Shift between carnivory and omnivory in stream stonefly predators

Omnivory can have profound effects on the trophic dynamics of communities and ecosystems, as they may interact with multiple trophic levels simultaneously. Some species of large-bodied stoneflies may be viewed as omnivores rather than true carnivores even at later nymphal stages. We evaluated the seasonal change in the diet of stonefly predators by analyzing their stable isotope ratio, gut contents, physiological activity, and food availability. A two-source-based mixing model based on stable isotope analysis revealed that stoneflies shifted their diet between carnivory in summer and omnivory in winter—despite the higher availability of animal prey in winter. The gut content analysis showed that swift prey (mayflies) were consumed in the summer, whereas sluggish prey (Chironomidae) were consumed in the winter. The physiological activity of stoneflies also declined markedly in winter. These results suggest that, in winter, stoneflies foraged on a mixture of Chironomidae and algae. It appears that omnivory in some stream consumers is related to the seasonal change in temperature-dependent physiological activity, rather than prey availability.     

Consequences of larval intraspecific competition to stonefly growth and fecundity

Summary We conducted experiments in replicated circular streams to measure the effect of intraspecific larval density on growth rates, size at emergence, timing of emergence, and fecundity of two species of predatory stoneflies (Megarcys signata and Kogotus modestus, Perlodidae). Early instars of both species showed no significant effect of intraspecific larval density on mean growth rates, despite the observation that in the absence of competitors stoneflies ate on average, significantly more prey (Baetis bicaudatus, Ephemeroptera, Baetidae) than in the presence of competitors. However, larval size of stoneflies held at higher densities (two per chamber) diverged over time, resulting in a greater size variability (coefficient of variation) among Kogotus than in treatments with low densities of stoneflies (one per chamber). The effect of doubling the density of early-instar Megarcys larvae was also asymmetrical, resulting in one larger and one smaller individual. In contrast, doubling the density of last-instar stoneflies whose feeding rates declined significantly prior to emergence had few measurable consequences, except that male Megarcys, which continued to feed throughout the last instar, had lower average feeding rates in high-density than in low-density chambers, and emerged at a significantly smaller mean size. We conclude that competition between early-instar stonefly larvae results in an asymmetry of body sizes, but that competitive effects are reduced as larvae slow or cease feeding before emergence. Since larger females of both stonefly species produced more eggs, the probable cost to females of early-instar larval competition was a reduction in their potential contribution of offspring to the next generation. The cost of attaining a smaller body size for male stoneflies is unknown; but if, as in many other insects, larger males have greater reproductive success, larval competition may increase the opportunity for sexual selection among males. This hypothesis remains to be tested experimentally.     

A new model estimating stonefly species production in the West Carpathian torrents

Samples of stoneflies were collected from 15 streams of Slovakia during 1976–2000. The model of growth rate is based on 219 data of 50 stonefly species. The non-linear relationship among growth rate (G) of stoneflies and monthly mean water temperature (T) and body mass (W) is described by equation: G = 0.014W ^−0.19 T ^0.25. The model estimates the species production of the families Taeniopterygidae, Nemouridae, Capniidae, Leuctridae and Perlodidae. This model is combined with Morin’s & Dumont’s (1994) model for Perlidae and Chloroperlidae. There is positive evidence that the total stonefly production of mountain and submountain streams increases with discharge. On the other hand, increasing altitude has a negative influence on production.     

Live cycles,drift, and standing stocks of some stoneflies (Insecta: Plecoptera) from streams in Minnesota,U.S.A.

From analyses of monthly length frequency distributions of stonefly (Plecoptera) nymphs from stream riffles, two types of univoltine life cycles (Hynes 1961, classification) were observed: slow seasonal forIsoperla transmarina, Isoperla signata, Isogenoides olivaceus, andAllocapnia rickerii, and fast seasonal forProstoia similis. The Perlodidae stoneflies (Isoperla andIsogenoides) showed greatest propensities to drift near the end of their life cycles. Drift rates of the other two species of stoneflies were correlated to standing stock estimates.     

