Social Insects Dominate Eastern US Temperate Hardwood Forest Macroinvertebrate Communities in Warmer Regions

Earthworms, termites, and ants are common macroinvertebrates in terrestrial environments, although for most ecosystems data on their abundance and biomass is sparse. Quantifying their areal abundance is a critical first step in understanding their functional importance. We intensively sampled dead wood, litter, and soil in eastern US temperate hardwood forests at four sites, which span much of the latitudinal range of this ecosystem, to estimate the abundance and biomass m⁻² of individuals in macroinvertebrate communities. Macroinvertebrates, other than ants and termites, differed only slightly among sites in total abundance and biomass and they were similar in ordinal composition. Termites and ants were the most abundant macroinvertebrates in dead wood, and ants were the most abundant in litter and soil. Ant abundance and biomass m⁻² in the southernmost site (Florida) were among the highest values recorded for ants in any ecosystem. Ant and termite biomass and abundance varied greatly across the range, from <1% of the total macroinvertebrate abundance (in the northern sites) to >95% in the southern sites. Our data reveal a pronounced shift to eusocial insect dominance with decreasing latitude in a temperate ecosystem. The extraordinarily high social insect relative abundance outside of the tropics lends support to existing data suggesting that ants, along with termites, are globally the most abundant soil macroinvertebrates, and surpass the majority of other terrestrial animal (vertebrate and invertebrate) groups in biomass m⁻². Our results provide a foundation for improving our understanding of the functional role of social insects in regulating ecosystem processes in temperate forest.     

Similar breakdown rates and benthic macroinvertebrate assemblages on native and Eucalyptus globulus leaf litter in Californian streams

1. Eucalyptus globulus, a tree species planted worldwide in many riparian zones, has been reported to affect benthic macroinvertebrates negatively. Although there is no consensus about the effects of Eucalyptus on aquatic macrobenthos, its removal is sometimes proposed as a means of ecological restoration. 2. We combined the sampling of macroinvertebrates with measurement of the colonisation of leaf packs in mesh bags, to examine the effects of riparian Eucalyptus and its litter on benthic macroinvertebrates in three small streams in California, U.S.A. Each stream included one reach bordered by Eucalyptus (E‐site) and a second bordered by native vegetation (N‐site). 3. The macrobenthos was sampled and two sets of litter bags were deployed at each site: one set with Eucalyptus litter (Euc‐bags) and one with mixed native tree litter (Nat‐bags) containing Quercus, Umbellularia, Acer and Alnus. Bags were exposed for 28, 56 and 90 days and this experiment was repeated in the autumn, winter and spring to account for effects of changing stream flow and insect phenology. 4. Litter input (average dry mass: 950 g m^?2 year^?1 in E‐sites versus 669 g m^?2 year^?1 in N‐sites) was similar, although in‐stream litter composition differed between E‐ and N‐sites. Litter broke down at similar rates in Euc‐bags and Nat‐bags (0.0193 day^?1 versus 0.0134 day^?1), perhaps reflecting the refractory nature of some of the leaves of the native trees (Quercus agrifolia). 5. Summary metrics for macroinvertebrates (taxon richness, Shannon diversity, pollution tolerance index) did not differ significantly between the E and N sites, or between Euc‐bags and Nat‐bags. No effect of exposure time or site was detected by ordination of the taxa sampled. However, distinct seasonal ordination clusters were observed in winter, spring and autumn, and one of the three streams formed a separate cluster. 6. The presence of Eucalyptus was less important in explaining the taxonomic composition of the macrobenthos than either ‘season’ or ‘stream’. Similarly, these same two factors (but not litter species) also helped explain the variation in leaf breakdown. We conclude that patches of riparian Eucalyptus and its litter have little effect on stream macrobenthos in this region.     

Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Effects of stream restoration and wastewater treatment plant effluent on fish communities in urban streams

1. Fish community characteristics, resource availability and resource use were assessed in three headwater urban streams in Piedmont North Carolina, U.S.A. Three site types were examined on each stream; two urban (restored and unrestored) and a forested site downstream of urbanisation, which was impacted by effluent from a wastewater treatment plant (WWTP). Stream basal resources, aquatic macroinvertebrates, terrestrial macroinvertebrates and fish were collected at each site. 2. The WWTPs affected isotope signatures in the biota. Basal resource, aquatic macroinvertebrate and fish δ¹⁵N showed significant enrichments in the downstream sites, although δ¹³C signatures were not greatly influenced by the WWTP. Fish were clearly deriving a significant part of their nutrition from sewage effluent‐derived sources. There was a trend towards lower richness and abundance of fish at sewage‐influenced sites compared with urban restored sites, although the difference was not significant. 3. Restored stream sites had significantly higher fish richness and a trend towards greater abundance compared with unrestored sites. Although significant differences did not exist between urban restored and unrestored areas for aquatic and terrestrial macroinvertebrate abundances and biotic indices of stream health, there appeared to be a trend towards improvements in restored sites for these parameters. Additional surveys of these sites on a regular basis, along with maintenance of restored features are vital to understanding and maximising restoration effectiveness. 4. A pattern of enriched δ¹³C in fish in restored and unrestored streams in conjunction with enriched δ¹³C of terrestrial invertebrates at these sites suggests that these terrestrial subsidies are important to the fish, a conclusion also supported by isotope cross plots. Furthermore, enriched δ¹³C observed for terrestrial invertebrates is consistent with some utilisation of the invasive C4 plants that occur in the urban riparian areas.     

Aquatic macroinvertebrate community structure of aNymphoides peltata-dominated and macrophyte-free site in an oxbow lake

The aquatic macroinvertebrates in two freshwater biotopes,viz. aNymphoides peltata-dominated site and a macrophyte-free site, were studied quantitatively in a shallow alkaline oxbow lake of the river Waal, the main branch of the river Rhine in The Netherlands. The research comprised the analysis of water, sediment and macrophyte samples.In the macrophyte-free site Oligochaeta and Nematocera, particularly of the collector gatherer functional feeding group, dominated the prevailing benthic community. The total macroinvertebrate biomass ranged here from 0.3 to 0.9 g ash-free dry weight per m² of biotope.Species richness, densities, and biomass of macroinvertebrates were considerably higher in the biotope dominated byNymphoides peltata. Many taxa were found associated with the aboveground macrophyte. The sediment compartment, however, contributed most to the total density and biomass of macroinvertebrates. Nematocera and Oligochaeta were the most abundant fauna groups, whereas the largest share in total biomass was provided by clams (Mollusca). The biomass of the total macroinvertebrate community in theNymphoides-dominated site ranged from 6.2 to 7.5 g ash-free dry weight per m² of biotope. The biomass of the aboveground phytophilous fauna ranged from 0.1 to 0.6 g ash-free dry weight per m² of biotope. In September, when theNymphoides peltata vegetation was in its senescent phase, the largest numbers and the highest biomass of phytophilous macroinvertebrates were observed. The contribution of the shredder functional feeding group was high in this period. This, and the overall high abundance of fauna with a detritivorous mode of life, indicates the importance of macrophyte detritus as input to food chains.     

Stream invertebrate diversity reduces with invasion of river banks by non‐native plants

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Food web structure and function in two arctic streams with contrasting disturbance regimes

1. We studied the effect of substratum movement on the communities of adjacent mountain and spring tributaries of the Ivishak River in arctic Alaska (69°1′N, 147°43′W). We expected the mountain stream to have significant bed movement during summer because of storm flows and the spring stream to have negligible bed movement because of constant discharge. 2. We predicted that the mountain stream would be inhabited only by taxa able to cope with frequent bed movement. Therefore, we anticipated that the mountain stream would have lower macroinvertebrate species richness and biomass and a food web with fewer trophic levels and lower connectance than the spring stream. 3. Substrata marked in situ indicated that 57–66% of the bed moved during summer in the mountain stream and 4–20% moved in the spring stream. 4. Macroinvertebrate taxon richness was greater in the spring (25 taxa) than in the mountain stream (20 taxa). Mean macroinvertebrate biomass was also greater in the spring (4617 mg dry mass m^?2) than in the mountain stream (635 mg dry mass m^?2). Predators contributed 25% to this biomass in the spring stream, but only 7% in the mountain stream. 5. Bryophyte biomass was >1000 times greater in the spring stream (88.4 g ash‐free dry mass m^?2) than the mountain stream (0.08 g ash‐free dry mass m^?2). We attributed this to differences in substratum stability between streams. The difference in extent of bryophyte cover between streams probably explains the high macroinvertebrate biomass in the spring stream. 6. Mean food‐web connectance was similar between streams, ranging from 0.18 in the spring stream to 0.20 in the mountain stream. Mean food chain length was 3.04 in the spring stream and 1.83 in the mountain stream. Dolly Varden char (Salvelinus malma) was the top predator in the mountain stream and the American dipper (Cinclus mexicanus) was the top predator in the spring stream. The difference in mean food chain length between streams was due largely to the presence of C. mexicanus at the spring stream. 7. Structural differences between the food webs of the spring and mountain streams were relatively minor. The difference in the proportion of macroinvertebrate biomass contributing to different trophic levels was major, however, indicating significant differences in the volume of material and energy flow between food‐web nodes (i.e. food web function).     

