Stream hydrology and a pulse subsidy shape patterns of fish foraging.

Pulsed subsidy events create ephemeral fluxes of hyper-abundant resources that can shape annual patterns of consumption and growth for recipient consumers. However, environmental conditions strongly affect local resource availability for much of the year, and can heavily impact consumer foraging and growth patterns prior to pulsed subsidy events. Thus, a consumer's capacity to exploit pulse subsidy resources may be influenced by antecedent environmental conditions, but this has rarely been shown in nature and is unknown in aquatic ecosystems. Here, we sought to understand the importance of hydrologic variation and a salmon pulse subsidy on the foraging and growth patterns of two stream salmonids in a coastal southeast Alaska drainage. To do this, we sampled fish stomach contents at a high temporal frequency (daily-weekly measurements) and analyzed fish consumption rates in relation to streamflow and pulse subsidy resource availability. We then explored the influence of interannual hydrologic variation on access to pulse subsidy resources (i.e. whether fish exceeded an egg consumption gape limit) in a bioenergetic simulation. Prior to Pink Salmon spawning, Dolly Varden and Coho Salmon displayed distinct and nonlinear flow-foraging relationships, where forage for both species consisted primarily of macroinvertebrates. During this time period, consumption maxima coincided with baseflow and the highest observed flow conditions, and consumption minima were observed at severe low-water and intermediate flow values. After salmon spawning began, forage was not significantly related to flow and consisted primarily of salmon eggs. Further, consumption rates increased overall, and foraging patterns did not appear to be affected by flow in either species. Bioenergetic simulations revealed that patterns of interannual hydrologic variation may shift Coho Salmon growth trajectories among years. Together, our results suggest that access to marine pulse subsidy resources may depend on whether antecedent hydrologic conditions are suitable for juvenile salmonids to grow large enough to consume salmon eggs by the onset of spawning.

A melting cryosphere constrains fish growth by synchronizing the seasonal phenology of river food webs.

Mountain watersheds often contain a mosaic of glacier-, snow-, and rain-fed streams that have distinct hydrologic, temperature, and biogeochemical regimes. However, as glaciers diminish and precipitation shifts from snow to rain, the physical and chemical characteristics that make glacial or snowmelt streams distinct from rain-fed streams will fade. Among the unforeseen consequences of this hydrologic homogenization could be the loss of unique food webs that sustain aquatic consumers. To explore the impacts of a melting cryosphere on stream food webs, we parameterized an aquatic food web model with empirical physicochemical data from glacier-, snow-, and rain-fed streams in southeast Alaska and used the model to simulate the seasonal biomass dynamics of aquatic primary producers and consumers and the growth of juvenile salmon. Model results suggest that glacier-, snow-, and rain-fed streams exhibit seasonal asynchronies in the timing of biofilm and aquatic invertebrate abundance. Although warmer rain-fed streams were more productive during the summer (June through September), colder glacial and snowmelt streams provided enhanced foraging and growth opportunities throughout the remainder of the year. For juvenile salmon that can track peaks in resource abundance within river networks, the loss of meltwater streams strongly constrained modeled growth opportunities by removing spatially and temporally distinct foraging habitats within a watershed. These findings suggest that climate change induced homogenization of high latitude river networks may result in the loss of unique food web dynamics, which could diminish the capacity of watersheds to sustain mobile consumers.

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs.

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier-marine habitats by developing a multi-trophic level Bayesian three-isotope mixing model. We utilized large gradients in stable (δ13 C, δ15 N, δ2 H) and radiogenic (Δ14 C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial-marine habitats. We also compared isotope ratios between glacial-marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic-level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier-nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest-nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100-1500 years BP 14 C-age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP 14 C-age to modern). Thus terrestrial-derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial-marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate-driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.

Spatial Variation in the Origin of Dissolved Organic Carbon in Snow on the Juneau Icefield, Southeast Alaska.

Dissolved organic carbon (DOC) plays a fundamental role in the biogeochemistry of glacier ecosystems. However, the specific sources of glacier DOC remain unresolved. To assess the origin and nature of glacier DOC, we collected snow from 10 locations along a transect across the Juneau Icefield, Alaska extending from the coast toward the interior. The Δ(14)C-DOC of snow varied from -743 to -420‰ showing progressive depletion across the Icefield as δ(18)O of water became more depleted (R(2) = 0.56). Older DOC corresponded to lower DOC concentrations in snow (R(2) = 0.31) and a decrease in percent humic-like fluorescence (R(2) = 0.36), indicating an overall decrease in modern DOC across the Icefield. Carbon isotopic signatures ((13)C and (14)C) combined with a three-source mixing model showed that DOC deposited in snow across the Icefield reflects fossil fuel combustion products (43-73%) and to a lesser extent marine (21-41%) and terrestrial sources (1-26%). Our finding that combustion aerosols are a large source of DOC to the glacier ecosystem during the early spring (April-May) together with the pronounced rates of glacier melting in the region suggests that the delivery of relic DOC to the ocean may be increasing and consequently impacting the biogeochemistry of glacial and proglacial ecosystems in unanticipated ways.