Amphibiotic insects of mountain lakes and small watercourses in the Urals

A total of 127 species of stoneflies, dayflies, caddisflies, and chironomids were found in the lakes and small watercourses of the Northern and Pre-Polar Urals. The highest species diversity (78 species) and number were the characteristics of chironomids. Along with widely distributed species, Siberian and Arctic species were revealed in the fauna. The Arctic stonefly Capnia zaicevi Klap and 24 chironomid species were found for the first time on the territory of Komi Republic. Lists of registered insect species and data on their occurrence, abundance, and biomass are given.     

Global diversity of stoneflies (Plecoptera; Insecta) in freshwater

Plecoptera, or stoneflies, is a small order of hemimetabolous insects: according to our data, more than 3,497 species have been described so far in the world. The total number of species has enormously increased in the last 30 years (2,000 species estimated in 1976) and, if the trend continues, then it will nearly double in the near future. The order is divided into the suborders Arctoperlaria and Antarctoperlaria, and includes 16 families: 12 arctoperlarian and 4 antarctoperlarian. The Arctoperlaria account for a total number of 3,179 species, and Antarctoperlaria, only 318 species. The total number of genera is 286. We give in this article the estimated number of species for each family. The fauna and diversity of stonefly in North America (650 species reported) and Europe (426 species) are best known. Nevertheless, in the last 25 years, a mean of 2.6 Plecoptera species per year were described in Europe. Stonefly-faunas of Australia (191 species, Tasmania included) and New Zealand (104 species) are relatively well-known, while our knowledge of the Plecoptera of Central and South America (95 and 378 species respectively) is poor and still not representative of the real diversity. Africa has a reduced stonefly fauna (126 species). Asian stonefly diversity (approximately 1,527 species) is much greater than that of Europe or North America despite the fact that, except for Japan and Asiatic Russia that have been well studied, our knowledge of the remaining Asiatic areas is extremely poor. Even though our data indicate the Holarctic Region as the diversity hot-spot for the order, the analysis of the specific diversity divided by family suggests also an important role of tropical stoneflies. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

A laboratory study of predator-prey interactions of stoneflies and mayflies

SUMMARY. 1. Reactions of individual predatory stoneflies to contact by individual mayfly prey, and vice versa, were studied in the laboratory using two sets of species, one from a stream in southern Ontario. Canada, and the other from a stream in North Wales. 2. Based largely on information received via their antennae, the stoneflies were capable of distinguishing between conspecifics and mayfly prey and between prey species. A clear preference for certain prey species was evident but less preferred species were made more attractive when damaged or when the stoneflies were very hungry. Detection of conspecifics by stoneflies was confused in the presence of prey or the local smell of prey body fluids and this led to interference during hunting and feeding at high predator densities. 3. Some mayfly species were capable of distinguishing between conspecifics. other mayflies and predators. The cerci were important in intra- and interspecific contact encounters between mayflies and were used in a fencing fashion to establish a minimum distance between nymphs. However, the cerci did not appear to play an anti-predation role as stoneflies captured and ate mayflies with and without cerci in equal numbers. 4. Reactions to conspecifics by the Palaearetie mayflies Rhithrogena semicolorata (Curt.) and Baetis rhodani (Pictet) were similar to those of the ecologically-equivalent Nearctic species Stenonema vicarium (Walker) and Baetis vagans McDunnough. 5. In encounters between mayflies, intra- and interspecific responses were similar. However, the responses of some mayfly species to conspeeifics were modified in the physical presence of a predator or the water from its holding tank. Different mayfly species made specific responses to predaeeous stoneflies. Responses to a herbivorous stonefly were either similar to conspeeific encounters or were intermediate between them and encounters with predators. 6. Experiments to reveal the exact nature of the stonefly stimulus detected by the mayfly nymphs strongly suggested that there were two components to the stimulus. One was physical contact by the predator's antennae, the other was chemical in nature and could be detected only over short distances (up to a few mm). Independently each could elicit a response, but combined their effect appeared to be synergistic. 7. Different mayfly species responded differently to the stonefly Phasganophora capitata (Pictet) and this sequence of sensitivity was mirrored by the predator's choice of prey species—the most sensitive to the predator being the favoured prey and the less sensitive the least favoured. The significance of these findings to optimal foraging theory, as applied to tactile predators, is discussed.     

