Christopher T. Solomon

Strong evidence for terrestrial support of zooplankton in small lakes based on stable isotopes of carbon, nitrogen, and hydrogen

Cross-ecosystem subsidies to food webs can alter metabolic balances in the receiving (subsidized) system and free the food web, or particular consumers, from the energetic constraints of local primary production. Although cross-ecosystem subsidies between terrestrial and aquatic systems have been well recognized for benthic organisms in streams, rivers, and the littoral zones of lakes, terrestrial subsidies to pelagic consumers are more difficult to demonstrate and remain controversial. Here, we adopt a unique approach by using stable isotopes of H, C, and N to estimate terrestrial support to zooplankton in two contrasting lakes. Zooplankton (Holopedium, Daphnia, and Leptodiaptomus) are comprised of ≈20–40% of organic material of terrestrial origin. These estimates are as high as, or higher than, prior measures obtained by experimentally manipulating the inorganic 13C content of these lakes to augment the small, natural contrast in 13C between terrestrial and algal photosynthesis. Our study gives credence to a growing literature, which we review here, suggesting that significant terrestrial support of pelagic crustaceans (zooplankton) is widespread.

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.

Ecosystem Consequences of Changing Inputs of Terrestrial Dissolved Organic Matter to Lakes: Current Knowledge and Future Challenges

Lake ecosystems and the services that they provide to people are profoundly influenced by dissolved organic matter derived from terrestrial plant tissues. These terrestrial dissolved organic matter (tDOM) inputs to lakes have changed substantially in recent decades, and will likely continue to change. In this paper, we first briefly review the substantial literature describing tDOM effects on lakes and ongoing changes in tDOM inputs. We then identify and provide examples of four major challenges which limit predictions about the implications of tDOM change for lakes, as follows: First, it is currently difficult to forecast future tDOM inputs for particular lakes or lake regions. Second, tDOM influences ecosystems via complex, interacting, physical-chemical-biological effects and our holistic understanding of those effects is still rudimentary. Third, non-linearities and thresholds in relationships between tDOM inputs and ecosystem processes have not been well described. Fourth, much understanding of tDOM effects is built on comparative studies across space that may not capture likely responses through time. We conclude by identifying research approaches that may be important for overcoming those challenges in order to provide policy- and management-relevant predictions about the implications of changing tDOM inputs for lakes.

Stable Isotope Turnover and Half-Life in Animal Tissues: A Literature Synthesis

Stable isotopes of carbon, nitrogen, and sulfur are used as ecological tracers for a variety of applications, such as studies of animal migrations, energy sources, and food web pathways. Yet uncertainty relating to the time period integrated by isotopic measurement of animal tissues can confound the interpretation of isotopic data. There have been a large number of experimental isotopic diet shift studies aimed at quantifying animal tissue isotopic turnover rate λ (%·day-1, often expressed as isotopic half-life, ln(2)/λ, days). Yet no studies have evaluated or summarized the many individual half-life estimates in an effort to both seek broad-scale patterns and characterize the degree of variability. Here, we collect previously published half-life estimates, examine how half-life is related to body size, and test for tissue- and taxa-varying allometric relationships. Half-life generally increases with animal body mass, and is longer in muscle and blood compared to plasma and internal organs. Half-life was longest in ecotherms, followed by mammals, and finally birds. For ectotherms, different taxa-tissue combinations had similar allometric slopes that generally matched predictions of metabolic theory. Half-life for ectotherms can be approximated as: ln (half-life) = 0.22*ln (body mass) + group-specific intercept; n = 261, p<0.0001, r2 = 0.63. For endothermic groups, relationships with body mass were weak and model slopes and intercepts were heterogeneous. While isotopic half-life can be approximated using simple allometric relationships for some taxa and tissue types, there is also a high degree of unexplained variation in our models. Our study highlights several strong and general patterns, though accurate prediction of isotopic half-life from readily available variables such as animal body mass remains elusive.

quantitative approaches to the analysis OF stable isotope food web data

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.

Understanding Regional Change: A Comparison of Two Lake Districts

ABSTRACT We compared long-term change in two lake districts, one in a forested rural setting and the other in an urbanizing agricultural region, using lakes as sentinel ecosystems. Human population growth and land-use change are important drivers of ecosystem change in both regions. Biotic changes such as habitat loss, species invasions, and poorer fishing were prevalent in the rural region, and lake hydrology and biogeochemistry responded to climate trends and landscape position. Similar biotic changes occurred in the urbanizing agricultural region, where human-caused changes in hydrology and biogeochemistry had conspicuous effects. Feedbacks among ecosystem dynamics, human uses, economics, social dynamics, and policy and practice are fundamental to understanding change in these lake districts. Sustained support for interdisciplinary collaboration is essential to build understanding of regional change.

Terrestrial carbon is a resource, but not a subsidy, for lake zooplankton

Inputs of terrestrial organic carbon (t-OC) into lakes are often considered a resource subsidy for aquatic consumer production. Although there is evidence that terrestrial carbon can be incorporated into the tissues of aquatic consumers, its ability to enhance consumer production has been debated. Our research aims to evaluate the net effect of t-OC input on zooplankton. We used a survey of zooplankton production and resource use in ten lakes along a naturally occurring gradient of t-OC concentration to address these questions. Total and group-specific zooplankton production was negatively related to t-OC. Residual variation in zooplankton production that was not explained by t-OC was negatively related to terrestrial resource use (allochthony) by zooplankton. These results challenge the designation of terrestrial carbon as a resource subsidy; rather, the negative effect of reduced light penetration on the amount of suitable habitat and the low resource quality of t-OC appear to diminish zooplankton production. Our findings suggest that ongoing continental-scale increases in t-OC concentrations of lakes will likely have negative impacts on the productivity of aquatic food webs.

