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.     

Climate change effects on upland stream macroinvertebrates over a 25-year period

Climate change effects on some ecosystems are still poorly known, particularly where they interact with other climatic phenomena or stressors. We used data spanning 25 years (1981–2005) from temperate headwaters at Llyn Brianne (UK) to test three hypotheses: (1) stream macroinvertebrates vary with winter climate; (2) ecological effects attributable to directional climate change and the North Atlantic Oscillation (NAO) are distinguishable and (3) climatic effects on macroinvertebrates depend on whether streams are impacted by acidification. Positive (i.e. warmer, wetter) NAO phases were accompanied by reduced interannual stability (=similarity) in macroinvertebrate assemblage in all streams, but associated variations in composition occurred only in acid moorland. The NAO and directional climate change together explained 70% of interannual variation in temperature, but forest and moorland streams warmed respectively by 1.4 and 1.7°C (P<0.001) between 1981 and 2005 after accounting for NAO effects. Significant responses among macroinvertebrates were confined to circumneutral streams, where future thermal projections (+1, +2, +3°C) suggested considerable change. Spring macroinvertebrate abundance might decline by 21% for every 1°C rise. Although many core species could persist if temperature gain reached 3°C, 4–10 mostly scarce taxa (5–12% of the species pool) would risk local extinction. Temperature increase in Wales approaches this magnitude by the 2050s under the Hadley HadCM3 scenarios. These results support all three hypotheses and illustrate how headwater stream ecosystems are sensitive to climate change. Altered composition and abundance could affect conservation and ecological function, with the NAO compounding climate change effects during positive phases. We suggest that acidification, in impacted streams, overrides climatic effects on macroinvertebrates by simplifying assemblages and reducing richness. Climatic processes might, nevertheless, exacerbate acidification or offset biological recovery.     

Growth and emergence of the stonefly Leuctra nigra in coniferous forest streams with contrasting pH

1. Growth and emergence of the acidification tolerant stonefly Leuctra nigra were studied in four coniferous forest streams with contrasting pH. Invertebrate samples were taken bimonthly during May 1997 to April 1998. Emerging L. nigra were captured in two of the streams during April to July 1998 using pyramidal emergence traps. In addition, a laboratory experiment was conducted to evaluate growth of L. nigra in response to variations in food quality (neutral or acid conditioned leaves) and water pH. 2. The invertebrate community in low pH streams had lower diversity and density compared with neutral streams. Abundance of L. nigra was also lower, despite the dominance of this stonefly in the invertebrate community in the acidic streams. 3. Secondary production of L. nigra was 2.04 and 3.07 g DW m^–2 year^–1 in the neutral streams compared with 1.02 and 1.27 g DW m^–2 year^–1 in the acidic streams. The P/B ratio differed only slightly among streams. 4. More adult L. nigra emerged in the neutral stream compared with the acidic stream, despite the density of the pre‐emerging fifth instar nymphs of L. nigra being similar in the two streams. Adult size (expressed as wing length) and female fitness were also significantly greater in the neutral stream. 5. Laboratory growth of L. nigra nymphs was significantly lower when fed acid conditioned alder leaves rather than neutral conditioned leaves. There was no difference in growth when nymphs were fed neutral conditioned leaves in acid and neutral water. 6. Our study shows multiple, sub‐lethal effects on life history of the acidification tolerant stonefly L. nigra in coniferous streams with low pH. The results suggest that these effects could be mediated primarily by low food quality. These findings add to the existing evidence that coniferous plantation forestry has a negative impact on stream ecosystems.     

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.     

Use of stream order and biological indices to assess water quality in the Osage and Black river basins of Missouri

Significant variations in four biological measures of water quality with stream order and river basin were demonstrated for streams of the Black and Osage river basins of Missouri. Water quality criteria specific for each order and basin were then developed.Benthic macroinvertebrates from springs and stream orders 3–8 in the two river basins were sampled quarterly for one year with riffle nets and artificial substrate samplers. A total of 548 samples were taken at 137 stations. The average annual macroinvertebrate density, index of diversity, number of taxa, and number of mayfly and stonefly taxa were determined for each station. These measures showed significant differences (p < 0.05) across stream order within and between the two river basins. Total taxa, total mayfly and stonefly taxa, and diversity were highest in orders 4 and 5 with decreased values in lower and higher stream orders. Maximum organism densities occurred in intermediate order streams. These differences were attributed to the succession of physical changes from headwaters to mouth within each river and to the unique geomorphology of each catchment basin.Water quality criteria based on three of the four measures described above (with 95% confidence limits) were established for each stream order in each river basin. Criteria for the Osage River basin were then used to identify three streams in the basin affected by environmental disturbances (stream impoundment, channelization and sewage discharge). The use of order- and basin-specific criteria assures that the biological differences between streams caused by environmental disturbance can be distinguished from the natural biological differences between streams of different orders and drainages.     

