Predation risk and resource abundance mediate foraging behaviour and intraspecific resource partitioning among consumers in dominance hierarchies

Dominance hierarchies and the resulting unequal resource partitioning among individuals are key mechanisms of population regulation. The strength of dominance hierarchies can be influenced by size‐dependent tradeoffs between foraging and predator avoidance whereby competitively inferior subdominants can access a larger proportion of limiting resources by accepting higher predation risk. Foraging‐predation risk tradeoffs also depend on resource abundance. Yet, few studies have manipulated predation risk and resource abundance simultaneously; consequently, their joint effect on resource partitioning within dominance hierarchies are not well understood. We addressed this gap by measuring behavioural responses of masu salmon Oncorhynchus masou ishikawae to experimental manipulations of predation risk and resource abundance in a natural temperate forest stream. Responses to predation risk depended on body size and social status such that larger fish (often social dominants) exhibited more risk‐averse behaviour (e.g. lower foraging and appearance rates) than smaller subdominants after exposure to a simulated predator. The magnitude of this effect was lower when resources were elevated, indicating that dominant fish accepted a higher predation risk to forage on abundant resources. However, the influence of resource abundance did not extend to the population level, where predation risk altered the distribution of foraging attempts (a proxy for energy intake) from being skewed towards large individuals to being skewed towards small individuals after predator exposure. Our results imply that size‐dependent foraging–predation risk tradeoffs can weaken the strength of dominance hierarchies by allowing competitively inferior subdominants to access resources that would otherwise be monopolized.     

Alpha and beta diversity of connected benthic–subsurface invertebrate communities respond to drying in dynamic river ecosystems

Drying disturbances are the primary determinant of aquatic community biodiversity in dynamic river ecosystems. Research exploring how communities respond to disturbance has focused on benthic invertebrates in surface sediments, inadequately representing a connected community that extends into the subsurface. We compared subsurface and benthic invertebrate responses to drying, to identify common and context‐dependent spatial patterns. We characterized community composition, alpha diversity and beta diversity across a gradient of drying duration. Subsurface communities responded to drying, but these responses were typically less pronounced than those of benthic communities. Despite compositional changes and in contrast to reductions in benthic alpha diversity, the alpha diversity of subsurface communities remained stable except at long drying durations. Some primarily benthic taxa were among those whose subsurface frequency and abundance responded positively to drying. Collectively, changing composition, stable richness and taxon‐specific increases in occurrence provide evidence that subsurface sediments can support persistence of invertebrate communities during drying disturbances. Beta‐diversity patterns varied and no consistent patterns distinguished the total diversity, turnover or nestedness of subsurface compared to benthic communities. In response to increasing drying duration, beta diversity increased or remained stable for benthic communities, but remained stable or decreased for subsurface communities, likely reflecting contrasts in the influence of mass effects, priority effects and environmental filtering. Dissimilarity between subsurface and benthic communities remained stable or increased with drying duration, suggesting that subsurface communities maintain distinct biodiversity value while also supporting temporary influxes of benthic taxa during drying events. As temporary rivers increase in extent due to global change, we highlight that recognizing the connected communities that extend into the subsurface sediments can enable holistic understanding of ecological responses to drying, the key determinant of biodiversity in these dynamic ecosystems.     

Stream-bank shade and larval distribution of the Philippine malaria vector Anopheles flavirostris

The principal malaria vector in the Philippines, Anopheles flavirostris (Ludlow) (Diptera: Culicidae), is regarded as 'shade-loving' for its breeding sites, i.e. larval habitats. This long-standing belief, based on circumstantial observations rather than ecological analysis, has guided larval control methods such as 'stream-clearing' or the removal of riparian vegetation, to reduce the local abundance of An. flavirostris. We measured the distribution and abundance of An. flavirostris larvae in relation to canopy vegetation cover along a stream in Quezon Province, the Philippines. Estimates of canopy openness and light measurements were obtained by an approximation method that used simplified assumptions about the sun, and by hemispherical photographs analysed using the program HEMIPHOT. The location of larvae, shade and other landscape features was incorporated into a geographical information system (GIS) analysis. Early larval instars of An. flavirostris were found to be clustered and more often present in shadier sites, whereas abundance was higher in sunnier sites. For later instars, distribution was more evenly dispersed and only weakly related to shade. The best predictor of late-instar larvae was the density of early instars. Distribution and abundance of larvae were related over time (24 days). This pattern indicates favoured areas for oviposition and adult emergence, and may be predictable. Canopy measurements by the approximation method correlated better with larval abundance than hemispherical photography, being economical and practical for field use. Whereas shade or shade-related factors apparently have effects on larval distribution of An. flavirostris, they do not explain it completely. Until more is known about the bionomics of this vector and the efficacy and environmental effects of stream-clearing, we recommend caution in the use of this larval control method.     