Stoneflies as ecological engineers – hungry predators reduce fine sediments in stream beds

1. We conducted experiments in a Colorado Rocky Mountain stream to measure the effect of foraging by predatory stoneflies ( Megarcys signata ) on fine sediment deposition and removal.
2. Cages containing one of four treatments were placed in the stream substratum and removed 3 days after fine sediment load to the stream was experimentally increased. Cages containing a stonefly but no prey accumulated less sediment than cages initially with no benthic invertebrates. Although cages with Megarcys plus prey also tended to have less sediment than controls, sediments were not reduced in cages with prey only.
3. Distance from sediment source, depth and current velocity at cages, final prey density and numbers of prey per predator gut at the end of the experiment had no effects on sediment accumulation in cages.
4. To determine the mechanisms underlying this effect, behavioural tests were conducted in a stream-side artificial stream system. To simulate the different hunger levels of stoneflies in cages, Megarcys were fed or starved for 3 days before behavioural trials that were repeated during high (night) and low (day) stonefly feeding periods. During night trials, foraging movements by starved Megarcys were more prolonged and active than those of fed stoneflies, regardless of the presence of prey. However, neither hunger level nor prey presence affected stonefly foraging behaviour during day trials.
5. Results of field experiments suggest that the presence of stoneflies enhances sediment removal from interstitial spaces. Behavioural observations indicate that nocturnal searching for prey by hungry Megarcys dislodges fine sediments from interstitial spaces.
6. Future studies should explore beyond the direct impacts of predators on stream invertebrate communities, and investigate the potential impact of predators on benthic microhabitat quality.     

Stoneflies as ecological engineers – hungry predators reduce fine sediments in stream beds

1. We conducted experiments in a Colorado Rocky Mountain stream to measure the effect of foraging by predatory stoneflies ( Megarcys signata ) on fine sediment deposition and removal.
2. Cages containing one of four treatments were placed in the stream substratum and removed 3 days after fine sediment load to the stream was experimentally increased. Cages containing a stonefly but no prey accumulated less sediment than cages initially with no benthic invertebrates. Although cages with Megarcys plus prey also tended to have less sediment than controls, sediments were not reduced in cages with prey only.
3. Distance from sediment source, depth and current velocity at cages, final prey density and numbers of prey per predator gut at the end of the experiment had no effects on sediment accumulation in cages.
4. To determine the mechanisms underlying this effect, behavioural tests were conducted in a stream-side artificial stream system. To simulate the different hunger levels of stoneflies in cages, Megarcys were fed or starved for 3 days before behavioural trials that were repeated during high (night) and low (day) stonefly feeding periods. During night trials, foraging movements by starved Megarcys were more prolonged and active than those of fed stoneflies, regardless of the presence of prey. However, neither hunger level nor prey presence affected stonefly foraging behaviour during day trials.
5. Results of field experiments suggest that the presence of stoneflies enhances sediment removal from interstitial spaces. Behavioural observations indicate that nocturnal searching for prey by hungry Megarcys dislodges fine sediments from interstitial spaces.
6. Future studies should explore beyond the direct impacts of predators on stream invertebrate communities, and investigate the potential impact of predators on benthic microhabitat quality.     

Can current velocity mediate trophic cascades in a mountain stream?