Chlorophyll‐a concentrations and macroinvertebrate declines coincide with the collapse of overwintering diving duck populations in a large eutrophic lake

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Macroinvertebrate colonization and biofilm development on leaves and wood in a boreal river

• 1 Biofilms, the accumulation of micro-organisms, exoenzymes, and detritus particles on submerged surfaces, may change the quality of leaves and wood as sources of food and/or habitat for invertebrates. We examined the relationship between macroinvertebrate assemblages and biofilm development on leaves and wood in a boreal river.
• 2 Arrays of white birch ice-cream sticks and sugar maple leaves were placed at fast- and slow-current sites. Samples were collected periodically, and assayed for microbial biomass (ATP, ergosterol, chlorophyll a) and macroinvertebrate colonizers.
• 3 Microbial biomass and macroinvertebrate density were consistently greater on wood than leaves. Taxon richness was similar for all substratum/current combinations, but taxon density (number of taxa m^?2) was greater on wood. Macroinvertebrate density and taxon richness correlated with all microbial indicators when data from both substrata were pooled.
• 4 Leaves did not support as great a density of invertebrates as wood perhaps because of their faster breakdown rate and truncated biofilm development. Greater stability and the potential for surface complexity may make wood a site of higher macroinvertebrate diversity than leaves.

     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Community–habitat relationships in coastal streams in Big Sur, California, USA: travertine influences macroinvertebrate abundance and community structure

Travertine deposition occurs in streams worldwide but its effects on stream communities are poorly understood. I sampled benthic macroinvertebrates, periphyton, and reach-scale environmental variables in coastal streams in Big Sur, central California, USA, to determine the specific effects of travertine that occurred at some sites as well as to provide a broader assessment of community–habitat relationships. Total density and biomass of macroinvertebrates varied 6- and 9-fold across sites, respectively, and chlorophyll a concentrations varied 10-fold, but invertebrate and periphyton abundances were not correlated. Baetis tricaudatus (Ephemeroptera), Simuliidae (Diptera), and Chironomidae (Diptera) dominated macroinvertebrate communities across all sites, although differences in the relative abundances of these and other taxa produced moderate variation in community structure among sites (Bray-Curtis similarity coefficients of 47–84). Variation in community structure was related to a number of habitat features, notably travertine but also including variables reflecting channel morphology, flow, substrate size, and riparian tree type. Median density and biomass of macroinvertebrates were more than twice as high at sites without travertine than sites with travertine. Taxa richness also was higher at sites without travertine, and community structure differed moderately between sites with and without travertine, although there were no particular assemblages associated with either group. Non-metric multidimensional scaling (MDS) and cluster analysis of similarities in community structure appeared to separate sites with either travertine or high fines from sites without those conditions. These results demonstrate that travertine can have strong effects on stream communities, and additional studies are needed to identify the full range of effects on ecosystems and to evaluate the potential consequences of travertine for conservation efforts such as biomonitoring programs and threatened species management. Handling editor: R. Bailey     

Does rapid glacial recession affect feeding habits of alpine stream insects?

