physical and chemical parameters. Temperature was hourly registered. We studied life cycles and larval feeding of four mayfly species [Serratella ignita (Poda, 1761), Serratella spinosa nevadensis (Alba-Tercedor, 1981), Alainites muticus (Linnaeus, 1758), and Baetis alpinus Pictet, 1843] and two stonefly species [Amphinemura triangularis (Ris, 1902) and Capnioneura mitis Despax, 1932]. Some species changed their strategies (growth rate, life cycle duration, and/or voltinism) in accordance with different conditions (depending on the altitude and temperature). The gut content data showed that species fed mainly on detritus (as collector-gatherers or as shredders), except B. alpinus that behaved as a scraper in one of the streams. Differences in the larval diet of the same species in different streams, and even through its life, support the idea of not using general models to extrapolate the function of a species from a higher taxonomical level. Handling editor: D. Dudgeon     

The status of neutral insects,mosquitoes in the food of natural enemies in tea gardens

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The influence of spatial heterogeneity on the behavior and growth of two herbivorous stream insects

Environmental heterogeneity can affect the behavior of organisms, but the consequences of patchiness for organismal energetics (e.g., growth, fitness) are not well understood. This study demonstrates that spatial heterogeneity can affect the growth of aquatic stream insects in laboratory streams, and reveals the behavioral mechanisms for these effects. In a 2×2 factorial design, I experimentally manipulated resource distribution (homogeneous vs. patchy, with the same overall resource levels) and current velocity (fast vs. slow) to investigate the direct and interactive effects of these factors on the drift behavior and growth of two mobile stream grazers, the mayflies Baetis bicaudatis and Epeorus deceptivus. B. bicaudatis nymphs grew larger in environments with homogeneously distributed resources than in patchy environments, and both species grew larger in fast than slow current environments. Patterns of drift behavior over the course of the study corresponded to observed differences in growth. Both species grew to larger body size in treatments where they drifted more successfully among substrates (fast-current treatments) and where they entered the drift less frequently (fast current for both species, and homogeneous treatments for B. bicaudatis). Overall, these results demonstrate that patchiness can significantly influence both the behavior of aquatic insects and the size to which these insects grow. In the light of previously published relationships between nymphal mayfly body mass and fecundity, these results suggest that patchiness in streams may have important consequences for mayfly populations.     

Fitness and community consequences of avoiding multiple predators

We investigated the fitness and community consequences of behavioural interactions with multiple predators in a four-trophic-level system. We conducted an experiment in oval flow-through artificial-stream tanks to examine the single and interactive sublethal effects of brook trout and stoneflies on the size at emergence of Baetis bicaudatus (Ephemeroptera: Baetidae), and the cascading trophic effects on algal biomass, the food resource of the mayflies. No predation was allowed in the experiment, so that all effects were mediated through predator modifications of prey behaviour. We reared trout stream Baetis larvae from just before egg development until emergence in tanks with four treatments: (1) water from a holding tank with two brook trout (trout odour), (2) no trout odour + eight stoneflies with glued mouthparts, (3) trout odour + stoneflies and (4) no trout odour or stoneflies. We ended the experiment after 3 weeks when ten male and ten female subimagos had emerged from each tank, measured the size of ten male and ten female mature nymphs (with black wing pads), and collected algal samples from rocks at six locations in each tank. To determine the mechanism responsible for sublethal and cascading effects on lower trophic levels we made day and night observations of mayfly behaviour for the first 6 days by counting mayflies drifting in the water column and visible on natural substrata in the artificial streams. Trout odour and stoneflies similarly reduced the size of male and female Baetis emerging from artificial streams, with non-additive effects of both predators. While smaller females are less fecund, a fitness cost of small male size has not been determined. The mechanism causing sublethal effects on Baetis differed between predators. While trout stream Baetis retained their nocturnal periodicity in all treatments, stoneflies increased drift dispersal of mayflies at night, and trout suppressed night-time feeding and drift of mayflies. Stoneflies had less effect on Baetis behaviour when fish odour was present. Thus, we attribute the non-additivity of effects of fish and stoneflies on mayfly growth to an interaction modification whereby trout odour reduced the impact of stoneflies on Baetis behaviour. Since stonefly activity was also reduced in the presence of fish odour, this modification may be attributed to the effect of fish odour on stonefly behaviour. Only stoneflies delayed Baetis emergence, suggesting that stoneflies had a greater sublethal effect on Baetis fitness than did trout. Delayed emergence may reduce Baetis fitness by increasing risks of predation and parasitism on larvae, and increasing competition for mates or oviposition sites among adults. Finally, algal biomass was higher in tanks with both predators than in the other three treatments. These data implicate a behavioural trophic cascade because predators were not allowed to consume prey. Therefore, differences in algal biomass were attributed to predator-induced changes in mayfly behaviour. Our study demonstrates the importance of considering multiple predators when measuring direct sublethal effects of predators on prey fitness and indirect effects on lower trophic levels. Identification of an interaction modification illustrates the value of obtaining detailed information on behavioural mechanisms as an aid to understanding the complex interactions occurring among components of ecological communities. Received: 20 March 1997 / Accepted: 29 September 1997     

