James M. Hood

Fish distributions and nutrient cycling in streams: can fish create biogeochemical hotspots?

Rates of biogeochemical processes often vary widely in space and time, and characterizing this variation is critical for understanding ecosystem functioning. In streams, spatial hotspots of nutrient transformations are generally attributed to physical and microbial processes. Here we examine the potential for heterogeneous distributions of fish to generate hotspots of nutrient recycling. We measured nitrogen (N) and phosphorus (P) excretion rates of 47 species of fish in an N-limited Neotropical stream, and we combined these data with population densities in each of 49 stream channel units to estimate unit- and reach-scale nutrient recycling. Species varied widely in rates of N and P excretion as well as excreted N:P ratios (6-176 molar). At the reach scale, fish excretion could meet >75% of ecosystem demand for dissolved inorganic N and turn over the ambient NH4 pool in <0.3 km. Areal N excretion estimates varied 47-fold among channel units, suggesting that fish distributions could influence local N availability. P excretion rates varied 14-fold among units but were low relative to ambient concentrations. Spatial variation in aggregate nutrient excretion by fish reflected the effects of habitat characteristics (depth, water velocity) on community structure (body size, density, species composition), and the preference of large-bodied species for deep runs was particularly important. We conclude that the spatial distribution of fish could indeed create hotspots of nutrient recycling during the dry season in this species-rich tropical stream. The prevalence of patchy distributions of stream fish and invertebrates suggests that hotspots of consumer nutrient recycling may often occur in stream ecosystems.

INTERACTIONS BETWEEN HERBIVOROUS FISHES AND LIMITING NUTRIENTS IN A TROPICAL STREAM ECOSYSTEM

Ecologists have long been interested in understanding the strengths of con- sumer and resource limitation in influencing communities. Here we ask three questions concerning the relative importance of nutrients and grazing fishes to primary producers of a tropical Andean stream: (1) Are stream algae nutrient limited? (2) Are top-down and bottom-up forces of dual importance in limiting primary producers? (3) Do grazing fishes modulate the degree of resource limitation? We obtained several lines of evidence suggesting that Andean stream algae are nitrogen limited. Addition of nitrogen in flow-through channels resulted in major increases in algal standing crop, whereas there were no measurable effects of phosphorus enrichment. Inter- estingly, the N2-fixing cyanobacteria Anabaena was one of the taxa that responded most dramatically to the addition of nitrogen. Moreover, nutrient uptake rates were significantly higher for inorganic nitrogen (NO3-N and NH4-N) compared to phosphorus (PO4-P). Nutrients and the presence of grazing fishes were manipulated simultaneously in a series of experiments by using nutrient-diffusing substrates in fish exclusions vs. open cages accessible to the natural fish assemblage. We observed strong effects of both nitrogen addition and consumers on algal standing crop, although consumer limitation was found to be of considerably greater magnitude than resource limitation in influencing algal biomass and composition. Finally, the degree of resource limitation varied as a consequence of grazing fishes. Experiments examining nutrient limitation in the presence and absence of fishes showed that the response to nitrogen enrichment was significantly greater on sub- strates accessible to natural fish assemblages compared to substrates where grazing fishes were excluded. These experiments demonstrate simultaneous and interactive effects of top- down and bottom-up factors in limiting primary producers of tropical Andean streams. Whereas other studies have shown that consumers affect nutrient supply in ecosystems, our findings suggest that consumers can play an important role in influencing nutrient

Impacts of Warming on the Structure and Functioning of Aquatic Communities : Individual-to Ecosystem-Level Responses

Environmental warming is predicted to rise dramatically over the next century, yet few studies have investigated its effects in natural, multi-species systems. We present data collated over an 8-year period from a catchment of geothermally heated streams in Iceland, which acts as a natural experiment on the effects of warming across different organisational levels and spatiotemporal scales. Body sizes and population biomasses of individual species responded strongly to temperature, with some providing evidence to support temperature size rules. Macroinvertebrate and meiofaunal community composition also changed dramatically across the thermal gradient. Interactions within the warm streams in particular were characterised by food chains linking algae to snails to the apex predator, brown trout These chains were missing from the colder systems, where snails were replaced by much smaller herbivores and invertebrate omnivores were the top predators. Trout were also subsidised by terrestrial invertebrate prey, which could have an effect analogous to apparent competition within the aquatic prey assemblage. Top-down effects by snails on diatoms were stronger in the warmer streams, which could account for a shallowing of mass-abundance slopes across the community. This may indicate reduced energy transfer efficiency from resources to consumers in the warmer systems and/or a change in predator-prey mass ratios. All the ecosystem process rates investigated increased with temperature, but with differing thermal sensitivities, with important implications for overall ecosystem functioning (e.g. creating potential imbalances in elemental fluxes). Ecosystem respiration rose rapidly with temperature, leading to increased heterotrophy. There were also indications that food web stability may be lower in the warmer streams.

