Jacques C. Finlay

STABLE‐CARBON‐ISOTOPE RATIOS OF RIVER BIOTA:IMPLICATIONS FOR ENERGY FLOW IN LOTIC FOOD WEBS

Stable-isotope ratios of carbon ( 13 C/ 12 Co rd 13 C) have been widely used to determine the energy base of stream food webs, but such use is controversial due to un- explained variability in algal d 13 C. I used published d 13 C data from temperate headwater streams through medium-sized rivers (0.2-4000 km 2 watershed area) collected during sum- mer baseflows and original data from streams in northern California to analyze energy pathways through river food webs. The analyses showed three important results. First, epilithic algal d 13 C and watershed area are positively related, suggesting that effects of carbon limitation on algal carbon uptake result in 13 C enrichment of algal d 13 C in larger, more productive rivers. Second, epilithic algae and terrestrial detritus d 13 C values are often distinct in small shaded streams but overlap in some larger unshaded streams and rivers. Measurements of d 13 C values may be most useful in distinguishing algal and terrestrial energy sources in unproductive streams with supersaturated dissolved CO2 concentrations, and some productive rivers where CO2 concentrations are low relative to photosynthetic rates. Finally, consumer d 13 C values are more strongly related to algal d 13 C than terrestrial d 13 C. The relative contribution of terrestrial and algal carbon sources often varied by func- tional feeding group within and between sites. However, with the exception of shredders and scrapers, which respectively relied on terrestrial and algal carbon sources, patterns of consumer d 13 C clearly show a transition from terrestrial to algal carbon sources for many lotic food webs in streams with

Dams and downstream aquatic biodiversity: Potential food web consequences of hydrologic and geomorphic change

10 km 2 watershed area. The observed transition to algal carbon sources is likely related to increasing primary production rates as forest canopy cover declines in larger streams, although decreasing retention or quality of terrestrial carbon may also play a role. Improved analyses of algal d 13 C and d 15 N combined with quantitative study of organic matter dynamics and food web structure should allow the relative im- portance of these factors to be distinguished in future food web studies.

Controls of streamwater dissolved inorganic carbon dynamics in a forested watershed

Responses of rivers and river ecosystems to dams are complex and varied, as they depend on local sediment supplies, geomorphic constraints, climate, dam structure and operation, and key attributes of the biota. Therefore, “one-size-fits-all” prescriptions cannot substitute for local knowledge in developing prescriptions for dam structure and operation to protect local biodiversity. One general principle is self-evident: that biodiversity is best protected in rivers where physical regimes are the most natural. A sufficiently natural regime of flow variation is particularly crucial for river biota and food webs. We review our research and that of others to illustrate the ecological importance of alternating periods of low and high flow, of periodic bed scour, and of floodplain inundation and dewatering. These fluctuations regulate both the life cycles of river biota and species interactions in the food webs that sustain them. Even if the focus of biodiversity conservation efforts is on a target species rather than whole ecosystems, a food web perspective is necessary, because populations of any species depend critically on how their resources, prey, and potential predators also respond to environmental change. In regulated rivers, managers must determine how the frequency, magnitude, and timing of hydrologic events interact to constrain or support species and food webs. Simple ecological modeling, tailored to local systems, may provide a framework and some insight into explaining ecosystem response to dams and should give direction to mitigation efforts.

SPATIAL SCALES OF CARBON FLOW IN A RIVER FOOD WEB

Iinvestigated controls of stream dissolved inorganic carbon (DIC) sources andcycling along a stream size and productivity gradient in a temperate forestedwatershed in northern California. Dissolved CO2 (CO2(aq))dynamics in heavily shaded streams contrasted strongly with those of larger,open canopied sites. In streams with canopy cover > 97%, CO2 (aq)was highest during baseflow periods (up to 540 μM) and wasnegatively related to discharge. Effects of algal photosynthesis on CO2(aq) were minimal and stream CO2 (aq) was primarily controlledby groundwater CO2 (aq) inputs and degassing losses to theatmosphere. In contrast to the small streams, CO2 (aq) in larger,open-canopied streams was often below atmospheric levels at midday duringbaseflow and was positively related to discharge. Here, stream CO2(aq) was strongly influenced by the balance between autotrophic andheterotrophic processes. Dynamics of HCO3− werelesscomplex. HCO3− and Ca2+ were positivelycorrelated, negatively related to discharge, and showed no pattern with streamsize. Stable carbon isotope ratios of DIC (i.e. δ13C DIC)increased with stream size and discharge, indicating contrasting sources of DICto streams and rivers. During summer baseflows, δ13C DIC were13C-depleted in the smallest streams (minimum of−17.7‰) due to the influence of CO2 (aq) derived frommicrobialrespiration and HCO3− derived from carbonateweathering. δ13C DIC were higher (up to −6.6‰)inthe larger streams and rivers due to invasion of atmospheric CO2enhanced by algal CO2 (aq) uptake. While small streams wereinfluenced by groundwater inputs, patterns in CO2 (aq) and evidencefrom stable isotopes demonstrate the strong influence of stream metabolism andCO2 exchange with the atmosphere on stream and river carbon cycles.