Nitrogen Export from Forested Watersheds in the Oregon Coast Range: The Role of N<Subscript>2</Subscript>-fixing Red Alder

Variations in plant community composition across the landscape can influence nutrient retention and loss at the watershed scale. A striking example of plant species importance is the influence of N₂-fixing red alder (Alnus rubra) on nutrient cycling in the forests of the Pacific Northwest. To understand the influence of red alder on watershed nutrient export, we studied the chemistry of 26 small watershed streams within the Salmon River basin of the Oregon Coast Range. Nitrate and dissolved organic nitrogen (DON) concentrations were positively related to broadleaf cover (dominated by red alder: 94% of basal area), particularly when near-coastal sites were excluded (r ² = 0.65 and 0.68 for nitrate-N and DON, respectively). Nitrate and DON concentrations were more strongly related to broadleaf cover within entire watersheds than broadleaf cover within the riparian area alone, which indicates that leaching from upland alder stands plays an important role in watershed nitrogen (N) export. Nitrate dominated over DON in hydrologic export (92% of total dissolved N), and nitrate and DON concentrations were strongly correlated. Annual N export was highly variable among watersheds (2.4–30.8 kg N ha^–1 y^–1), described by a multiple linear regression combining broadleaf and mixed broadleaf–conifer cover (r² = 0.74). Base cation concentrations were positively related to nitrate concentrations, which suggests that nitrate leaching increases cation losses. Our findings provide evidence for strong control of ecosystem function by a single plant species, where leaching from N saturated red alder stands is a major control on N export from these coastal watersheds.     

Parallels and contrasts in the effects of drought on stream macroinvertebrate assemblages

• 1 It is axiomatic that unusually long dry periods (droughts) adversely affect aquatic biota. Recovery after drought is rapid by macroinvertebrates that possess strategies to survive drying or are highly mobile but other taxa take longer to recolonise depending on the timing, intensity, and duration of the dry phase.
• 2 Although drought acts as a sustained ‘ramp’ disturbance, impacts may be disproportionately severe when certain critical thresholds are exceeded. For example, ecological changes may be gradual while a riffle dries but cessation of flow causes abrupt loss of a specific habitat, alteration of physicochemical conditions in pools downstream, and fragmentation of the river ecosystem. Many ecological responses to drought within these habitats apparently depend on the timing and rapidity of hydrological transitions across these thresholds, exhibiting a ‘stepped’ response alternating between gradual change while a threshold is approached followed by a swift transition when a habitat disappears or is fragmented.
• 3 In two Australian intermittent streams, drought conditions eliminated or decimated several groups of macroinvertebrates, including atyid shrimps, stoneflies and free‐living caddisflies. These taxa persisted during the early stages of the drought but did not recruit successfully the following year, despite a return to higher‐than‐baseflow conditions. This ‘lag effect’ in response to drought emphasises the value of long‐term survey data. Although changes in faunal composition were inconsistent among sites, marked shifts in taxa richness, abundance and trophic organisation after the riffle habitat dried provide evidence for a stepped response.
• 4 Responses by macroinvertebrate assemblages to droughts of differing severity in English chalk streams were variable. The prolonged 1988–92 drought had a greater impact than shorter droughts in the early 1970s but recovery over the next 3 years was swift. Effects of the 1995 summer drought were buffered by sustained groundwater discharge from the previous winter. These droughts tended to reduce available riverine habitats, especially via siltation, but few taxa were eliminated because they could recolonise from perennial sections of the chalk streams.
• 5 In the contrasting environments of the intermittent streams studied in England and Australia, there are parallels in the rapid rates of recolonisation. However, recruitment by taxa that lack desiccation‐resistant stages or have limited mobility is delayed. Currently, long‐term data on these systems may be insufficient to indicate persistent effects of droughts or predict the impacts of excessive surface or groundwater abstraction or the increased frequency and duration of droughts expected with global climate change.

     

Nitrogen uptake and transformation in a midwestern U.S. stream: A stable isotope enrichment study

This study presents a comprehensive analysis ofnitrogen (N) cycling in a second-order forestedstream in southern Michigan that has moderatelyhigh concentrations of ammonium (mean,16 g N/L) and nitrate (17 g N/L). Awhole-stream ¹⁵NH₄ ⁺ addition wasperformed for 6 weeks in June and July, and thetracer ¹⁵N was measured downstream inammonium, nitrate, and detrital and livingbiomass. Ancillary measurements includedbiomass of organic matter, algae, bacteria andfungi, nutrient concentrations, hydrauliccharacteristics, whole-stream metabolism, andnutrient limitation assays. The resultsprovide insights into the heterotrophic natureof woodland streams and reveal the rates atwhich biological processes alter nitrogentransport through stream systems.Ammonium uptake lengths were 766–1349 m anduptake rates were 41–60 g N m^–2min^–1. Nitrate uptake could not bedetected. Nitrification rates were estimatedfrom the downstream increase in¹⁵N-enriched nitrate using a simulationmodel. The ammonium was removed bynitrification (57% of total uptake),heterotrophic bacteria and fungi associatedwith detritus (29%), and epilithic algae(14%). Growth of algae was likely limited bylight rather than nutrients, and dissolvedO₂ revealed that the stream metabolism washeterotrophic overall (P:R = 0.2). Incubationsof detritus in darkened chambers showed thatuptake of ¹⁵N was mostly heterotrophic.Microbial N in detritus and algal N inepilithon appeared to reach isotopic steadystate with the dissolved ammonium, but theisotopic enrichment of the bulk detritus andepilithon did not approach that of ammonium,probably due to a large fraction of organic Nin the bulk samples that was not turning over. The actively cycling fraction of total N inorganic compartments was estimated from theisotopic enrichment, assuming uptake ofammonium but not nitrate, to be 23% forepilithon, 1% for fine benthic organic matter,5% for small woody debris, and 7% for leaves. These percentages agree with independentestimates of epilithic algal biomass, whichwere based on carbon:chlorophyll ratios in bulksamples and in algal fractions separated bydensity-gradient centrifugation in colloidalsilica, and of microbial N in the detritus,which were based on N released by chloroformfumigations.     

