William B. Richardson

AGRICULTURAL PONDS SUPPORT AMPHIBIAN POPULATIONS

In some agricultural regions, natural wetlands are scarce, and constructed agricultural ponds may represent important alternative breeding habitats for amphibians. Properly managed, these agricultural ponds may effectively increase the total amount of breeding habitat and help to sustain populations. We studied small, constructed agricultural ponds in southeastern Minnesota to assess their value as amphibian breeding sites. Our study examined habitat factors associated with amphibian reproduction at two spatial scales: the pond and the landscape surrounding the pond. We found that small agricultural ponds in southeastern Minnesota provided breeding habitat for at least 10 species of amphibians. Species richness and multispecies reproductive success were more closely associated with characteristics of the pond (water quality, vegetation, and predators) compared with char- acteristics of the surrounding landscape, but individual species were associated with both pond and landscape variables. Ponds surrounded by row crops had similar species richness and reproductive success compared with natural wetlands and ponds surrounded by non- grazed pasture. Ponds used for watering livestock had elevated concentrations of phos- phorus, higher turbidity, and a trend toward reduced amphibian reproductive success. Spe- cies richness was highest in small ponds, ponds with lower total nitrogen concentrations, tiger salamanders ( Ambystoma tigrinum) present, and lacking fish. Multispecies reproduc- tive success was best in ponds with lower total nitrogen concentrations, less emergent vegetation, and lacking fish. Habitat factors associated with higher reproductive success varied among individual species. We conclude that small, constructed farm ponds, properly managed, may help sustain amphibian populations in landscapes where natural wetland habitat is rare. We recommend management actions such as limiting livestock access to the pond to improve water quality, reducing nitrogen input, and avoiding the introduction of fish.

Past, Present, and Future Concepts in Large River Ecology How rivers function and how human activities influence river processes

This article reviews the current concepts on how flowing water, or lotic, systems function and suggest ways to expand these concepts to increase understanding of large river systems. Topics covered include the following: past and present approaches to physical and biological concepts of stream organization; the river-continuum concept; the flood-pulse concept; deficiencies of the current hypotheses (linear versis hierachical, physical verses biological control, equilibrium versus disequilibrium); future directions for large river research. 63 refs., 4 figs.

Nitrification in the Upper Mississippi River: patterns, controls, and contribution to the NO3− budget

Abstract We measured nitrification rates in sediment samples collected from a variety of aquatic habitats in Navigation Pool 8 of the Upper Mississippi River (UMR) 7 times between May 2000 and October 2001. We also conducted nutrient-enrichment experiments and analyzed vertical profiles of sediment to determine factors regulating nitrification. Nitrification rates were relatively high compared to other ecosystems (ranging from 0–8.25 μg N cm−2 h−1) and exhibited significant temporal and spatial patterns. Nitrification rates were greatest during the summer and spring compared to autumn and winter (ANOVA, p < 0.05) and were greater in contiguous backwater and impounded habitats compared to main and side-channel habitats (p < 0.05). Regression analysis indicated that nitrification rates were weakly (r2 = 0.18, p < 0.0001) related to temperature and exchangeable NH4+ of the sediment. However, nutrient-enrichment experiments showed that NH4+ availability did not limit nitrification in 3 sediment types with variable organic matter. Vertical profiles of sediment cores demonstrated that oxygen concentration and nitrification had similar patterns suggesting that nitrification may be limited by oxygen penetration into sediments. We conclude that temperature and sediment NH4+ can be useful for predicting broad-scale temporal and spatial nitrification patterns, respectively, but oxygen penetration into the sediments likely regulates nitrification rates in much of the UMR. Overall, we estimated that nitrification produces 6982 mt N/y of NO3− or 7% of the total annual NO3− budget.

Nitrogen and phosphorus in the Upper Mississippi River: transport, processing, and effects on the river ecosystem

Existing research on nutrients (nitrogen and phosphorus) in the Upper Mississippi River (UMR) can be organized into the following categories: (1) Long-term changes in nutrient concentrations and export, and their causes; (2) Nutrient cycling within the river; (3) Spatial and temporal patterns of river nutrient concentrations; (4) Effects of elevated nutrient concentrations on the river; and (5) Actions to reduce river nutrient concentrations and flux. Nutrient concentration and flux in the Mississippi River have increased substantially over the last century because of changes in land use, climate, hydrology, and river management and engineering. As in other large floodplain rivers, rates of processes that cycle nitrogen and phosphorus in the UMR exhibit pronounced spatial and temporal heterogeneity because of the complex morphology of the river. This spatial variability in nutrient processing creates clear spatial patterns in nutrient concentrations. For example, nitrate concentrations generally are much lower in off-channel areas than in the main channel. The specifics of in-river nutrient cycling and the effects of high rates of nutrient input on UMR have been less studied than the factors affecting nutrient input to the river and transport to the Gulf of Mexico, and important questions concerning nutrient cycling in the UMR remain. Eutrophication and resulting changes in river productivity have only recently been investigated the UMR. These recent studies indicate that the high nutrient concentrations in the river may affect community composition of aquatic vegetation (e.g., the abundance of filamentous algae and duckweeds), dissolved oxygen concentrations in off-channel areas, and the abundance of cyanobacteria. Actions to reduce nutrient input to the river include changes in land-use practices, wetland restoration, and hydrological modifications to the river. Evidence suggests that most of the above methods can contribute to reducing nutrient concentration in, and transport by, the UMR, but the impacts of mitigation efforts will likely be only slowly realized.

