A long-term stream fertilization experiment was performed to evaluate the potential eutrophication of an arctic stream ecosystem. During 16 years of summer phosphorus (H 3 PO 4 ) fertilization, we observed a dramatic change in the community structure of the Kuparuk River on the North Slope of Alaska. A positive response to fertilization was observed at all trophic levels with increases in epilithic algal stocks, some insect densities, and fish growth rates. After approximately eight years of P fertilization, bryophytes (mosses) replaced epilithic diatoms as the dominant primary producers in the Kuparuk River. The moss impacted NH 4 ϩ uptake rates, benthic gross primary production, habitat structure, and insect abundance and species composition. This study documents the long-term changes in an arctic tundra stream in response to nutrient enrichment. Predicting stream ecosystem responses to chronic perturbation requires long-term observation and experiments.

Abstract:We predicted that substratum freezing and instability are major determinants of the variability of stream community structure in Arctic Alaska. Their effects were conceptualized as a two-dimensional habitat template that was assessed using a natural experiment based on five stream types (mountain-spring, tundra-spring, tundra, mountain, glacier). Detrended correspondence analysis (DCA) indicated distinct macroinvertebrate assemblages for each stream type. The contribution of functional feeding groups to assemblage biomass varied systematically among stream types, indicating that structure and function are linked. Assemblage position within a DCA biplot was used to assess factors controlling its structure. Springs separated from other stream types along a gradient of nutrient concentration and freezing probability. Glacier and mountain streams separated from springs and tundra streams along a gradient of substratum instability and freezing probability. Owing to differences in sources of discharge to streams, the effects of nutrients and substratum stability could not be separated from freezing. Although many factors likely contribute to the variability of Arctic stream communities, the major determinants may be conceptualized as a template structured by gradients in (i) nutrient supply and substratum freezing and (ii) substratum instability and substratum freezing. This template provides a basis for predicting the response of Arctic stream communities to climate change. Résumé :Nous avons prédit que le gel et l'instabilité du substrat sont des causes majeures de la variabilité de la structure des communautés des cours d'eau en Alaska arctique. Nous avons conceptualisé ces effets dans un cadre bidimensionnel de l'habitat qui a été évalué au cours d'une expérience en nature dans cinq types de cours d'eau (source montagneuse, source de toundra, toundra, montagne et glacier). Une analyse de correspondance redressée (DCA) révèle des peuplements caractéristiques de macroinvertébrés dans chacun des types de cours d'eau. La contribution des diffé-rentes guildes alimentaires fonctionnelles à la biomasse du peuplement varie systématiquement selon le type de cours d'eau, ce qui montre un lien entre la structure et la fonction. La position de chaque peuplement sur le graphique DCA bidimensionnel sert à évaluer les facteurs qui contrôlent sa structure. Les sources se séparent des autres types de cours d'eau le long d'un gradient de concentration de nutriments et de probabilité de gel. Les cours d'eau de montagne et de glaciers se distinguent des sources et des cours d'eau de toundra sur un gradient d'instabilité du substrat et de probabilité de gel. À cause des différentes sources des apports vers les cours d'eau, il est impossible de distinguer les effets des nutriments et de la stabilité du substrat de ceux du gel. Bien que de nombreux facteurs contribuent vraisemblablement à la variabilité des communautés des cours d'eau arctiques, on peut imaginer les principales causes déterminantes sur un cadre structuré par de...

Primary producers form the base of food webs but also affect other ecosystem characteristics, such as habitat structure, light availability, and microclimate. Here, we examine changes caused by 5-30+ years of nutrient addition and resulting increases in net primary productivity (NPP) in tundra, streams, and lakes in northern Alaska. The Arctic provides an important opportunity to examine how ecosystems characterized by low diversity and low productivity respond to release from nutrient limitation. We review how responses of algae and plants affect light availability, perennial biotic structures available for consumers, oxygen levels, and temperature. Sometimes, responses were similar across all three ecosystems; e.g., increased NPP significantly reduced light to the substrate following fertilization. Perennial biotic structures increased in tundra and streams but not in lakes, and provided important new habitat niches for consumers as well as other producers. Oxygen and temperature responses also differed. Life history traits (e.g., longevity) of the primary producers along with the fate of detritus drove the responses and recovery. As global change persists and nutrients become more available in the Arctic and elsewhere, incorporating these factors as response variables will enable better prediction of ecosystem changes and feedbacks in this biome and others.

1. Nutrient enrichment and resulting eutrophication is a widespread anthropogenic influence on freshwater ecosystems, but recovery from nutrient enrichment is poorly understood, especially in stream environments. We examined multi-year patterns in community recovery from experimental low-concentration nutrient enrichment (N + P or P only) in three reaches of two Arctic tundra streams (Kuparuk River and Oksrukuyik Creek) on the North Slope of Alaska (U.S.A.). 2. Rates of recovery varied among community components and depended on duration of enrichment (2-13 consecutive growing seasons). Biomass of epilithic algae returned to reference levels rapidly (within 2 years), regardless of nutrients added or enrichment duration. Aquatic bryophyte cover, which increased greatly in the Kuparuk River only after long-term enrichment (8 years), took 8 years of recovery to approach reference levels, after storms had scoured most remnant moss in the recovering reach. 3. Multi-year persistence of bryophytes in the Kuparuk River appeared to prevent recovery of insect populations that had either been positively (e.g. the mayfly Ephemerella, most chironomid midge taxa) or negatively (e.g. the tube-building chironomid Orthocladius rivulorum) affected by this shift in dominant primary producer. These lags in recovery (of >3 years) were probably driven by the persistent effect of bryophytes on physical benthic habitat. 4. Summer growth rates of Arctic grayling (both adults and young-of-year) in Oksrukuyik Creek (fertilised for 6 years with no bryophyte colonisation), which were consistently increased by nutrient addition, returned to reference rates within 1-2 years. 5. Rates of recovery of these virtually pristine Arctic stream ecosystems from low-level nutrient enrichment appeared to be controlled largely by duration of enrichment, mediated through physical habitat shifts caused by eventual bryophyte colonisation, and subsequent physical disturbance that removed bryophytes. Nutrient enrichment of oligotrophic Arctic stream ecosystems caused by climate change or local anthropogenic activity may have dramatic and persistent consequences if it results in the colonisation of long-lived primary producers that alter physical habitat.

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