Interregional comparisons of sediment microbial respiration in streams

• 1 The rate of microbial respiration on fine‐grained stream sediments was measured at 371 first to fourth‐order streams in the Central Appalachian region (Maryland, Pennsylvania, Virginia, and West Virginia), Southern Rocky Mountains (Colorado), and California's Central Valley in 1994 and 1995.
• 2 Study streams were randomly selected from the United States Environmental Protection Agency's (USEPA) River Reach File (RF3) using the sample design developed by USEPA's Environmental Monitoring and Assessment Program (EMAP).
• 3 Respiration rate ranged from 0 to 0.621 g O₂ g^‐1 AFDM h^‐1 in Central Appalachian streams, 0‐0.254 g O₂ g^‐1 AFDM h^‐1 in Rocky Mountain streams, and 0‐0.436 g O₂ g^‐1 AFDM h^‐1 in Central Valley streams.
• 4 Respiration was significantly lower in Southern Rocky Mountain streams and in cold water streams (< 15 °C) of the Central Appalachians.
• 5 Within a defined index period, respiration was not significantly different between years, and was significantly correlated with stream temperature and chemistry (DOC, total N, total P, K, Cl, and alkalinity).
• 6 The uniformity of respiration estimates among the three study regions suggests that sediment microbial respiration may be collected at any number of scales above the site‐level for reliable prediction of respiration patterns at larger spatial scales.

     

Species richness and assemblage structure of Trichoptera in Danish streams

• 1 Species richness and assemblages of Trichoptera from 157 ‘pristine’ Danish lowland stream sites were analyzed, for dependence on geographical position of the sites and simple physical variables, using two way indicator species analysis and detrended correspondence analysis.
• 2 A total of 106 species were recorded, representing ≈ 90% of the species pool known from Danish streams. Only seven species occurred at more than half the sites, whereas an additional 11 species were found at more than a quarter of the sites.
• 3 Although sites showed significant regional differences in environmental variables (stream order, width, slope and presence/absence of riparian forest), species richness and assemblages were primarily correlated with stream order, width and slope. Maximum richness was found at the largest (5th order) stream sites.
• 4 Regional differences in species assemblages were found, with several species being absent from the islands Funen and Bornholm. Species assemblages also differed between forested and non‐forested stream sites.
• 5 We found evidence that stream temperature may be of only minor importance in determining Trichoptera species richness and assemblage composition in Danish streams compared to other size‐related physical factors.

     

Canonical variables of aquatic bryophyte combinations for predicting water trophic level

Concentrations of soluble reactive phosphorus (SRP), nitrate, and soluble reactive silicon (SRSi) were monitored in 12 streams draining small catchments (<10 km2) in the English Lake District. The catchments varied with respect to underlying geology, soil type and land cover. Average concentrations of SRP were in the range 0.5–11.2 μg P l-1, and estimated loads ranged from 0.01 to 0.14 kg P ha-1 a-1. The higher concentrations and loads were associated with catchments containing improved pasture. Mean streamwater concentrations of nitrate varied from 55 to 660 μg N l-1, while loads were in the range 0.8–9.6 kg N ha-1 a-1; no general dependence on catchment properties was discerned. Concentrations of SRSi were similar in all the streams (0.8–2 mg Si l-1), and annual loads were in the range 10–26 kg Si ha-1 a-1. Loads of all three nutrients were greatest during the winter, because of higher discharges, but in some catchments containing improved pasture, considerable transport of P also took place during the summer. Concentrations of nitrate in streams draining unimproved moorland catchments are approximately twice those reported for samples taken from similar streams in 1973 and 1974, possibly because of increased atmospheric deposition of inorganic nitrogen (ammonium and nitrate). Concentrations of SRP in such streams were similar to those reported for the earlier samples. Comparisons of stream loads of SRP and nitrate with estimated inputs suggest that catchment soils retain substantial amounts of these nutrients. Implications for surface water eutrophication of changes in P retention by soils are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.