EFFECTS OF AMMONIA ON JUVENILE UNIONID MUSSELS ( LAMPSILIS CARDIUM ) IN LABORATORY SEDIMENT TOXICITY TESTS

Ammonia is a relatively toxic compound generated in water and sediments by heterotrophic bacteria and accumulates in sediments and pore water. Recent data suggest that unionid mussels are sensitive to un-ionized ammonia (NH3) relative to other organisms. Existing sediment exposure systems are not suitable for ammonia toxicity studies with juvenile unionids; thus, we modified a system to expose juveniles to ammonia that was continuously infused into sediments. This system maintained consistent concentrations of ammonia in pore water up to 10 d. Juvenile Lampsilis cardium mussels were exposed to NH3 in pore water in replicate 96-h and 10-d sediment toxicity tests. The 96-h median lethal concentrations (LC50s) were 127 and 165 microg NH3-N/L, and the 10-d LC50s were 93 and 140 microg NH3-N/L. The median effective concentrations (EC50s) (based on the proportion affected, including dead and inactive mussels) were 73 and 119 microg NH3-N/L in the 96-h tests and 71 and 99 microg NH3-N/L in the 10-d tests. Growth rate was substantially reduced at concentrations between 31 and 76 microg NH3-N/L. The lethality results (when expressed as total ammonia) are about one-half the acute national water quality criteria for total ammonia, suggesting that existing criteria may not protect juvenile unionids.

Variability and regulation of denitrification in an Upper Mississippi River backwater

Abstract Sediments in the backwaters of the Upper Mississippi River (UMR) are highly organic and provide an optimal environment for N removal. We monitored an 8.6-ha UMR backwater site near La Crosse, Wisconsin, for nearly 3 y to assess temporal variability, seasonal trends, and the factors regulating denitrification. We measured rates of unamended denitrification (DEN) and denitrification enzyme activity (DEA) rates at ambient temperature and DEA at 30°C (DEA30). Seasonal mean (±1 SE) DEN rates ranged from 0.041 ± 0.015 to 0.47 ± 0.23 μg N cm−2 h−1 and were highest in winter and lowest in autumn. Seasonal rates of DEA exhibited a different pattern with the highest rates in summer (25.6 ± 3.4 μg N cm−2 h−1) and the lowest rates in winter (10.6 ± 2.1 μg N cm−2 h−1). The overall mean DEA30 rate was 31.0 ± 1.9 μg N cm−2 h−1 but showed no significant seasonal pattern. Short-term (weekly) and seasonal variability exhibited by rates of DEN and DEA were best explained by water-column NO3− concentration and temperature, respectively. No environmental variables explained a significant amount of variability in DEA30. Our results suggest that nutrient (i.e., NO3−) availability and temperature are both regulators of denitrification, with NO3− concentration being the most important limiting factor in this system. The high DEN rates during winter were in response to elevated NO3− concentrations resulting from a chain reaction beginning with algal blooms creating oxic conditions that stimulated nitrification. Increasing hydrological connectivity in large rivers as a river management tool to reduce N flux to downstream areas may be beneficial.

Effects of zebra mussels on food webs: interactions with juvenile bluegill and water residence time

We evaluated how water residence time mediated the impact of zebramussels Dreissena polymorpha and bluegill sunfish Lepomis macrochirus on experimental food webs established in1100-l outdoor mesocosms. Water residence time was manipulated asa surrogate for seston resupply – a critical variable affectinggrowth and survival of suspension-feeding invertebrates. We used a2×2×2 factorial experimental design witheight treatment combinations (3 replicates/treatment) including thepresence or absence of Dreissena (2000 per m2), juvenilebluegill (40 per mesocosm), and short (1100 l per d) or long (220 lper d) water residence time. Measures of seston concentration(chlorophyll a, turbidity and suspended solids) were greaterin the short- compared to long water-residence mesocosms, butintermediate in short water-residence mesocosms containing Dreissena. Abundance of rotifers (Keratella and Polyarthra) was reduced in Dreissena mesocosms and elevatedin short residence time mesocosms. Cladocera abundance, in general,was unaffected by the presence of Dreissena; densities werehigher in short-residence time mesocosms, and reduced in thepresence of Lepomis. The growth of juvenile Lepomiswere unaffected by Dreissena because of abundant benthicfood. The final total mass of Dreissena was significantlygreater in short- than long-residence mesocosms. Impacts of Dreissena on planktonic food webs may not only depend on thedensity of zebra mussels but also on the residence time of thesurrounding water and the resupply of seston.

