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  1. Abstract

    Ecosystem engineers modify habitats in ways that facilitate other community members by ameliorating harsh conditions. The strength of such facilitation is predicted to be influenced by both beneficiary traits and abiotic context. One key trait of animals that could control the strength of facilitation is beneficiary body size because it should determine how beneficiaries fit within and exploit stress ameliorating habitat modifications. However, few studies have measured how beneficiary body size relates to facilitation strength along environmental gradients.

    We examined how the strength of facilitation by net‐spinning caddisflies on invertebrate communities in streams varied along an elevation gradient and based on traits of the invertebrate beneficiaries. We measured whether use of silk retreats as habitat concentrated invertebrate density and biomass compared to surrounding rock surface habitat and whether the use of retreat habitat varied across body sizes of community members along the gradient.

    We found that retreats substantially concentrated the densities of a diversity of taxa including eight different Orders, and this effect was greatest at high elevations. Caddisfly retreats also concentrated invertebrate biomass more as elevation increased. Body size of invertebrates inhabiting retreats was lower than that of surrounding rock habitats at low elevation sites, however, body size between retreats and rocks converged at higher elevation sites. Additionally, the body size of invertebrates found in retreats varied within and across taxa. Specifically, caddisfly retreats functioned as a potential nursery for taxa with large maximal body sizes. However, the patterns of this taxon‐specific nursery effect were not influenced by elevation unlike the patterns observed based on community‐level body size.

    Collectively, our results indicate that invertebrates use retreats in earlier life stages or when they are smaller in body size independent of life stage. Furthermore, our analysis suggests that facilitation strength intensifies as elevation increases within stream invertebrate communities.

    Further consideration of how trait variation and environmental gradients interact to determine the strength and direction of biotic interactions will be important as species ranges and environmental conditions continue to shift.

     
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  2. Abstract

    Ecosystem engineers can generate hotspots of ecological structure and function by facilitating the aggregation of both resources and consumers. However, nearly all examples of such engineered hotspots come from long‐lived foundation species, such as marine and freshwater mussels, intertidal cordgrasses, and alpine cushion plants, with less attention given to small‐bodied and short‐lived animals. Insects often have rapid life cycles and high population densities and are among the most diverse and ubiquitous animals on earth. Although these taxa have the potential to generate hotspots and heterogeneity comparable to that of foundation species, few studies have examined this possibility. We conducted a mesocosm experiment to examine the degree to which a stream insect ecosystem engineer, the net‐spinning caddisfly (Tricoptera:Hydropsychidae), creates hotspots by facilitating invertebrate community assembly. Our experiment used two treatments: (1) stream benthic habitat with patches of caddisfly engineers present and (2) a control treatment with no caddisflies present. We show that compared to controls, caddisflies increased local resource availability measured as particulate organic matter (POM) by 43%, ecosystem respiration (ER) by 70%, and invertebrate density, biomass, and richness by 96%, 244%, and 72%, respectively. These changes resulted in increased spatial variation of POM by 25%, invertebrate density by 76%, and ER by 29% compared to controls, indicating a strong effect of caddisflies on ecological heterogeneity. We found a positive relationship between invertebrate density and ammonium concentration in the caddisfly treatment, but no such relationship in the control, indicating that either caddisflies themselves or the invertebrate aggregations they create increased nutrient availability. When accounting for the amount of POM, caddisfly treatments increased invertebrate density by 48% and richness by 40% compared to controls, suggesting that caddisflies may also enhance the nutritional quality of resources for the invertebrate assemblage. The caddisfly treatment also increased the rate of ecosystem respiration as a function of increasing POM compared to the control. Our study demonstrates that insect ecosystem engineers can generate heterogeneity by concentrating local resources and consumers, with consequences for carbon and nutrient cycling.

