Impacts of marine-derived nutrients on stream ecosystem functioning

Energy and nutrient subsidies transported across ecosystem boundaries are increasingly appreciated as key drivers of consumer-resource dynamics. As purveyors of pulsed marine-derived nutrients (MDN), spawning salmon are one such cross-ecosystem subsidy to freshwaters connected to the north Pacific. We examined how salmon carcasses influenced detrital processing in an oligotrophic stream. Experimental manipulations of MDN inputs revealed that salmon carcasses indirectly reduced detrital processing in streams through temporarily decoupling the detrital resource-consumer relationship, in which detrital consumers shifted their diet to the high-nutrient resource, i.e. salmon carcasses. The average decomposition rate of alder leaves with salmon carcass addition was significantly lower than that without the carcass, which was associated with lower abundance and biomass of detritivorous Trichoptera on the carcass-treated leaves. There were generally larger in size Trichopteran detritivores on the carcasses than on leaves. These results imply that cross-boundary MDN subsidies indirectly retard the ecosystem processing of leaf litter within the short term, but may enhance those food-limited detritivorous consumers. Because unproductive freshwaters in the Pacific northwest are highly dependent upon the organic matter inputs from surrounding forests, this novel finding has implications for determining conservation and management strategies of salmon-related aquatic ecosystems, in terms of salmon habitat protection and fisheries exploitation.     

Sea to sky: impacts of residual salmon-derived nutrients on estuarine breeding bird communities

Pacific salmon (Oncorhynchus spp.) returning to streams around the North Pacific Rim provide a nutrient subsidy to these ecosystems. While many species of animals feed directly on salmon carcasses each autumn, salmon-derived nutrients can also be stored in coastal habitats throughout the year. The effects of this storage legacy on vertebrates in other seasons are not well understood, especially in estuaries, which can receive a large portion of post-spawning salmon nutrients. We examine the effects of residual salmon-derived nutrients, forest habitats and landscape features on summer breeding birds in estuary forests. We compared models containing environmental variables and combined chum (Oncorhynchus keta) and pink (Oncorhynchus gorbuscha) salmon biomass to test predictions concerning bird density and diversity. We discovered that total bird, insectivore, golden-crowned kinglet and Pacific wren densities and Shannon's diversity in the summer were strongly predicted by salmon biomass in the autumn. For most metrics, this relationship approaches an asymptote beyond 40 000 kg of salmon biomass. Foliage height diversity, watershed catchment area and estuary area were also important predictors of avian communities. Our study suggests that the legacy of salmon nutrients influences breeding bird density and diversity in estuaries that vary across a wide gradient of spawning salmon biomass.     

Aquatic insect subsidies influence microbial composition and processing of detritus in near-shore subarctic heathland

Insects are major conduits of resources moving from aquatic to terrestrial systems. While the ecological impacts of insect subsidies are well documented, the underlying mechanisms by which these resources change recipient ecosystems remain poorly understood. Most subsidy inputs enter terrestrial systems as detritus; thus, soil microbes will likely influence the processing of insect subsidies, with implications for plant community composition and net primary productivity (NPP). In a subarctic ecosystem near Lake Mývatn, Iceland where midge (Diptera: Chironomidae) deposition to land is high, we investigated how insect subsidies affected litter processing and microbial communities. We also evaluated how those belowground effects related to changes in inorganic nitrogen, plant composition and NPP. We simulated subsidies by adding midge carcasses to 1-m² heathland plots, where we measured effects on decomposition rates and the plant community. We then studied how fertilization treatments (control, KNO₃ and midge-carcass addition) affected graminoid biomass and inorganic nitrogen in greenhouse experiments. Lastly, we conducted a soil-incubation study with a phospholipid fatty acid analysis (PLFA) to examine how midge addition to heathland soils affected microbial respiration, biomass and composition. We found that midge addition to heathland soils increased litter decomposition and graminoid plant cover by 2.6× and 2×, respectively. Greenhouse experiments revealed similar patterns, with midge carcasses increasing graminoid biomass by at least 2× and NH₄⁺ concentrations by 7×. Our soil-incubation study found that midge carcasses elevated microbial respiration by 64%, microbial biomass by 43% and shifted microbial functional composition. Our findings indicate that insect subsidies can stimulate soil microbial communities and litter decomposition in subarctic heathlands, leading to increased NPP and changes in plant community composition.     