Stream Physical Characteristics Impact Habitat Quality for Pacific Salmon in Two Temperate Coastal Watersheds.

Climate warming is likely to cause both indirect and direct impacts on the biophysical properties of stream ecosystems especially in regions that support societally important fish species such as Pacific salmon. We studied the seasonal variability and interaction between stream temperature and DO in a low-gradient, forested stream and a glacial-fed stream in coastal southeast Alaska to assess how these key physical parameters impact freshwater habitat quality for salmon. We also use multiple regression analysis to evaluate how discharge and air temperature influence the seasonal patterns in stream temperature and DO. Mean daily stream temperature ranged from 1.1 to 16.4°C in non-glacial Peterson Creek but only 1.0 to 8.8°C in glacial-fed Cowee Creek, reflecting the strong moderating influence glacier meltwater had on stream temperature. Peterson Creek had mean daily DO concentrations ranging from 3.8 to 14.1 mg L(-1) suggesting future climate changes could result in an even greater depletion in DO. Mean daily stream temperature strongly controlled mean daily DO in both Peterson (R2=0.82, P<0.01) and Cowee Creek (R2=0.93, P<0.01). However, DO in Peterson Creek was mildly related to stream temperature (R2=0.15, P<0.01) and strongly influenced by discharge (R2=0.46, P<0.01) on days when stream temperature exceeded 10°C. Moreover, Peterson Creek had DO values that were particularly low (<5.0 mg L(-1)) on days when discharge was low but also when spawning salmon were abundant. Our results demonstrate the complexity of stream temperature and DO regimes in coastal temperate watersheds and highlight the need for watershed managers to move towards multi-factor risk assessment of potential habitat quality for salmon rather than single factor assessments alone.

Characterizing dissolved organic matter using PARAFAC modeling of fluorescence spectroscopy: a comparison of two models.

We evaluated whether fitting fluorescence excitation-emission matrices (EEMs) to a previously validated PARAFAC model is an acceptable alternative to building an original model. To do this, we built a 10-component model using 307 EEMs collected from southeast Alaskan soil and streamwater. All 307 EEMs were then fit to the existing model (CM) presented in Cory and McKnight (Environ. Sci. Technol. 2005, 39, 8142-8149). The first approach for evaluating whether the EEMs were fit well to the CM model was an evaluation of the residual EEMs, and we found 22 EEMs were fit poorly by the CM model. Our second measure for verifying whether EEMs were fit well to the CM model was a comparison of correlations between the percent contribution of PARAFAC components and DOM measurements (e.g., dissolved nutrient concentrations), and we found no significant difference Ip > 0.05) between the two models. These results support the approach of fitting EEMs to an existing model when DOM is collected from similar environments, which can potentially reduce some of the problems when building an original PARAFAC model. However, it is important to recognize that some of the sensitivity or ecological interpretative power may be lost when fitting EEMs to an existing model.

An evaluation of freezing as a preservation technique for analyzing dissolved organic C, N and P in surface water samples.

Techniques for preserving surface water samples are recently in demand because of the increased interest in quantifying dissolved organic matter (DOM) in surface waters and the frequent collection of samples in remote locations. Freezing is a common technique employed by many researchers for preserving surface water samples; however, there has been little evaluation of the effects of freezing on DOM concentrations. Ten streams were sampled in southeast Alaska with a range of dissolved organic carbon (DOC) concentrations (1.5 to 39 mg C L(-1)) to evaluate the influence of freezing (flash and standard freeze) and filter pore size (0.2 and 0.7 mum nominal pore size) on dissolved organic C, N and P concentrations. We report a significant decrease in DOC (p<0.005) and total dissolved P (p<0.005) concentrations when streamwater samples were frozen, whereas concentrations of dissolved organic N did not significantly decrease after freezing (p=0.06). We further show that when surface water samples were frozen, there was a decrease in the specific ultraviolet absorbance (SUVA) of DOC that is particularly evident with high concentrations of DOC. This finding suggests that spectroscopic properties of DOC have the potential to be used as indicators of whether surface water samples can be frozen. Our results lead us to recommend that surface water samples with high DOC concentrations (>5 mg C L(-1)) and/or samples with high SUVA values (>3.5-4 L mg-C(-1) m(-1)) should be analyzed immediately and not frozen.