Seasonal change in the gut clearance rate of three stonefly species in a Japanese stream

We measured seasonal changes in the gut clearance rate of three stonefly species in a stream. The rate declined with water temperature. Based on the gut clearance rate obtained here, we estimated the half-reduction time of gut contents and consumption rate of stoneflies on algae. Irrespective of stonefly species, the half-reduction time in winter (2 days) was twice as long as that in summer (1 day). The areal consumption rates of stoneflies varied depending on species and season. These results imply that the seasonal consumption impact of aquatic insects can be different, despite their daily ration being similar among species.     

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.     

Triclads (Turbellaria: Tricladida) as predators of lake-dwelling stonefly and mayfly nymphs

It has been demonstrated by using a serological technique that Polycelis nigra (Müller) and P. tennis (Tjima) feed on mayfly and stonefly nymphs in unproductive lakes. Such feeding is less in productive lakes where these insects form a smaller proportion of the invertebrate fauna of the stony littoral zone. These nymphs form a significantly larger part of the diet of P. nigra than of P. tenuis; both triclad species feed more on stoneflies than mayflies.     

Life cycle and production of Skwala parallela (Frison) (Plecoptera: Perlodidae) in a Colorado montane stream

The life cycle and production of Skwala parallela, a perlodid stonefly, was investigated in a third-order Colorado montane stream. The species exhibited a univoltine life cycle with a distinct cohort. Small nymphs appeared in May. Rapid growth was exhibited throughout summer and autumn. During winter, growth slowed somewhat but was continuous until April. Maximum density of 34 nymphs/m² occurred in July. Based upon the instantaneous growth method, annual production was 395.3 mg/m² or 3.95 kg/ha dry weight with a P/B ratio of 4.4     

Forests and the temperature of upland streams in Wales: a modelling exploration of the biological effects

SUMMARY. 1. Daily temperature data from six streams in upland Wales were used to explore the thermal effects of afforestation on stream ecology. The data were linked to published biological models to simulate fish and invertebrate development.
2. Mean daily temperatures in forest streams were lower than those of moorland streams in spring and summer, and higher in winter. These spatial comparisons were supported by the results of experimental bank-side clearance at a forest site, where there was evidence that stream temperatures fell in winter and rose in spring following treatment.
3. Simulations indicated that brown trout (Salmo trutta) could weigh over 30% more by the end of their second growing season in a moorland compared with a forest stream. Several species of insects showed slower simulated egg development at forest sites. For two ephemeropteran species simulated nymphal growth was also retarded, suggesting significant alterations to the life cycle. Two plecopteran species were affected only slightly by the different temperature regimes.
4. Overall, the simulations suggested that afforestation, by reducing summer temperatures, could lead to marked reductions in rates of development of some invertebrates and fish.     

Influences of the atmospheric patterns on unstable climate-growth associations of western Mediterranean forests

Quantifying climate-growth associations is needed to evaluate how forest productivity will respond to climate change. Year-to-year fluctuations in forest productivity and radial growth are partly explained by local climatic conditions driven by large-scale atmospheric patterns. This is illustrated by Iberian forests in the western Mediterranean Basin, which are subjected to complex climatic and atmospheric influences such as Atlantic and Mediterranean cyclogenesis. The North Atlantic Oscillation (NAO) is one of the major atmospheric circulation patterns affecting Iberian forests since positive winter NAO phases lead to dry and warm conditions. The Western Mediterranean Oscillation (WeMO) may also explain Iberian forest growth in some areas since this index captures Mediterranean cyclogenesis and WeMO negative phases are linked to warm and wet spring to summer conditions. Here, we analyze the associations between atmospheric patterns, climate and tree growth and we determine if they are changing through time. We use dendrochronology to relate radial growth of four tree species (Pyrenean oak, Sweet chestnut, Maritime pine and Scots pine) growing in western Spain to climate conditions and the NAO and WeMO indices. Winter and early spring temperatures increased since the 1950s in the area whereas the negative association between winter precipitation and the NAO strengthened since then. However, mean temperature rise was particularly evident since the 1970s. Growth was reduced by dry conditions during the growing season (spring and summer), but also by cold and dry conditions during the previous autumn and winter. This explains why the NAO January and the WeMo April indices were negative to growth of three species excluding Pyrenean oak. The early 1970s reflected an inflection point in the instability of climate-growth associations in the study area. We conclude that the winter NAO is a relevant driver of forest growth in the western Iberian Peninsula forests but additional atmospheric patterns (WeMO) also affect, albeit to a minor extent, these forests.     