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Mountain stoneflies may tolerate warming streams: Evidence from organismal physiology and gene expression

Rapid glacier recession is altering the physical conditions of headwater streams. Stream temperatures are predicted to rise and become increasingly variable, putting entire meltwater‐associated biological communities at risk of extinction. Thus, there is a pressing need to understand how thermal stress affects mountain stream insects, particularly where glaciers are likely to vanish on contemporary timescales. In this study, we measured the critical thermal maximum (CT_MAX) of stonefly nymphs representing multiple species and a range of thermal regimes in the high Rocky Mountains, USA. We then collected RNA‐sequencing data to assess how organismal thermal stress translated to the cellular level. Our focal species included the meltwater stonefly, Lednia tumana, which was recently listed under the U.S. Endangered Species Act due to climate‐induced habitat loss. For all study species, critical thermal maxima (CT_MAX > 20°C) far exceeded the stream temperatures mountain stoneflies experience (<10°C). Moreover, while evidence for a cellular stress response was present, we also observed constitutive expression of genes encoding proteins known to underlie thermal stress (i.e., heat shock proteins) even at low temperatures that reflected natural conditions. We show that high‐elevation aquatic insects may not be physiologically threatened by short‐term exposure to warm temperatures and that longer‐term physiological responses or biotic factors (e.g., competition) may better explain their extreme distributions.     

Diet and predation by three leaf-associated stoneflies (Plecoptera) in an Arkansas mountain stream

SUMMARY. 1. Life cycles, food habits, mouthpart morphologies, prey preferences, and predator-prey size relationships were investigated for the three most common stoneflies in leaf packs in the Little Missouri River, Arkansas: Clioperla clio (Newman), Isoperla namata Prison, and Perlesta spp. Each species was univoltine and had a fast seasonal cycle, with extended egg or nymphal diapause, rapid nymphal growth and emergence in spring to early summer.
2. Foregut analysis showed predictable dietary shifts by C. clio and Perlesta spp., from detritus and diatoms in early instars to invertebrate prey in later instars. Isoperla namata fed facultatively on insects, detritus and diatoms, with no conspicuous ontogenetic shifts. The mouthparts of all three species strongly reflected their diet.
3. Electivity analysis indicated opportunistic feeding by the three species, with feeding rarely differing significantly from random.
4. Sizes of stonefly predators and their ingested prey were highly correlated ( P <0.01). However, rather than shifting toward larger prey sizes, growing predators expanded their size thresholds and continued to include numerous small prey items in their diet. Prey choice was governed most directly by prey availability, because stoneflies selected the most abundant groups, irrespective of size or taxon.
5. The likelihood of competition among the three stoneflies for prey was minimized by timing of their life cycles, differential food use and probably the temporary nature of leaf pack habitats.     

Filters and templates: stonefly (Plecoptera) richness in Ouachita Mountains streams, U.S.A.

1.  We collected adult stoneflies periodically over a 1-year period at 38 sites in two headwater catchments in the Ouachita Mountains, Arkansas, U.S.A. The 43 species collected were a subset of the Ozark-Ouachita fauna and the much larger fauna of the eastern U.S.A. We estimated 78–91% species coverage in the two catchments using jackknife extrapolation of species richness from our survey.
2.  Many streams, especially small ones, lacked surface water for months, but others, both small and large, flowed permanently.
3.  Using published regional presence–absence and coarse ecological data in a discriminant function analysis (DFA), we identified stream size (negative) and regional frequency of occurrence (positive) as predictors of presence in these headwater catchments. For the combined catchments, the extrapolated richness (51 spp.) was similar to an estimate (48 spp.) based on predicted absences from DFA and the Ouachita provincial total of known stonefly species (57 spp.).
4.  Local species richness (1–27 spp. per site) was correlated strongly with stream size (catchment area) but was independent of stream drying. Generic richness was correlated negatively with stream drying and positively, but less strongly, with stream size.
5.  Regionally endemic stoneflies dominated in drying streams, and widely distributed species dominated in more permanent streams. The composition of stonefly assemblages was associated with regional factors (species pools, regional abundance, evolution of tolerant endemic species, regional climate) and local factors (drying, stream size).     