     
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  3. Abstract

    Ecosystem engineers transform habitats in ways that facilitate a diversity of species; however, few investigations have isolated short‐term effects of engineers from the longer‐term legacy effects of their engineered structures. We investigated how initial presence of net‐spinning caddisflies (Hydropsychidae) and their structures that provide and modify habitat differentially influence benthic community colonization in a headwater stream by conducting an in situ experiment that included three treatments: (1) initial engineering organism with its habitat modification structure occupied (hereafter caddisfly); (2) initial habitat modification structure alone (hereafter silk); and (3) a control with the initial absence of both engineer and habitat modification structure (hereafter control). Total invertebrate colonization density and biomass was higher in caddisfly and silk treatments compared to controls (~25% and 35%, respectively). However, finer‐scale patterns of taxonomy revealed that density for one of the taxa, Chironomidae, was ~19% higher in caddisfly compared to silk treatments. Additionally, conspecific biomass was higher by an average of 50% in silk treatments compared to controls; however, no differences inHydropsychesp. biomass were detected between caddisfly treatments and controls, indicating initially abandoned silk structures elevated conspecific biomass. These findings suggest that the positive effects of the habitat modification structures that were occupied for the entirety of the experiment may outweigh any potential negative impacts from the engineer, which is known to be territorial. Importantly, these results reveal that the initial presence of the engineer itself may be important in maintaining the ecological significance of habitat modifications. Furthermore, the habitat modifications that were initially abandoned (silk) had similar positive effects on conspecific biomass compared to caddisfly treatments, suggesting legacy effects of these engineering structures may have pertinent intraspecific feedbacks of the same magnitude to that of occupied habitat modifications. Elucidating how engineers and their habitat modifications differentially facilitate organisms will allow for a clearer mechanistic understanding of the extent to which animal engineers and their actions influence aspects of community organization such as colonization.

     
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  4. Abstract

    Rising global temperatures are changing how energy and materials move through ecosystems, with potential consequences for the role of animals in these processes. We tested a central prediction of the metabolic scaling framework—the temperature independence of animal community production—using a series of geothermally heated streams and a comprehensive empirical analysis. We show that the apparent temperature sensitivity of animal production was consistent with theory for individuals (Epind = 0.64 vs. 0.65 eV), but strongly amplified relative to theoretical expectations for communities, both among (Epamong = 0.67 vs. 0 eV) and within (Epwithin = 1.52 vs. 0 eV) streams. After accounting for spatial and temporal variation in resources, we show that the apparent positive effect of temperature was driven by resource supply, providing strong empirical support for the temperatureindependenceof invertebrate production and the necessary inclusion of resources in metabolic scaling efforts.

     
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  5. Abstract

    As global temperatures continue to rise, assessment of how species within ecological communities respond to shifts in temperature has become increasingly important. However, such assessments require detailed long‐term observations or ecosystem‐level manipulations that allow for interactions among species and the potential for species dispersal and exchange with the regional species pool.

    We examined the effects of experimental whole‐stream warming on a larval black fly assemblage in southwest Iceland. We used a paired‐catchment design, in which we studied the warmed stream and a nearby reference stream for 1 year prior to warming and 2 years during warming and estimated population abundance, biomass, secondary production, and growth rates for larvae of three black fly species.

    Experimental warming by 3.8°C had contrasting effects on the three black fly species in the assemblage. The abundance, biomass, growth, and production ofProsimulium ursinumdecreased in the experimental stream during the warming manipulation. Despite increasing in the reference stream, the abundance, biomass, and production of another species,Simulium vernum, decreased in the experimental stream during warming.

    In contrast, warming had an overall positive effect onSimulium vittatum. While warming had little effect on the growth of overwintering cohorts ofS. vittatum, warming led to an additional cohort during the summer months and increased its abundance, biomass, and production. Overall, family‐level production was enhanced by warming, despite variation in species‐level responses.

    Our study illustrates that the effects of climate warming are likely to differ even among closely related species. Moreover, our study highlights the need for further investigation into the uneven effects of warming on individual species and how those variable effects influence food web dynamics and ecosystem function.

     
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  6. Abstract

    Climate warming is predicted to alter routing and flows of energy through food webs because of the critical and varied effects of temperature on physiological rates, community structure, and trophic dynamics. Few studies, however, have experimentally assessed the net effect of warming on energy flux and food web dynamics in natural intact communities. Here, we test how warming affects energy flux and the trophic basis of production in a natural invertebrate food web by experimentally heating a stream reach in southwest Iceland by ~4°C for 2 yr and comparing its response to an unheated reference stream. Previous results from this experiment showed that warming led to shifts in the structure of the invertebrate assemblage, with estimated increases in total metabolic demand but no change in annual secondary production. We hypothesized that elevated metabolic demand and invariant secondary production would combine to increase total consumption of organic matter in the food web, if diet composition did not change appreciably with warming. Dietary composition of primary consumers indeed varied little between streams and among years, with gut contents primarily consisting of diatoms (72.9%) and amorphous detritus (19.5%). Diatoms dominated the trophic basis of production of primary consumers in both study streams, contributing 79–86% to secondary production. Although warming increased the flux of filamentous algae within the food web, total resource consumption did not increase as predicted. The neutral net effect of warming on total energy flow through the food web was a result of taxon‐level variation in responses to warming, a neutral effect on total invertebrate production, and strong trophic redundancy within the invertebrate assemblage. Thus, food webs characterized by a high degree of trophic redundancy may be more resistant to the effects of climate warming than those with more diverse and specialized consumers.

     
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