Salmon species, density and watershed size predict magnitude of marine enrichment in riparian food webs

Resource subsidies across habitat boundaries can structure recipient communities and food webs. In the northern Pacific region, bears Ursus spp. foraging on anadromous salmon Oncorhynchus spp. provide a key link between marine and terrestrial ecosystems, with salmon density, fish size and watershed size as potential predictors of the magnitude of marine subsidy to terrestrial habitats. We use nitrogen and carbon stable isotopes to provide an assessment of the patterns of marine‐enrichment in riparian plants (11 species, 4 guilds) and litter invertebrates (4 guilds) sampled from 27 watersheds in coastal British Columbia, Canada. Watersheds occurred in three geographical regions (Vancouver Island, mainland midcoast and Haida Gwaii) and varied in size, and in biomass (kg m^?1 of spawning length) and species of salmon (chum O. keta, pink O. gorbuscha and coho O. kisutch). δ¹⁵N values in all plant species and invertebrate guilds were positively predicted by total salmon biomass (kg m^?1) and negatively predicted by watershed size. We observed replicated parallel slopes among plant species and invertebrate guilds across the gradient in salmon biomass, with differences in means hypothesized to be due to plant fractionation and animal trophic position. As such, we derived a watershed δ¹⁵N‐index averaged across guilds, and using an information theoretic approach we find that the biomass of chum salmon is a much stronger predictor of the δ¹⁵N‐index than either pink or coho salmon, or the sum biomass of all species. The top linear model contained chum biomass and watershed size. Chum salmon biomass independently predicted δ¹⁵N‐index variation in all three regions of British Columbia. Chum salmon are larger than pink or coho and provide an energetic reward for bears that facilitates carcass transfer, tissue selective foraging, and nutrient distribution by insect scavengers. Analyses of biodiversity and habitat data across many watersheds moves towards a long‐term goal in fisheries ecology to better integrate ecosystem values in salmon conservation.     

Trade-offs in community properties through time in a desert rodent community

Resource limitation represents an important constraint on ecological communities, which restricts the total abundance, biomass, and community energy flux a given community can support. However, the exact relationship among these three measures of biological activity remains unclear. Here we use a simple framework that links abundance and biomass with an energetic constraint. Under constant energetic availability, it is expected that changes in abundance and biomass can result from shifts in the distribution of individual masses. We test these predictions using long-term data from a desert rodent community. Total energy use for the community has not changed directionally for 25 years, but species composition has. As a result, the average body size has decreased by almost 50%, and average abundance has doubled. These results lend support to the idea of resource limitation on desert rodent communities and demonstrate that systems are able to maintain community energy flux in the face of environmental change, through changes in composition and structure.     

The ecology of terrestrial invertebrates on Pacific salmon carcasses

In coastal streams throughout the north Pacific region, spawning salmon (Oncorhynchus spp.) subsidize terrestrial communities with their nutrients and carcasses. We document the previously unreported composition and ecology of terrestrial invertebrates using salmon carcasses in forest habitats from two high salmon density watersheds in coastal British Columbia. From experimental placement of 186 carcasses, terrestrial Diptera-dominated salmon carcass decay (85.5% of carcasses). Overall, we recorded over 60 species from salmon carcasses, including saprophagous Diptera and Coleoptera (15 spp.), dipteran predators (eight spp.) and parasitoids (four spp.), and opportunistic predators, scavengers, and detritivores (24 spp.). Using stable isotopes of nitrogen and carbon, we reconstruct the dietary niches of select species relative to salmon muscle tissue and previously sampled non-salmon feeding invertebrate species. From comparisons across seasons, sampling locations and larval and adult life stages, we find evidence for a diet of salmon tissue in flies (Calliphora terraenovae and Dryomyza anilis), and beetles (Nicrophorus investigator and Anthobium fimetarium). The parasitic wasps Alysia alticola and Atractodes sp. had the highest levels of enrichment of all species, representing their larval diet of fly larvae that have fed on salmon carcasses 1 year prior to adult wasp sampling. Temporal and spatial isotopic variation in insect indicator species varies by species mobility and the pathway of salmon nutrient uptake. Cataloguing these associations may be useful for developing indices of intact salmon runs, bear foraging, and subsequent nutrient transfer in coastal watersheds.     