1. Glacial retreat, accompanied by shifts in riparian vegetation and glacier meltwater inputs, alters the energy supply and trophic structure of alpine stream food webs. Our goal in this study was to enhance understanding of dietary niches of macroinvertebrates inhabiting different alpine streams with contrasting glacial and non‐glacial (groundwater, precipitation, snowmelt) water inputs in conjunction with seasonal and habitat‐specific variation in basal resource availability.
2. We measured a range of stream physico‐chemical attributes as well as carbon and nitrogen isotopes (δ¹³C, δ¹⁵N) of macroinvertebrates and primary food sources at seven sites across seasons within a Swiss glaciated catchment (Val Roseg) undergoing rapid glacial retreat (1–2 km between 1997 and 2014). Sampling sites corresponded to streams used in a previous (1997/1998) study within the same alpine catchment.
3. Physico‐chemical attributes showed wide variation in environmental conditions across streams and seasons. Significant correlation among physico‐chemical proxies of glacier meltwater (phosphate‐P, total inorganic carbon, conductivity, turbidity) and macroinvertebrate δ¹³C, δ¹⁵N, and size‐corrected standard ellipse area (a proxy for feeding niche width) values showed that the extent of glacial water input shapes the energy base among alpine streams. Feeding niche differences among common alpine stream insect taxa (Chironomidae, Baetidae, Heptageniidae) were not significant, indicating that these organisms probably are plastic in feeding behaviour, opportunistically relying on food resources available in a particular stream and season.
4. Seasonal trends in macroinvertebrate δ¹³C largely followed patterns in periphyton δ¹³C values, indicating that autochthonous resources were the main consumer energy source within the stream network, as shown previously. The overall range in macroinvertebrate δ¹³C (?33.5 to ?18.4‰) and δ¹⁵N (?6.9 to 6.7‰) values also corresponded to values measured in the previous study, suggesting that macroinvertebrates altered diets in line with changes in environmental conditions and food resources during a period of rapid glacial retreat. Our results suggest that environmental changes brought on by rapid glacial retreat have not yet caused a profound change in the trophic structure within these fluvial networks.

     

Selective foraging by the crayfish Orconectes virilis and its impact on macroinvertebrates

SUMMARY. 1. The impact of crayfish predation on the abundance of macroinvertebrates was examined under semi-natural conditions. Female (Experiment 1) or male (Experiment 2) crayfish (Orconectes virilis) were held for 5 weeks in twelve small pools (4.67 m² surface area) at biomasses of 0. 5, 10 or 18 g m^?2 (live weight). The pools were stocked with known densities of macroinvertebrates. 2. Crayfish significantly affected the abundance of macroinvertebrates in the pools. Differences in the effects of crayfish on macroinvertebrates were related to crayfish sex, the presence of age-0 crayfish, and the species of macroinvertebrate. 3. The abundance of snails (Stagnicola elodes and Physa gyrina) was greatly reduced, in comparison with controls, by biomass of female crayfish ≥10 g m^?2 and by biomasses of male crayfish ≥5 g m^?2. The total density of non-molluscan invertebrates was inversely correlated with the biomass of female crayfish but the total biomass of non-molluscan invertebrates did not differ between treatments. This is consistent with our observation that small invertebrates (<2 mg wet weight) were less numerous, and large amphipods (32–64 mg) were more numerous, in pools stocked with female crayfish. In contrast, male crayfish had little apparent effect on the abundance of non-molluscan invertebrates. 4. Age-0 crayfish hatched at the end of Experiment 1 and were present in each pool at the start of Experiment 2. Surprisingly, male crayfish preyed little on age-0 crayfish. At the end of Experiment 2, the densities of age-0 crayfish varied between six and 116 individuals m^?2 and there was a strong inverse correlation between the mean biomass and density of age-0 crayfish recovered from the pools. This suggests age-0 crayfish were food limited in the pools and may explain the dominance of oligochaetes (which largely escape predation by burrowing) in the invertebrate community at the end of Experiment 2. 5. These results indicate that even relatively low densities of crayfish could greatly affect the abundance of macroinvertebrates in lakes. The introduction of crayfish into lakes (most lakes in Alberta currently have no crayfish) could substantially affect abundance and species composition of the macroinvertebrate community and, ultimately, the fish populations.     