Interactions of Adult Stoneflies (Plecoptera) with Riparian Zones II. Diet

The diets of three species of gripopterygid stonefly adults (Acroperla trivacuata, Zelandoperla decorata and Zelandobius furcillatus) from New Zealand were investigated using field studies and laboratory experiments. Guts of female stoneflies were generally fuller than those of males, suggesting that terrestrial feeding was particularly important for females. Adult A. trivacuata and Z. decorata collected from riparian vegetation had fuller guts than those collected from rocks, indicating that adults may move towards riparian vegetation to feed after emergence. Fungi and pollen formed the core food categories found in the guts of adult stoneflies collected from riparian vegetation. Stable isotope analyses of stoneflies and potential food sources collected from the near-stream riparian zone indicated that adult body carbon was derived primarily from autochthonous sources and/or from lichen growing on exposed stream channel rocks. The absence of mature females in our near-stream field collections may explain the lack of congruence between results of gut content and stable isotope analyses, as there would be a time lag between feeding following emergence and incorporation of any terrestrial carbon into body tissue. Assays for the presence of fructose were typically negative, even though pollen and sooty mould fungi were often ingested, probably because nectar or honeydew was rapidly assimilated after ingestion. Laboratory feeding trials indicated that adult Z. decorata lived significantly longer when fed a dilute sucrose solution than when fed a yeast extract solution or water. Our results demonstrate that terrestrial food sources are consumed by these adult gripopterygid stoneflies and that feeding is important for longevity. The provision of dietary requirements for the completion of adult aquatic insect life cycles may be an important consideration in riparian vegetation management. Dietary and assimilation studies of mature females which may move deep into riparian zones are required to resolve the strength of these linkages.     

Bill Ricker’s Entomological Contributions

Synopsis Bill Ricker was an internationally recognized expert on the stoneflies (Plecoptera). These insects have aquatic larvae that live mostly in cool and clear running water. They are of ecological significance in the breakdown of leaf-litter and the cycling of detritus in aquatic lotic habitats. Since they can account for a significant portion of the diet of some fishes in cold northern waters, they are linked with Ricker’s other work in fisheries. Within 1 year of taking up the study of stoneflies, Ricker was describing species new to science. He continued such activity through most of his life, describing or co-describing some 108 species and 46 genera. Ricker made other outstanding contributions, publishing in 1952 what is regarded as the most important publication dealing with stonefly systematics in the 20th century. Many names he coined in his studies clearly demonstrated his linguistic flair and humour. He made significant contributions to an understanding of the biogeography of stoneflies, and received many honours and citations for his research.     

Mechanisms of intra-and interspecific interference between larval stoneflies

Summary Behavior of focal individuals of two potentially competing sympatric stonefly species, Megarcys signata and Kogotus modestus (Perlodidae), was videotaped in flow-through plexiglass arenas placed in the East River, Gunnison County, Colorado. Focal individuals were observed alone and in pairs with conspecifics and allospecifics at four prey (Baetis bicaudatus, Baetidae, Ephemeroptera) densities to determine whether competitors and prey resource levels affected prey capture rates. Presence of conspecific or allospecific competitors reduced stonefly prey capture rates, especially for Kogotus, the smaller of the two species, due to a significant decline in predator-prey encounter rates with competitors present. This competitive effect was not observed at the lowest and highest prey densities due to very low or very high predator-prey encounter rates, respectively. Thus, interference affected feeding rates only at intermediate prey densities. Competitors had no effect on the probability of attacks per prey encounter or capture success per attack. Within each stonefly species the effects of intra-and interspecific interference on feeding rates were similar, even though behavioral responses by both stoneflies to interspecific encounters were more frequent than to encounters with conspecifics. Kogotus showed the highest levels of response to encounters with other stoneflies, maintaining those high levels of response to Megarcys over all prey densities. Further, male Kogotus, which are the smaller sex, responded more frequently to competitive interactions than did females. These data are consistent with the hypothesis that interspecific interference was asymmetrical with Megarcys, the larger species, being the superior competitor.     