乐清湾贝类群落组成及其粒径谱结构特征

为了解乐清湾贝类资源情况及群落稳定性特征,基于2016—2017年乐清湾定点调查数据,分析了不同季节乐清湾贝类组成、优势种、资源密度、以及物种多样性,并首次建立乐清湾海域贝类生物量粒径谱与标准化生物量粒径谱。结果显示：1)本次共采集到贝类共计45种,隶属于12目,25科,31属。焦河蓝蛤(Potamocorbula ustulata)为乐清湾绝对优势种;2)资源密度表现为夏季最高,其次为春季,秋季与冬季次之;4个季节的丰富度指数(D)、多样性指数(H′)和均匀度指数(J′)平均为3.26、1.61、0.45,3种多样性指数均处于较低水平;3)ABC曲线显示乐清湾贝类的生物群落处于中度干扰状态;4)乐清湾生物量粒径谱谱型复杂,优势种控制着生物量粒径谱的峰值,且与我国其他海域的结果也存在差异;5)标准化生物量粒径谱的斜率范围为-1.5539—-0.7373,在我国近海海域中属于偏低水平,表明营养循环水平较低。截距范围在16.673—21.597,高于我国其他海域,说明乐清湾贝类生产力水平较高。     

Spatial scaling laws do not structure strongyloid nematode communities in macropodid hosts

The characteristics of species within a community can influence the number of species that can coexist within that community. In particular, body size can constrain how many individuals can 'fit' into a community, and overlap in resource use between species depends on differences in their body sizes. Here, using data from 18 communities of strongyloid nematodes living in the stomachs of macropodid marsupials, we test key predictions derived from spatial scaling laws regarding the minimum similarity in body size between coexisting species believed to control how many species can coexist in a community. These communities are ideal systems for such a test: they consist of huge numbers of individuals from numerous species, all belonging to the same family (Chabertiidae) and living in the same host organ. Within these communities, we found that mean abundance correlated negatively with body size across all nematode species, whether body size was measured as length or volume. However, we found no support for the predictions of spatial scaling laws. First, the size ratios of pairs of adjacent-sized species did not decrease as a function of the size of the largest species in a pair. The few significant relationships observed were all positive, suggesting that the relative difference in size between adjacent species in the size hierarchy may in fact increase toward the upper end of the size spectrum. Second, the frequency distributions of body sizes were predominantly right-skewed amongst the communities investigated: within the size spectrum observed in a nematode community, small-bodied species greatly outnumber large-bodied ones, in sharp contrast to the predictions of spatial scaling laws. Nematode body size may thus determine the abundance achieved by a species but not how many species can coexist; the limiting similarity between coexisting species must depend on other biological traits.     