Linking water quality and well-being for improved assessment and valuation of ecosystem services

Spatial extents of food webs that support stream and river consumers are largely unknown, but such information is essential for basic understanding and management of lotic ecosystems. We used predictable variation in algal d 13 C with water velocity, and measurements of consumer d 13 C and d 15 N to examine carbon flow and trophic structure in food webs of the South Fork Eel River in Northern California. Analyses of d 13 C showed that the most abundant macroinvertebrate groups (collector-gatherers and scrapers) relied on algae from local sources within their riffle or shallow pool habitats. In contrast, filter- feeding invertebrates in riffles relied in part on algal production derived from upstream shallow pools. Riffle invertebrate predators also relied in part on consumers of pool-derived algal carbon. One abundant taxon drifting from shallow pools and riffles (baetid mayflies) relied on algal production derived from the habitats from which they dispersed. The trophic linkage from pool algae to riffle invertebrate predators was thus mediated through either predation on pool herbivores dispersing into riffles, or on filter feeders. Algal production in shallow pool habitats dominated the resource base of vertebrate predators in all habitats at the end of the summer. We could not distinguish between the trophic roles of riffle algae and terrestrial detritus, but both carbon sources appeared to play minor roles for vertebrate consumers. In shallow pools, small vertebrates, including three-spined stickleback ( Gasterosteus aculeatus), roach (Hesperoleucas symmetricus) , and rough-skinned newts (Taricha granulosa), relied on invertebrate prey derived from local pool habitats. During the most productive summer period, growth of all size classes of steelhead and resident rainbow trout (Oncorhynchus mykiss)in all habitats (shallow pools, riffles, and deep unproductive pools) was largely derived from algal production in shallow pools. Preliminary data suggest that the strong role of shallow pool algae in riffle steelhead growth during summer periods was due to drift of pool invertebrates to riffles, rather than movement of riffle trout. Data for d 15 N showed that resident rainbow trout (25-33 cm standard length) in deep pools preyed upon small size classes of juvenile steelhead that were most often found in riffles or shallow pools. While many invertebrate consumers relied primarily on algal production derived from local habitats, our study shows that growth of top predators in the river is strongly linked to food webs in adjacent habitats. These results suggest a key role for emigration of aquatic prey in determining carbon flow to top predators.

The role of discharge variation in scaling of drainage area and food chain length in rivers

Despite broad recognition of the value of the goods and services provided by nature, existing tools for assessing and valuing ecosystem services often fall short of the needs and expectations of decision makers. Here we address one of the most important missing components in the current ecosystem services toolbox: a comprehensive and generalizable framework for describing and valuing water quality-related services. Water quality is often misrepresented as a final ecosystem service. We argue that it is actually an important contributor to many different services, from recreation to human health. We present a valuation approach for water quality-related services that is sensitive to different actions that affect water quality, identifies aquatic endpoints where the consequences of changing water quality on human well-being are realized, and recognizes the unique groups of beneficiaries affected by those changes. We describe the multiple biophysical and economic pathways that link actions to changes in water quality-related ecosystem goods and services and provide guidance to researchers interested in valuing these changes. Finally, we present a valuation template that integrates biophysical and economic models, links actions to changes in service provision and value estimates, and considers multiple sources of water quality-related ecosystem service values without double counting.

Human Influences on Nitrogen Removal in Lakes

River Food Chains Food chain length, or the number of organisms involved in any eat-or-be-eaten relationship, is a key property of aquatic habitats that regulates nutrient cycling, energy flow, and carbon exchange between aquatic ecosystems and the atmosphere. Investigating river food webs in North American watersheds, Sabo et al. (p. 965, published online 14 October) found discharge variation governed the link between ecosystem size (watershed area) and food chain length. River-drying truncated food chain length because intermittent streams have much higher discharge variation than perennial rivers. These results could be important for the management of rivers as drought and climate change may increase the frequency of river drying and discharge variability. The lengths of river food chains are indirectly related to drainage area and directly affected by flow and discharge. Food chain length (FCL) is a fundamental component of food web structure. Studies in a variety of ecosystems suggest that FCL is determined by energy supply, environmental stability, and/or ecosystem size, but the nature of the relationship between environmental stability and FCL, and the mechanism linking ecosystem size to FCL, remain unclear. Here we show that FCL increases with drainage area and decreases with hydrologic variability and intermittency across 36 North American rivers. Our analysis further suggests that hydrologic variability is the mechanism underlying the correlation between ecosystem size and FCL in rivers. Ecosystem size lengthens river food chains by integrating and attenuating discharge variation through stream networks, thereby enhancing environmental stability in larger river systems.