Congruent population genetic structure but differing depths of divergence for three alpine stoneflies with similar ecology and geographic distributions

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Prey selection by a stonefly: the influence of hunger and prey size

Summary The influences of hunger and prey size on prey selection by the stonefly Hesperoperla pacifica (Perlidae) were explored in the laboratory by observing behavioral responses toward ten prey taxa and three nonprey taxa. Patterns of behavior were consistent with most assumptions and predictions of optimal foraging theory predicting sizebased prey selection by pursuing predators. Handling time appeared to increase as an exponential function of prey mass, and prey profitability (mg/s) was highest for small and intermediate-sized prey. Fasted stoneflies consumed a wide range of prey sizes, whereas well-fed stoneflies concentrated their attacks on intermediate-sized prey. Responses of H. pacifica to nonprey taxa, however, suggest that prey recognition and selection are not based on size alone.     

Interactions of Adult Stoneflies (Plecoptera) with Riparian Zones I. Effects of Air Temperature and Humidity on Longevity

The effects of constant air temperature and relative humidity on the longevity of three species of gripopterygid stonefly adults from New Zealand were investigated in laboratory experiments, and the results were compared to field measurements of air temperature and humidity obtained during summer. Greatest longevity for Zelandobius furcillatus, Zelandoperla decorata and Acroperla trivacuata was recorded in cool humid conditions (10°C, 100% humidity) for adults fed water and a 5% sucrose solution. Absence of feeding reduced survival by 37–73% at 17°C and 100% humidity. Survival decreased significantly with increasing constant air temperature (10, 17 and 25°C) and decreasing mean relative humidity (100, 81 and 15%). Males survived significantly longer than females in all temperature treatments for Z. furcillatus, but longevity was not influenced by gender in other species or in the humidity experiments. Interpolated LT 50 values over 96 h for female stoneflies in the temperature treatments averaged of 22–23°C. Field measurements at near-ground level and 1.5 m above the streambank during summer indicated that these air temperatures were exceeded for 25% of the time in a pasture catchment compared to &lt;0.1% of the time in a native forest catchment. These findings implicate air temperature as a factor potentially influencing the longevity of adult stoneflies, and suggest that maintenance of appropriate microclimate conditions should be a consideration in riparian management.     

Unidirectional prey-predator facilitation: apparent prey enhance predators' foraging success on cryptic prey

Food availability can strongly affect predator-prey dynamics. When change in habitat condition reduces the availability of one prey type, predators often search for other prey, perhaps in a different habitat. Interactions between behavioural and morphological traits of different prey may influence foraging success of visual predators through trait-mediated indirect interactions (TMIIs), such as prey activity and body coloration. We tested the hypothesis that foraging success of stream-dwelling cutthroat trout (Onchorhyncus clarki) on cryptically coloured, less-active benthic prey (larval mayfly; Paraleptophebia sp.) can be enhanced by the presence of distinctly coloured, active prey (larval stonefly shredder; Despaxia augusta). Cutthroat trout preyed on benthic insects when drifting invertebrates were unavailable. When stonefly larvae were present, the trout ate most of the stoneflies and also consumed a higher proportion of mayflies than under mayfly only treatment. The putative mechanism is that active stonefly larvae supplied visual cues to the predator that alerted trout to the mayfly larvae. Foraging success of visual predators on cryptic prey can be enhanced by distinctly coloured, active benthic taxa through unidirectional facilitation to the predators, which is a functional change of interspecific interaction caused by a third species. This study suggests that prey-predator facilitation through TMIIs can modify species interactions, affecting community dynamics.