Burying beetle Nicrophorus investigator reproduction on Pacific salmon carcasses

Abstract.  1. In many undisturbed watersheds along the Pacific Rim, anadromous salmon ( Oncorhynchus spp.) provide a predictable source of carrion to the riparian zone, largely due to horizontal transfer of salmon carcasses by bears ( Ursus spp.) and other vertebrates.
2. Burying beetles are important members of the north-temperate carrion fauna, and may utilise salmon carcasses and remnants for breeding. In this study, isotopic and observational data are reported that demonstrate previously unrecognised Nicrophorus investigator (Zetterstedt) reproduction on large salmon carcasses from five watersheds in coastal British Columbia.
3. Stable isotope signatures (δ¹⁵N and δ¹³C) of adult beetles collected in autumn indicate a diet of salmon origin in all but one individual from all watersheds, suggesting that this beetle–salmon association is widespread. Comparison of autumn isotope signatures to individuals collected randomly in summer suggests that isotope signatures represent the larval carrion source from the previous autumn rather than immediate adult diet.
4. In a survey of N. investigator use of salmon carcasses from two watersheds, 35 broods were observed on chum and pink salmon carcasses, including 16 natural brood complexes containing over 100 larvae, and five ranging from 250 to 750 larvae.
5. Overall, north-coastal populations of N. investigator breed on the rich and reliable salmon resource and may exhibit a system of communal breeding on these carcasses. This is most relevant when the dramatic reduction in salmon spawning biomass over the last century is considered.     

Influence of Tree Species Composition and Community Structure on Carbon Density in a Subtropical Forest

We assessed the impact of species composition and stand structure on the spatial variation of forest carbon density using data collected from a 4-ha plot in a subtropical forest in southern China. We found that 1) forest biomass carbon density significantly differed among communities, reflecting a significant effect of community structure and species composition on carbon accumulation; 2) soil organic carbon density increased whereas stand biomass carbon density decreased across communities, indicating that different mechanisms might account for the accumulation of stand biomass carbon and soil organic carbon in the subtropical forest; and 3) a small number of tree individuals of the medium- and large-diameter class contributed predominantly to biomass carbon accumulation in the community, whereas a large number of seedlings and saplings were responsible for a small proportion of the total forest carbon stock. These findings demonstrate that both biomass carbon and soil carbon density in the subtropical forest are sensitive to species composition and community structure, and that heterogeneity in species composition and stand structure should be taken into account to ensure accurate forest carbon accounting.     

跨越生境的资源补贴对生态系统的影响研究进展

跨越不同生态系统之间的物质、能量和营养元素流动,即资源补贴,是生态系统的基本属性,也是生态学研究的基本问题之一.常见的资源补贴包括落入水体的树叶凋落物和陆地昆虫、水生昆虫成虫、从海洋生殖洄游的鲑鱼等,这些外源性的资源补贴对接收生态系统的生物个体、种群、群落、生物多样性和生态系统功能都有影响,包括促进个体生长、增加物种丰度和多样性、改变群落结构、增加生态系统的生产力、改变食物链长度及影响食物网、影响生态系统的稳定性等.随着未来人类活动对环境扰动的增加,尤其在土地利用、气候变化、生物入侵方面,对跨越生态系统资源补贴的时空动态影响将加剧,因而生态系统将面临更加严峻的威胁.鉴于此,未来在该领域的基础研究应着重开展以下几方面研究:单一和多重环境胁迫对资源补贴和生态系统的影响;动态资源补贴在生态系统修复及管理中的应用;关注与污染物相关的资源补贴的负面影响;加强跨越生境资源补贴在热带和亚热带以及在我国的生态学基础研究.     

土壤微生物削弱了水生-陆地系统补贴对植物生长的正向影响

土壤微生物削弱了水生-陆地系统补贴对植物生长的正向影响 水生-陆地系统补贴形成的联结作用在构建群落和调节生态系统功能方面发挥重要作用。在营养贫瘠的生态系统中(例如密歇根湖周围的淡水沙丘),水生-陆生系统补贴显得尤为重要。春季成年蠓在密歇根湖涌出,成群交配,然后死亡。蠓尸体在植物的基部形成土丘状,通过输入营养提高植物的生产力。然而,水生-陆地系统补贴对植物生产力的影响可能取决于其他生物的交互作用,特别是土壤微生物可能通过促进养分转化为植物可利用的形式或与植物竞争养分而发挥关键作用。在温室实验中,我们检验了湖生蠓(Chironomidae)的尸体和土壤微生物如何独立和相互影响一种常见沙丘草(沙拂子茅,Calamovilfa longifolia)的生长表现。为确定蠓是否影响土壤非生物特性,我们检验了添加蠓如何影响土壤养分和土壤湿度。研究结果显示,蠓极大地增加了植物生物量,但其效应的大小受土壤微生物的影响。在没有土壤微生物的情况下,添加蠓的植物生物量比没有添加的高7倍,而在有土壤微生物的情况下,植物生物量提高了3倍。蠓对植物生长的促进作用可能由于它们向土壤中输入养分所导致,因为与沙丘土壤相比,蠓的氮、磷、钾含量分别高100倍、10倍和150倍。我们的研究结果表明,土壤微生物可能与植物竞争这些养分。总之,我们发现蠓是重要的水生-陆地系统补贴,对密歇根湖沿岸植物生产力产生强烈和正向的影响,但水生-陆地系统补贴作用必须在生态群落内发生的复杂相互作用的背景下考虑。     