Subsidy or Subtraction: How Do Terrestrial Inputs Influence Consumer Production in Lakes?

Abstract Cross-ecosystem fluxes are ubiquitous in food webs and are generally thought of as subsidies to consumer populations. Yet external or allochthonous inputs may in fact have complex and habitat-specific effects on recipient ecosystems. In lakes, terrestrial inputs of organic carbon contribute to basal resource availability, but can also reduce resource availability via shading effects on phytoplankton and periphyton. Terrestrial inputs might therefore either subsidise or subtract from consumer production. We developed and parameterised a simple model to explore this idea. The model estimates basal resource supply and consumer production given lake-level characteristics including total phosphorus (TP) and dissolved organic carbon (DOC) concentration, and consumer-level characteristics including resource preferences and growth efficiencies. Terrestrial inputs diminished primary production and total basal resource supply at the whole-lake level, except in ultra-oligotrophic systems. However, this system-level generalisation masked complex habitat-specific effects. In the pelagic zone, dissolved and particulate terrestrial carbon inputs were available to zooplankton via several food web pathways. Consequently, zooplankton production usually increased with terrestrial inputs, even as total whole-lake resource availability decreased. In contrast, in the benthic zone the dominant, dissolved portion of the terrestrial carbon load had predominantly negative effects on resource availability via shading of periphyton. Consequently, terrestrial inputs always decreased zoobenthic production except under extreme and unrealistic parameterisations of the model. Appreciating the complex and habitat-specific effects of allochthonous inputs may be essential for resolving the effects of cross-habitat fluxes on consumers in lakes and other food webs.

Distribution and community-level effects of the Chinese mystery snail (Bellamya chinensis) in northern Wisconsin lakes

Managing invasive species requires information about their distributions and potential effects, but community-level impacts of invasive animals remain poorly understood. The Chinese mystery snail (Bellamya chinensis) is a large invasive gastropod that achieves high densities in waters across North America, yet little is known about its ecological significance in invaded systems. We surveyed 44 lakes to describe the patterns and determinants of B. chinensis distributions in northern Wisconsin, USA, and to assess the likelihood of effects on native snail communities in the invaded systems. B. chinensis was widespread among surveyed lakes (21 of 42 lakes with snails) and its occurrence was correlated with indicators of lake productivity and anthropogenic dispersal vectors (boat landings, distance to population centers, shoreline housing density). Some native snail species tended not to occur at sites where B. chinensis was abundant; among these was Lymnaea stagnalis, which suffered reduced survival in the presence of B. chinensis in a recently published mesocosm study. However, there was no difference in overall snail assemblage structure at either the site or lake level as a function of B. chinensis presence or abundance. Lake occurrences of many snail species have apparently been lost over time, but a comparison to a 1930s survey showed that there was no increased likelihood of species loss in lakes invaded by B. chinensis (or by the invasive crayfish Orconectes rusticus). Although B. chinensis is widespread and sometimes abundant in northern Wisconsin lakes, it does not appear to have strong systematic impacts on native snail assemblages.

Sediment size and nutrients regulate denitrification in a tropical stream

Abstract Landuse changes might alter N cycling in tropical aquatic ecosystems, but understanding of N cycling in tropical streams is limited. We measured actual and potential denitrification rates during the dry season in Río Las Marías, a 4th-order Andean piedmont stream in Venezuela. Our objectives were to describe spatial and temporal variation in denitrification, quantify the effects of nutrient availability and substratum particle size on denitrification, and explore potential effects of anthropogenic sedimentation on denitrification. In 4 experiments, actual and potential denitrification rates ranged from 0 to 160 and from 0 to 740 μg N2O-N m−2 h−1, respectively. Rates were distributed approximately log-normally because of spatial variation. During a 1-mo period, actual denitrification rates decreased exponentially from 37 ± 39 to 5 ± 7 μg N2O-N m−2 h−1 (mean ± SD), probably because of a decline in water-column NO3-N concentration from 41 ± 14 to 12 ± 3 μg NO3-N/L. The texture (particle size) of stream substrata markedly affected denitrification rates. Actual rates were low in cobble, gravel, and fine sediments (<5 mm), but in fine sediments, rates increased in response to addition of excess NO3-N and organic C. In a 3-km stream reach, actual (but not potential) denitrification rates increased with the proportion of fine sediments (<2 mm) in mixed substrata. This increase was nonlinear, and the threshold value occurred at 37% fine particles, above which actual denitrification rates were almost always high. An experiment simulating the effects of anthropogenic sedimentation showed that topsoil inputs resulted in denitrification rates ∼8× higher than rates in trials where excess NO3-N and organic C were supplied. Denitrification is a small but potentially significant sink for available N in this N-limited system. Anthropogenic sedimentation associated with landuse change might significantly increase denitrification rates in streams.