A seasonal change in the distribution of a stream-dwelling stonefly nymph reflects oxygen supply and water flow

We examined the effects of oxygen availability, which has been viewed as a minor factor in streams, on the seasonal and spatial microhabitat distribution of a stonefly. Surveys were conducted in winter and summer in a mountain stream by collecting stones from the streambed and determining the presence or absence of the insect. At each stone sampling, we also measured physical conditions. The probability of the stonefly presence increased significantly with current velocity in summer, but not in winter. Because current influences oxygen renewal rates, our results suggest that the distribution of the insect could be restricted by oxygen.     

Ecology of streams draining forested and non-forested catchments in an area of central Scotland subject to acid precipitation

A study of 12 streams draining forested and non-forested catchments was made in an area of central Scotland where slow-weathering bedrock was predominantly quartzite, schists and slates. Sitka spruce (Picea sitchensis Carriere) was the most common tree species. Precipitation in the area had an annual mean pH in the range 4.3–4.5. Streams within the planted zone were always more acid than those outside and had higher concentrations of aluminium and manganese. With one exception, trout were absent from streams within long-established forests and planted salmon eggs (Salmo salar L.) died within a few weeks. A high proportion of such eggs survived in streams outside the forest. Siphlonurus lacustris Eaton was the only mayfly nymph found in the most acid streams in summer collections. In winter samples, mayfly nymphs, Heptagenia lateralis (Curtis) were found in only one forest stream but several species were present in the non-forested catchments. It is suggested that spruce forests can effectively collect acid pollutants which are subsequently washed out, thus accelerating the acidification of the soil. Streams therefore become increasingly acid as the neutralisation capacities of their catchments decrease.     

The effect of environmental factors on the mayfly communities of headwater streams in the Pieniny Mountains (West Carpathians)

A study of the species composition of mayfly communities in connection with environmental parameters was made in headwater streams of the Pieniny Mts. The rhithral zone is inhabited maximally by 19 mayfly species. In most of the streams studied the mayfly communities were found to be similar, however the vertical zonation which reflected human impact was visible (NMDS analysis). The main factors responsible for mayfly communities at all the sites studied were stream regulation and organic pollution, followed by type of bottom substrate (pebble and gravel), riparian vegetation (shrubs), pH and water temperature. At undisturbed sites the most important factors were pH, substrate type, distance from the source, current velocity and riparian vegetation (CCA analysis). Analysis of mayfly communities and environmental characteristics in different seasons showed that occurrence of mayfly species varied substantially depending on the season. Only in early spring and autumn do mayfly communities occur which are dependent on many environmental factors, the most significant of which are substrate type, phosphate, distance from source and altitude (CCA analysis).     

Effects of temperature on growth rate and behavior of Epeorus albertae (Ephemeroptera: Heptageniidae) nymphs

Anthropogenic disturbances affect temperature in river systems. Temperature potentially affects life histories of macroinvertebrates and alters behavior and biological functions. Temperature preferences and tolerance ranges for key taxa are therefore critical for understanding impacts of human-induced changes to water temperatures on river ecosystems. The objective of this study was to examine the effect of water temperature on growth rate and behavior of Epeorus albertae (McDunnough) nymphs. Nymphs were collected from the Umatilla River in eastern Oregon, and exposed to temperatures of 18, 22, and 28 °C. Nymphs held at 28 °C exhibited increased growth rates compared with individuals held at 18 and 22 °C. However, at 28 °C the accumulation of nymphal tissues was not consistent with that of nymphs held in lower temperatures; ratios of head capsule width to total body length were significantly lower in individuals at 28 °C compared with those held at the lower temperatures. This indicates that the nymphs held at the high temperature had longer total body length relative to the developmental stage, represented by head capsule width, when compared with insects in cooler temperatures. To examine the effect of water temperature on behavior, active drift of mayflies was examined in experimental chambers held at 12, 18, 22, and 28 °C. The number of drifting insects observed was significantly higher at 28 °C compared with 22, 18, and 12 °C. These results indicate that temperature is a factor influencing growth and behavior of E. albertae and is likely to lead to limitations in habitat use of this mayfly.     