Human activities cause distinct dissolved organic matter composition across freshwater ecosystems.

Dissolved organic matter (DOM) composition in freshwater ecosystems is influenced by the interactions among physical, chemical, and biological processes that are controlled, at one level, by watershed landscape, hydrology, and their connections. Against this environmental template, humans may strongly influence DOM composition. Yet, we lack a comprehensive understanding of DOM composition variation across freshwater ecosystems differentially affected by human activity. Using optical properties, we described DOM variation across five ecosystem groups of the Laurentian Great Lakes region: large lakes, Kawartha Lakes, Experimental Lakes Area, urban stormwater ponds, and rivers (n = 184 sites). We determined how between ecosystem variation in DOM composition related to watershed size, land use and cover, water quality measures (conductivity, dissolved organic carbon (DOC), nutrient concentration, chlorophyll a), and human population density. The five freshwater ecosystem groups had distinctive DOM composition from each other. These significant differences were not explained completely through differences in watershed size nor spatial autocorrelation. Instead, multivariate partial least squares regression showed that DOM composition was related to differences in human impact across freshwater ecosystems. In particular, urban/developed watersheds with higher human population densities had a unique DOM composition with a clear anthropogenic influence that was distinct from DOM composition in natural land cover and/or agricultural watersheds. This nonagricultural, human developed impact on aquatic DOM was most evident through increased levels of a microbial, humic-like parallel factor analysis component (C6). Lotic and lentic ecosystems with low human population densities had DOM compositions more typical of clear water to humic-rich freshwater ecosystems but C6 was only present at trace to background levels. Consequently, humans are strongly altering the quality of DOM in waters nearby or flowing through highly populated areas, which may alter carbon cycles in anthropogenically disturbed ecosystems at broad scales.

EFFECTS OF PORE-WATER AMMONIA ON IN SITU SURVIVAL AND GROWTH OF JUVENILE MUSSELS (LAMPSILIS CARDIUM ) IN THE ST. CROIX RIVERWAY, WISCONSIN, USA

We conducted a series of in situ tests to evaluate the effects of pore-water ammonia on juvenile Lampsilis cardium in the St. Croix River (WI, USA). Threats to this river and its associated unionid fauna have accelerated in recent years because of its proximity to Minneapolis-St. Paul, Minnesota, USA. In 2000, caged juveniles were exposed to sediments and overlying water at 12 sites for 10 d. Survival and growth of juveniles was significantly different between sediment (mean, 47%) and water column (mean, 86%) exposures; however, these effects were unrelated to pore-water ammonia. During 2001, juveniles were exposed to sediments for 4, 10, and 28 d. Pore-water ammonia concentrations ranged from 0.3 to 62.0 microg NH3-N/L in sediments and from 0.5 to 140.8 microg NH3-N/L within exposure chambers. Survival (mean, 45, 28, and 41% at 4, 10, and 28 d, respectively) and growth (range, 3-45 microm/d) of juveniles were highly variable and generally unrelated to ammonia concentrations. Although laboratory studies have shown unionids to be quite sensitive to ammonia, further research is needed to identify the route(s) of ammonia exposure in unionids and to understand the factors that contribute to the spatial variability of ammonia in rivers.

Zebra mussels (Dreissena polymorpha) limit food for larval fish (Pimephales promelas) in turbulent systems: a bioenergetics analysis*

We conducted a factorial experiment, in outdoor mesocosms, on the effects of zebra mussels and water column mixing (i.e., turbulence) on the diet, growth, and survival of larval fathead minnows (Pimephales promelas). Significant (P< 0.05) larval mortality occurred by the end of the experiment with the highest mortality (90%) occurring in the presence of both turbulence and zebra mussels, whereas mortality was 37% in treatment with turbulence and 17% and 18% in the zebra mussels treatment, and the control, respectively. The size of individual fish was significantly different among treatments at the end of the experiment and was inversely related to survival. Levels of trophic resources (i.e., phyto and zooplankton) varied among treatments and were treatment specific. Turbulent mixing facilitated removal of phytoplankton by zebra mussels by making the entire water column of the tanks available to these benthic filter feeders. Early in the experiment (Day = 0 to 14) the physical process of turbulent mixing likely caused a reduction in standing stocks of zooplankton. The interactive effect of turbulence and mussels reduced copepod and rotifer stocks, through physical processes and through filtration by zebra mussels, relative to the turbulence treatment. The reductions in the number of total zooplankton in the turbulent mixing mesocosms and the further reduction of rotifer and copepod in the turbulence and mussels treatment coincided with a period of increased reliance of larval fathead minnows on these prey. Estimates of consumption from bioenergetics modeling and measured prey standing stocks indicated caloric resources of suitable prey in turbulence treatments during the early weeks of the experiment were insufficient to prevent starvation. Early mortality in the turbulence and mussels treatment likely released surviving fish from intense intraspecific competition and resulted in higher individual growth rates. A combination of high abundance of zebra mussels in an environment with a well-mixed water column can have significant effects on larval fish survival and growth.