Habrophlebia antoninoi sp.n., a New Species from Spain,with an Account of the European Species of Habrophlebia Eaton, 1881 (Ephemeroptera: Leptophlebiidae,Habrophlebiinae)

The adults and egg of Habrophlebia antoninoi sp. n. are described and illustrated from material collected in southern Spain. Although the genitalia of the new species is characteristic of Habrophlebia, the chorionic sculpturing of the egg and the shape and venation of hind wings are unique among European members of the genus. A discussion of its relationships with other European species of Habrophlebia is included and a key to male imagines of European species of Habrophlebia is provided.     

Mayflies (ephemeroptera), the most “primitive” winged insects,have telotrophic meroistic ovaries

Germ line cell cluster formation in ovarioles of three different stages, each from a different mayfly species, was studied using ultra-thin serial sectioning. In the analysed ovariole of Cloeön sp., only one linear, zigzag germ line cell cluster was found, consisting of sibling cells connected by intercellular bridges which represent remnants of preceding synchronized mitotic cycles followed by incomplete cytokinesis. A polyfusome stretched through all sibling cells. At the tip of the ovariole, cytokinesis occurred without preceding division of nuclei; thus, intercellular bridges were lined up but the remaining cytoplasm between the bridges had no nuclei. The analysed Siphlonurus armatus vitellarium contained five oocytes at different stages of development. Each oocyte in the vitellarium was connected via a nutritive cord to the linear cluster of its sibling cells in the terminal trophic chamber. Each cluster had the same architecture as was found in Cloëon. The 3-dimensional arrangement and distribution of closed intercellular bridges strongly suggest that all five clusters are derived from a single primary clone. The position of oocytes within each cluster is random. However, each oocyte is embraced by follicular or prefollicular cells whilst all other sibling cells are enclosed by somatic inner sheath cells, clearly distinguishable from prefollicular cells. In the analysed ovariole of Ephemerella ignita, two small linear clusters were found in the tropharium beside two single cells, two isolated cytoplasmic bags with intercellular bridges but no nuclei, and some degenerating aggregates. One cluster was still connected to a growing oocyte via a nutritive cord. In all species the nurse cells remained small and no indications of polyploidization were found. We suggest that this ancient and previously unknown telotrophic meroistic ovary has evolved directly from panoistic ancestors.     

Developmental responses to predation risk in morphologically defended mayflies

Densities and species composition of predators could affect morphological defences, larval development and the timing of emergence of their prey. To address this issue we studied the morphology and life history of an ephemerellid mayfly, Ephemerella invaria, from two streams in a deciduous forested drainage basin in central New York. Both streams contained predatory fish, but densities and species composition of fish differed. A field survey provided evidence that Ephemerella inhabiting a stream with 10 fish species and high relative densities of fish emerged several weeks earlier and at smaller sizes than Ephemerella inhabiting a nearby tributary with ~2 fish species and low relative densities of fish. However, the two populations of mayflies showed no differences in defensive morphology or growth rates. In laboratory rearing experiments, we exposed Ephemerella larvae from these two locations to fish chemical cues or control water (no fish) over 2 months to test whether differences in life histories could be attributed to fish. Fish cues induced faster larval development, but also smaller size of mature Ephemerella individuals from both high and low predator locations. Although shorter development times in more dangerous environments could increase larval survival, smaller size of females results in a fecundity cost associated with this life history shift. Consistent with the field studies, laboratory rearing experiments revealed no effects of fish cues on Ephemerella's morphological defences. These data suggest that variation in the density or species composition of predators may favour the evolution of developmental plasticity to reduce mortality in the larval environment.