Distribution Patterns,Species Assemblages and Habitat Selection of the Stoneflies (Plecoptera) from Two Mediterranean River Basins in Southern Spain

The stonefly distribution and its relationship with habitat changes, based on stream physico chemical features, were investigated in two Mediterranean river basins (Guadaira and Guadalete, S Spain). Plecopteran nymph distribution varied along an elevation gradient. Both stonefly species richness and species diversity increased with altitude. Stoneflies were able to tolerate a wide range of environmental conditions but were absent from low altitude sites, temporary streams with high salinity values and high water temperature. A total of 13 stonefly species were found in the study area, from which Tyrrhenoleuctra minuta (Klapálek ) and Protonemura sp. were dominant. The most widespread species were Nemoura lacustris and Capnioneura mitis. Three distinct species assemblages were observed according to their distribution patterns, but with similar environmental preferences. Capnioneura petitpierreae and T. minuta showed preferences to low altitude sites, whereas C. mitis and N. lacustris were found throughout all sites. T. minuta and N. lacustris inhabited permanent and temporary sites, while the same species and C. mitis also were found in semipermanent streams. Temporally, most stoneflies were caught during winter (85.9%), especially at the low altitude sites.     

The lateral extent of the subsidy from an upland stream to riparian lycosid spiders

Adult aquatic insects emerging from streams can subsidize riparian food webs, but little is known of the spatial extent of these subsidies. Stable isotope (¹⁵N) enrichment of aquatic insects, principally a species of stonefly (Plecoptera: Leuctridae), emerging from an upland stream was used to trace the subsidy from the stream ecosystem to riparian spiders (Lycosidae). The downstream profile of spider δ¹⁵N correlated closely with that of adult stoneflies, indicating that they were deriving nutrition from aquatic sources. The contribution of adult aquatic insects to spider diets was determined using a two-source mixing model. Adult aquatic insects made up over 40% of spider diets adjacent to the stream, but <1% at 20 m from the stream. Enrichment of riparian spiders declined exponentially with distance from the stream channel. Aquatic-terrestrial subsidies were spatially restricted, but locally important, to riparian lycosid spiders at the study site.     

Trematodes and nematodes parasitizing the benthic insect community of an Andean Patagonian stream,with emphasis on plagiorchiid metacercariae

In freshwater systems, parasitological studies have mainly been carried out on vertebrates and molluscs, but little is known about parasites of aquatic insects. We describe the trematodes and nematodes parasitizing the benthic insects of an Andean Patagonian stream and the presence of parasites in the terrestrial adult stages. Members of 3 of 20 insect taxa were found to be parasitized by larval nematodes, and members of six taxa harbored metacercariae of digeneans. In benthic samples, chironomids, simuliids (Order Diptera), and baetids (Order Ephemeroptera) harbored mermithid larvae (Nematoda). The stonefly Antarctoperla michaelseni (Order Plecoptera), the caddisfly Smicridea annulicornis (Order Trichoptera), a watersnipe fly (Order Diptera: Athericidae), and three species of leptophlebiid mayflies (Order Ephemeroptera) were parasitized by encysted plagiorchiid metacercariae (Order Plagiorchiida). Most metacercariae were found in the three species of mayflies with prevalences ranging 15–63% and mean intensities ranging 1.2–4.9. Prevalence declined from summer to early winter, probably because of the emergence of infected nymphs and the recruitment of uninfected new cohorts. The imagos had live metacercariae with higher prevalences and intensities of infection than nymphs. We suggest that these plagiorchiids have an allogenic life cycle, involving a terrestrial definitive host.     

Local and regional climate variables driving spring phenology of tortricid pests: a 36 year study

1. Insect phenology is driven by local climate variables, most notably temperature. Increased warming has been linked to advancements in critical phenophases such as the spring flight of reproductive adults in the mid‐Atlantic region of the U.S.A. 2. Local climate is governed by the fluctuations of large‐scale climate oscillations. In the northern hemisphere, both the North Atlantic Oscillation (NAO) and the Arctic Oscillation (AO) control the local autumn and winter severity. Low NAO and AO indices are associated with colder autumns and winters, which can delay spring phenology. 3. In this study, 36 years of data from experimental fruit orchards in Biglerville, Pennsylvania, were used to run partial least‐squares regressions in order to determine the climate variables related to the spring phenology of five tortricid pest species. 4. The phenology of the tortricid pests did not advance, even though there was evidence of warming at the research site. 5. Spring temperatures were found to be the most significant climate variables in determining the timing of the spring flights. However, autumn–winter temperatures were also important. 6. For the NAO and the AO, it was found that these oscillations affected the tortricid moths by influencing autumn–winter conditions. The oscillations of the NAO and AO can obscure long‐term changes in phenology. 7. These findings suggest that the inclusion of large‐scale climate oscillations can provide important insights into how climate conditions can influence insect phenology, and presents an opportunity for improving the ability to forecast spring emergence.