A shift of thermokarst lakes from carbon sources to sinks during the Holocene epoch

Unlucky Lakes The negative consequences of increased loading of nitrogen and phosphorus into aquatic ecosystems are well known. Management strategies aimed at reducing the sources of these excess nutrients, such as fertilizer runoff or sewage outflows, can largely mitigate the increases in nitrogen and phosphorus levels; however, it is unclear if these strategies are influencing other spects of these ecosystems. Using a global lake data set, Finlay et al. (p. 247; see the Perspective by Bernhardt) found that reducing phosphorus inputs reduced a lakes ability to export reactive nitrogen, exacerbating nitrate pollution. Successful control of phosphorus levels results in nitrogen accumulation in many of Earth’s largest lakes. [Also see Perspective by Bernhardt] Human activities have increased the availability of reactive nitrogen in many ecosystems, leading to negative impacts on human health, biodiversity, and water quality. Freshwater ecosystems, including lakes, streams, and wetlands, are a large global sink for reactive nitrogen, but factors that determine the efficacy of freshwater nitrogen removal rates are poorly known. Using a global lake data set, we show that the availability of phosphorus, a limiting nutrient, affects both annual nitrogen removal rate and efficiency. This result indicates that increased phosphorus inputs from human activities have stimulated nitrogen removal processes in many lakes. Recent management-driven reductions in phosphorus availability promote water column accumulation and export of nitrogen from large lakes, an unintended consequence of single-element management that argues for greater control of nitrogen as well as phosphorus sources.

The interactive effects of excess reactive nitrogen and climate change on aquatic ecosystems and water resources of the United States

Thermokarst lakes formed across vast regions of Siberia and Alaska during the last deglaciation and are thought to be a net source of atmospheric methane and carbon dioxide during the Holocene epoch. However, the same thermokarst lakes can also sequester carbon, and it remains uncertain whether carbon uptake by thermokarst lakes can offset their greenhouse gas emissions. Here we use field observations of Siberian permafrost exposures, radiocarbon dating and spatial analyses to quantify Holocene carbon stocks and fluxes in lake sediments overlying thawed Pleistocene-aged permafrost. We find that carbon accumulation in deep thermokarst-lake sediments since the last deglaciation is about 1.6 times larger than the mass of Pleistocene-aged permafrost carbon released as greenhouse gases when the lakes first formed. Although methane and carbon dioxide emissions following thaw lead to immediate radiative warming, carbon uptake in peat-rich sediments occurs over millennial timescales. We assess thermokarst-lake carbon feedbacks to climate with an atmospheric perturbation model and find that thermokarst basins switched from a net radiative warming to a net cooling climate effect about 5,000 years ago. High rates of Holocene carbon accumulation in 20 lake sediments (47 ± 10 grams of carbon per square metre per year; mean ± standard error) were driven by thermokarst erosion and deposition of terrestrial organic matter, by nutrient release from thawing permafrost that stimulated lake productivity and by slow decomposition in cold, anoxic lake bottoms. When lakes eventually drained, permafrost formation rapidly sequestered sediment carbon. Our estimate of about 160 petagrams of Holocene organic carbon in deep lake basins of Siberia and Alaska increases the circumpolar peat carbon pool estimate for permafrost regions by over 50 per cent (ref. 6). The carbon in perennially frozen drained lake sediments may become vulnerable to mineralization as permafrost disappears, potentially negating the climate stabilization provided by thermokarst lakes during the late Holocene.

INTRODUCED TROUT SEVER TROPHIC CONNECTIONS IN WATERSHEDS: CONSEQUENCES FOR A DECLINING AMPHIBIAN

Nearly all freshwaters and coastal zones of the US are degraded from inputs of excess reactive nitrogen (Nr), sources of which are runoff, atmospheric N deposition, and imported food and feed. Some major adverse effects include harmful algal blooms, hypoxia of fresh and coastal waters, ocean acidification, long-term harm to human health, and increased emissions of greenhouse gases. Nitrogen fluxes to coastal areas and emissions of nitrous oxide from waters have increased in response to N inputs. Denitrification and sedimentation of organic N to sediments are important processes that divert N from downstream transport. Aquatic ecosystems are particularly important denitrification hotspots. Carbon storage in sediments is enhanced by Nr, but whether carbon is permanently buried is unknown. The effect of climate change on N transport and processing in fresh and coastal waters will be felt most strongly through changes to the hydrologic cycle, whereas N loading is mostly climate-independent. Alterations in precipitation amount and dynamics will alter runoff, thereby influencing both rates of Nr inputs to aquatic ecosystems and groundwater and the water residence times that affect Nr removal within aquatic systems. Both infrastructure and climate change alter the landscape connectivity and hydrologic residence time that are essential to denitrification. While Nr inputs to and removal rates from aquatic systems are influenced by climate and management, reduction of N inputs from their source will be the most effective means to prevent or to minimize environmental and economic impacts of excess Nr to the nation’s water resources.