Biogeochemical transformation of a nutrient subsidy: salmon, streams, and nitrification

Migratory animals can alter ecosystem function via the provision of nutrient subsidies. These subsidies are heterogeneous in space and time, which may create hot spots or hot moments in biogeochemical transformations, in turn altering the ecosystem effect of the subsidy by changing the form of the nutrients. Annual migrations of Pacific salmon (Oncorhynchus spp.) transport nutrients from the marine environment to their natal freshwater ecosystems. Salmon subsidies provide high quality nutrients (e.g., nitrogen, phosphorus, carbon) that may also be large in quantity where salmon migrations are near historic levels. We hypothesized that the nutrient subsidy provided via the excretion of ammonium (NH₄ ⁺) by live salmon would stimulate microbially mediated nitrification rates in stream sediments and increase streamwater nitrate (NO₃ ^?) concentrations. We quantified sediment nitrification in seven streams in Southeast Alaska before and during the salmon run in 2007 and 2008. Nitrification rates increased 3-fold from before to during the salmon run (mean ± SE = 0.07 ± 0.01 to 0.24 ± 0.02 mgN gAFDM^?1 d^?1, respectively). The variation in nitrification was explained by both streamwater and exchangeable NH₄ ⁺ concentrations (R ² = 0.50 and 0.71, respectively), which were low before salmon and increased relative to the size of the salmon run. To experimentally test the effect of salmon subsidies on nitrification rates, we staked senesced salmon carcasses on stream sediments for 3 weeks during the salmon run and then measured nitrification rates directly under the carcasses. Sediment nitrification was 2–5 times higher under the carcasses compared to nearby sediments without the direct carcass influence. Our results confirm that biogeochemical transformations alter the form of salmon-derived nitrogen, representing an overlooked aspect in the dynamics of this subsidy. Therefore, animal-derived nutrient subsidies are not passively retained or exported in recipient ecosystems, but also transformed, thereby influencing the form and incorporation of these nutrient subsidies.     

Resources and global avian assemblage structure in forests

Explaining spatial variation in a number of bird species, particularly from temperate to tropical regions, has been a longstanding challenge. We test at a global scale whether species‐rich forest assemblages are associated with division of a larger resource pool, a finer division of that pool, or some combination of the two. Species richness increases with increasing assemblage abundance, biomass and energy use. As assemblage abundance, biomass and energy use increase with increasing energy availability, and as per species numbers of individuals, biomass and energy use do not decrease with increasing energy availability, we provide direct evidence that the avian species–energy relationship in forests is associated foremost with an increase in the size of the resource pool and not with a finer level of its subdivision.     

Litter Biomass and Nutrient Determinants of Ant Density, Nest Size, and Growth in a Costa Rican Tropical Wet Forest

The nutritional demands of animals vary by taxon. Across landscapes, communities of animals experience variability in the stoichiometry of carbon and nutrients within their resource base. Thus, we expect stoichiometry to contribute to the spatial variance in the demographic parameters of animal communities. Here, we measure how the composition of a litter-nesting tropical rainforest ant community is influenced by spatial variation in environmental stoichiometry relative to litter biomass, a known predictor of ant density. We found the density of ants and their nests were strongly related to litter biomass and carbon: phosphorus stoichiometry. The spatial variation in soil nutrients, which determines leaf litter stoichiometry, was an excellent predictor of nest size in the two most common genera of ants. We found a negative relationship between species' growth rate and local soil stocks of phosphorus. Overall, the density of litter-dwelling ants varied greatly across this tropical forest landscape and environmental stoichiometry can account for limits on ant density independent of the biomass of the leaf litter resource base.     