Nutrient recycling by two phosphorus-rich grazing catfish: the potential for phosphorus-limitation of fish growth

In ecosystems where excretion by fish is a major flux of nutrients, the nitrogen (N) to phosphorus (P) ratio released by fish can be important in shaping patterns of algal biomass, community composition, primary production, and nutrient limitation. Demand for N and P as well as energy influences N/P excretion ratios and has broad implications in ecosystems where nutrient recycling by fishes is substantial. Bioenergetics and stoichiometric models predict that natural fish populations are generally energy-limited and therefore N/P recycling by fishes is relatively invariant. Yet, the potential for P limitation of growth has not been examined in herbivorous fishes, which are common in many aquatic habitats. We examined N/P excretion ratios and P demand in two P-rich herbivorous catfishes of the family Loricariidae, Ancistrus triradiatus (hereafter Ancistrus) and Chaetostoma milesi (hereafter Chaetostoma). Both fishes are common grazers in the Andean piedmont region of Venezuela where we conducted this study. Mass balance (MB) models indicate that these fishes have a high P demand. In fact, our Ancistrus’ P MB model predicted negative P excretion rates, indicating that Ancistrus did not consume enough P to meet its P demand for growth. Direct measurement of excretion rates showed positive, but very low P excretion rates and high N/P excretion ratios for both taxa. To obtain measured P excretion rates of Ancistrus from the MB model, gross growth efficiency must be reduced by 90%. Our results suggest that growth rates of both of these herbivorous and P-rich fish are likely P-limited. If P limitation of growth is common among herbivorous fish populations, herbivorous fishes recycle likely at high N/P ratios and act to diminish the quality of their food.

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits--growth, respiration, body size, and elemental content--that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population-level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature-nutrient interactions influence processes at the whole-ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature-nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.

Light-mediated thresholds in stream-water nutrient composition in a river network

The elemental composition of solutes transported by rivers reflects combined influences of surrounding watersheds and transformations within stream networks, yet comparatively little is known about downstream changes in effects of watershed loading vs. in-channel processes. In the forested watershed of a river under a mediterranean hydrologic regime, we examined the influence of longitudinal changes in environmental conditions on water-column nutrient composition during summer base flow across a network of sites ranging from strongly heterotrophic headwater streams to larger, more autotrophic sites downstream. Small streams (0.1-10 km2 watershed area) had longitudinally similar nutrient concentration and composition with low (approximately 2) dissolved nitrogen (N) to phosphorus (P) ratios. Abrupt deviations from this pattern were observed in larger streams with watershed areas > 100 km2 where insolation and algal abundance and production rapidly increased. Downstream, phosphorus and silica concentrations decreased by > 50% compared to headwater streams, and dissolved organic carbon and nitrogen increased by approximately 3-6 times. Decreasing dissolved P and increasing dissolved N raised stream-water N:P to 46 at the most downstream sites, suggesting a transition from N limitation in headwaters to potential P limitation in larger channels. We hypothesize that these changes were mediated by increasing algal photosynthesis and N fixation by benthic algal assemblages, which, in response to increasing light availability, strongly altered stream-water nutrient concentration and stoichiometry in larger streams and rivers.

Climate change and geothermal ecosystems: natural laboratories, sentinel systems, and future refugia

Understanding and predicting how global warming affects the structure and functioning of natural ecosystems is a key challenge of the 21st century. Isolated laboratory and field experiments testing global change hypotheses have been criticized for being too small-scale and overly simplistic, whereas surveys are inferential and often confound temperature with other drivers. Research that utilizes natural thermal gradients offers a more promising approach and geothermal ecosystems in particular, which span a range of temperatures within a single biogeographic area, allow us to take the laboratory into nature rather than vice versa. By isolating temperature from other drivers, its ecological effects can be quantified without any loss of realism, and transient and equilibrial responses can be measured in the same system across scales that are not feasible using other empirical methods. Embedding manipulative experiments within geothermal gradients is an especially powerful approach, informing us to what extent small-scale experiments can predict the future behaviour of real ecosystems. Geothermal areas also act as sentinel systems by tracking responses of ecological networks to warming and helping to maintain ecosystem functioning in a changing landscape by providing sources of organisms that are preadapted to different climatic conditions. Here, we highlight the emerging use of geothermal systems in climate change research, identify novel research avenues, and assess their roles for catalysing our understanding of ecological and evolutionary responses to global warming.