The Effect of Forest Type on Benthic Macroinvertebrate Structure and Ecological Function in a Pine Plantation in the North Carolina Piedmont

We examined the impact of small-scale commercial forestry on the structure and function of 6 headwater streams in the North Carolina Piedmont. During 2001–2003 terrestrial organic matter inputs, temperature, macroinvertebrate community composition and tolerance, leaf breakdown rate, and food web structure were quantified for 2 streams draining mature stands of managed loblolly pine, 2 streams draining mature hardwood forests, and 2 streams draining 3-year-old clear cuts, which had been replanted with loblolly pine. Streams in the clear-cuts and pine plantations were bordered by a 15 m hardwood buffer. Despite differences in watershed land-use, there were no significant differences in the organic matter supply or temperature between streams draining different forest types. However, algal biomass was significantly higher in clear-cut sites than forested sites, and was also higher in hardwood sites than pine sites. Streams draining the clear-cut sites contained lower macroinvertebrate richness and diversity, and fewer intolerant species, than streams draining pine and hardwood stands. Despite the differences in macroinvertebrates community composition, there was no difference among forest types in leaf-pack breakdown rates. Analysis of δ¹⁵N and δ¹³C natural abundance of functional feeding group indicated that the shredders and predators collected from streams draining clear-cuts had a δ¹⁵N value that was enriched relative to the macroinvertebrates of streams draining pine and hardwood forests. This difference in δ¹⁵N signature appears to be the result of the incorporation of riparian grass species in the clear-cuts, which have a higher δ¹⁵N, into the diet of shredders. Pine sites had similar food webs to natural hardwood sites. Our results suggest that clear-cutting changes both the trophic dynamics and macroinvertebrate composition of low-order Piedmont streams in North Carolina despite the presence of hardwood buffers. However, large differences were not found between older pine and hardwood stands, indicating rapid recovery following re-growth of forest vegetation, when hardwood buffer strips were present.     

The Structure of a Macroinvertebrate Community in a Northern German Lake Outlet (Lake Belau,Schleswig-Holstein) with Special Emphasis on Abundance,Biomass and Secondary Production

The structure of macroinvertebrate communities was studied at I I sampling sites of the outlet of Lake Belau in the lowlands of northern Germany. To describe the structures of macrobenthic animal communities three different units were examined: abundance, biomass, and secondary production. 112 taxa were collected from the entire stream. The numbers of species ranged from 31 (fine sand) to 70 (submerged macrophytes). For the stream, average macroinvertebrate density was 18.400 ind. M⁻¹. Density was highest at the macrophytes amounting to 35,630 individuals per m², and lowest in the pure sand with only 3,900 ind. M⁻². Average biomass (dry mass) was 194 g DM m⁻² varying from 9.8 (peat) to 381 g DM m⁻² (gravel with mollusk shells near the upstream lake). For the stream, average annual production was 129 g DM m⁻² varying from 15 (peat) to 286 g DM m⁻² (macrophytes). The highest values for each unit were found in stream sections with gravel and submerged macrophytes. Lower values occured in sections that contained peat and sand. Usually, a single structure of the macroinvertebrate community was dominated by less than ten taxa, which varied at each sampling site depending on the units observed.     

Correlates of habitat favourability for benthic macroinvertebrates at five stream sites in an Appalachian Mountain drainage basin, U.S.A.

SUMMARY.

• 1 Based on monthly samples taken over a 1-year period, average density (individuals m^-2). average standing biomass and annual production of benthic macroinvertebrates were estimated at five sites within an Appalachian Mountain drainage basin. Two sites were on first order streams and differed from the three second order sites: they were smaller and more shallow and they were depressed in pH and chemical richness.
• 2 Patterns of abundance of individual taxa, of higher taxonomic groups and of functional (feeding) groups differed according to whether abundance was measured as density, as standing biomass or as annual production. Standing biomass was chosen as the measure of macroinvertebrate abundance because available evidence indicates that only standing biomass is consistently, positively correlated with survivorship, and thus with habitat favourability.
• 3 Two non-insect taxa (the crayfish Cambarus and the snail Leptoxis carinata) dominated standing biomass at each site. Consequently, differences among sites in total macroinvertebrate standing biomass and differences within and among sites in standing biomass of functional groups were determined by differences in estimated standing biomass of these two taxa. Differences in estimates of crayfish standing biomass were consistent with an explanation based on the availability of refuges created by large substrate particles. The abundance of L. carinata appeared to be controlled primarily by water chemistry and possibly secondarily by predators. A number of insect taxa exhibited patterns of standing biomass consistent with hypotheses based on effects of annual depth-flow regimes. Hypotheses based on differences in food resource and on competition appeared, in general, to be inconsistent with observed patterns of macroinvertebrate abundance.
• 4 Contrary to predictions of the River Continuum Concept, the shredder functional group in the Guys Run drainage and in other temperate woodland streams was found to be a minor part of total macroinvertebrate standing biomass. Further, in a majority of small forested stream sites studied to date, standing biomass of grazers has been determined to be greater than that of shredders.