Environmental,structural, and taxonomic diversity factors drive aboveground carbon stocks in semi-deciduous tropical rainforest strata in Cameroon

Forest stratification plays a crucial role in the interception of light and plants' photosynthetic activities. However, there is still a lack of information on the contribution of tropical forest stratification to its functioning, despite the increasing number of studies. Here, we analysed from a perspective of the whole tree community (WTC) and forest strata (i.e., large trees, understory trees, and small stems), the relationship between abiotic, biotic factors and aboveground Carbon (AGC). The abiotic factors-AGC relationships were positive for all strata and WTC. However, soil factors-AGC relationship was stronger for small stems and understorey, while topography factor-AGC relationship was stronger for large trees and WTC. Tree size inequality-AGC relationship was positive and much stronger for WTC, large trees and small stems. In addition, a species diversity-AGC relationship was found positive only for large trees and WTC. These results highlight the niche complementarity effect for driving positive relationships of species diversity and individual tree size variation with aboveground biomass at large tree strata and WTC. The lack of positive effect of species diversity on AGC for understorey and small stems strata might be attributable to the selection effect or resource complementarity among species.     

Resource availability mediates the importance of priority effects in plant community assembly and ecosystem function

Assembly history, including the order in which species arrive into a community, can influence long‐term community structure; however we know less about how timing of species arrival may alter assembly especially under varying resource conditions. To explore how the timing of species arrival interacts with resource availability to alter community assembly, we constructed experimental plant communities and manipulated the interval between plantings of groups of seedlings (0, 5, 10, 15 or 20 days) at low and high levels of soil nutrient supply. To see if community changes influenced ecosystem‐scale processes, we measured parameters across the plant–soil continuum (e.g. plant biomass and net ecosystem carbon dioxide exchange). We found that the timing of species arrival had a large impact on community assembly, but the size of the effect depended on soil fertility. As planting interval increased, plant communities diverged further from the control, but the divergence was stronger at high than at low nutrient supply. Our data suggest that at high nutrient supply, early‐planted species preempted light resources more quickly, thus preventing the successful establishment of later arriving species even at short planting intervals. Finally, we found that assembly related divergence in plant communities scaled to impact ecosystem‐level characteristics such as green leaf chemistry, but had little effect on total community biomass and net ecosystem exchange of CO₂ and water vapor. Our data indicate that the effect of a stochastic factor, here the timing of species arrival on community composition, depends on the resource level under which the community assembles.     

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of～4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.     

Fecundity and egg size variation in North American Pacific salmon (Oncorhynchus)

We examined regional and latitudinal variation in fecundity and egg weight for five species of Pacific salmon ( Oncorhynchus ) along the Pacific coast of North America. Data were examined for 24 chum salmon, 15 pink salmon, 34 sockeye salmon, 44 chinook salmon, and 40 coho salmon populations from published sources, unpublished Canadian hatchery records, our own laboratory investigations, and other unpublished sources. Substantial regional variation in fecundity and egg weight was observed, with salmon on the Queen Charlotte Islands and Vancouver Island in British Columbia generally having lower fecundity and larger egg size than nearby mainland populations. The relative distance of freshwater migration to the spawning grounds generally had a marked effect on both fecundity and egg size, with populations spawning in the upper portions in the drainages of large rivers like the Fraser River in British Columbia having reduced fecundity and egg size compared with coastal spawning populations. Fecundity was generally higher and egg size generally lower in more northern populations of sockeye, chinook, and coho salmon compared with southern ones. We suggest that egg size tends to be lower in northern populations of some species as a result of increased fecundity due to their older ages at maturity and a limited amount of energy that can be expended on egg production.     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.