Diet Mixing: Do Animals Integrate Growth or Resources across Temporal Heterogeneity?

Animals commonly experience spatial and temporal variation in resource quality, thus experiencing temporally variable diets. Methods for scaling up growth in component patches to long‐term growth across heterogeneity are seldom explicitly considered. Long‐term growth is sometimes considered to be a weighted average of growth rates on component diets (growth integration). However, if animals integrate resources across high‐ and low‐quality diets, their long‐term growth may be greater than predicted from diet‐specific growth rates (resource integration). We measured biomass growth rates of seven Daphnia species exposed to different types of diel variation in algal phosphorus (P) content. Support for resource integration was found for four of the seven species, which achieved near maximal growth when high‐P food was available for at least 12 h. In contrast, no support for resource integration was found for the other three species. These three species achieved only one‐half maximal growth rate under the same conditions and could be considered growth integrators. The type of integration could be predicted from the degree of stoichiometric homeostasis. Species with weak homeostatic regulation exhibited a capacity for resource integration. Resource integrators should have an advantage in heterogeneous environments.

Coupling of dietary phosphorus and growth across diverse fish taxa: a meta‐analysis of experimental aquaculture studies

Imbalances in phosphorus (P) intake relative to demand negatively affect animal growth, but their consequences are less understood for vertebrates, in which bone represents a significant and potentially flexible pool of P. Flexibility in body-P content could buffer vertebrates from the effects of imbalances between P intake and demand, reducing the likelihood of a sharp stoichiometric “knife-edge” in the relationship between growth rate and diet-P level. We conducted a meta-analysis of published aquaculture experiments that tested effects of diet %P on fish growth rate (49 studies, 28 species) and body-P content (27 of the studies in the main data set, 20 species). Our meta-analysis revealed significant P limitation of growth, as well as significant negative effects of excess P on growth rate. Diet-P thresholds for these effects occurred at ecologically relevant levels (optimal diet-P of 1.2% ± 0.45%, mean ± SD, under experimental conditions of high ration). Finally, the analysis also suggested a pattern ...

Does N2 fixation amplify the temperature dependence of ecosystem metabolism

Variation in resource supply can cause variation in temperature dependences of metabolic processes (e.g., photosynthesis and respiration). Understanding such divergence is particularly important when using metabolic theory to predict ecosystem responses to climate warming. Few studies, however, have assessed the effect of temperature-resource interactions on metabolic processes, particularly in cases where the supply of limiting resources exhibits temperature dependence. We investigated the responses of biomass accrual, gross primary production (GPP), community respiration (CR), and N2 fixation to warming during biofilm development in a streamside channel experiment. Areal rates of GPP, CR, biomass accrual, and N2 fixation scaled positively with temperature, showing a 32- to 71-fold range across the temperature gradient (approximately 7 degrees-24 degrees C). Areal N2-fixation rates exhibited apparent activation energies (1.5-2.0 eV; 1 eV = approximately 1.6 x 10(-19) J) approximating the activation energy of the nitrogenase reaction. In contrast, mean apparent activation energies for areal rates of GPP (2.1-2.2 eV) and CR (1.6-1.9 eV) were 6.5- and 2.7-fold higher than estimates based on metabolic theory predictions (i.e., 0.32 and 0.65 eV, respectively) and did not significantly differ from the apparent activation energy observed for N2 fixation. Mass-specific activation energies for N2 fixation (1.4-1.6 eV), GPP (0.3-0.5 eV), and CR (no observed temperature relationship) were near or lower than theoretical predictions. We attribute the divergence of areal activation energies from those predicted by metabolic theory to increases in N2 fixation with temperature, leading to amplified temperature dependences of biomass accrual and areal rates of GPP and R. Such interactions between temperature dependences must be incorporated into metabolic models to improve predictions of ecosystem responses to climate change.