     

Macroinvertebrate Responses to Algal and Bacterial Manipulations in Streams

Our study was designed to assess the relative importance of algae and bacteria as sources of energy for stream macroinvertebrates. In one experiment, we manipulated algae by artificially shading six sections in each of two streams, one stream with an open canopy (clear-cut drainage basin) and the other with a closed canopy (forested drainage basin); both streams were in Hubbard Brook Experimental Forest, New Hampshire, USA. Chlorophyll a concentrations were reduced from 0.2 to 0.05 μg/cm² in artificially shaded sections of both streams. However, macroinvertebrates showed no response to these algal manipulations in either the clear-cut or forested stream. Nutrient concentrations (N and P) were low and limiting to primary production in both the clear-cut and forested streams. Additionally, both streams had relatively low macroinvertebrate densities suggesting bottom-up controls were important in macroinvertebrate abundance. However, the forested stream did have higher macroinvertebrate densities presumably because of higher inputs of coarse particulate organic matter from the riparian vegetation. In a second experiment, in Augusta Creek, Michigan, we manipulated both algae and bacteria. To reduce algae, we artificially shaded experimental stream channels so that chlorophyll a was reduced from natural levels of 3.0–5.6 to 0.4–0.7 μg/cm². Half of the shaded channels had dissolved organic carbon (DOC – sucrose) dripped into them to raise DOC levels by 2–3 mg/l and thus stimulate bacterial abundance. Open channels, with higher algal abundance, had higher densities of Ephemerella, but only in November when nymphs were larger. Channels with increased DOC had higher bacterial abundances, higher densities of Chironomidae and lower densities of Heptageniidae. Several other macroinvertebrate taxa that were at relatively low abundance in our samples showed no significant response to these manipulations. Our results suggest that early instar Ephemerella may not rely as heavily on algae as later instars. Also, certain taxa were able to use the heterotrophic microbial community, especially chironomids which increased in numbers when bacterial density increased; thus, the bacterial carbon source may be more important to some stream macroinvertebrates than previous studies have suggested.     

Two lowland stream riffles – linkages between physical habitats and macroinvertebrates across multiple spatial scales

The physical structure of two riffles in a lowland Danish stream was studied and its importance for the composition and density of the macroinvertebrate communities was evaluated. The two riffles were visually assessed to be very similar, but measurements revealed that they differed in overall hydraulic conditions, stability, substratum composition and consolidation. Differences affected abundance of both burrowing and surface dwelling macroinvertebrates. The unstable unconsolidated riffle had higher total macroinvertebrate abundance (4137 m⁻² vs. 1698 m⁻²), diptera abundance (2329 m⁻² vs. 386 m⁻²) and total estimated species richness (31.7 vs. 28.8) as well as lower evenness (0.77 vs. 0.83) than the compact riffle. Among samples within the unconsolidated riffle, variations in macroinvertebrate communities were related to differences in mean substratum particle size. Here a linear log–log relationship existed between macroinvertebrate abundance, the abundance of EPT taxa and the median particle size (r ² _total = 0.46, p = 0.002; r ² _EPT = 0.73, p < 0.001). No similar relationships were evident on the consolidated riffle. Moreover, macroinvertebrate communities on the unconsolidated riffle were dominated by species with a high colonising potential. Despite being assessed to the same morphological unit, physical variation between riffles was surprisingly high as the riffles differed substantially with respect to consolidation, substratum heterogeneity and overall hydraulic structure. Macroinvertebrate community structure and composition also differed between riffles despite being drawn from the same species pool. The findings address the question if we use the correct methods and parameters when assessing the macroinvertebrate communities at the scale of the morphological unit.     

Adapting streams for climate change using riparian broadleaf trees and its consequences for stream salmonids

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Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.