Brian C. Weidel

Early Warnings of Regime Shifts: A Whole-Ecosystem Experiment

High-frequency monitoring of manipulated and reference lakes enabled early detection of subsequent catastrophic regime shift. Catastrophic ecological regime shifts may be announced in advance by statistical early warning signals such as slowing return rates from perturbation and rising variance. The theoretical background for these indicators is rich, but real-world tests are rare, especially for whole ecosystems. We tested the hypothesis that these statistics would be early warning signals for an experimentally induced regime shift in an aquatic food web. We gradually added top predators to a lake over 3 years to destabilize its food web. An adjacent lake was monitored simultaneously as a reference ecosystem. Warning signals of a regime shift were evident in the manipulated lake during reorganization of the food web more than a year before the food web transition was complete, corroborating theory for leading indicators of ecological regime shifts.

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.

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.

Overcompensatory response of a smallmouth bass (Micropterus dolomieu) population to harvest: release from competition?

An intensive seven-year removal of adult, juvenile, and young-of-the-year smallmouth bass (Micropterus dolo- mieu) from a north temperate lake (Little Moose Lake, New York, USA) resulted in an increase in overall population abundance, primarily due to increased abundance of immature individuals. We developed a density-dependent, stage-struc- tured model to examine conditions under which population control through harvest could result in the increase of a tar- geted species. Parameter values were derived from a 54-year data set collected from another north temperate lake (Lake Opeongo, Ontario, Canada) smallmouth bass population. Sensitivity analyses identified the demographic conditions that could lead to increased abundance in response to harvest. An increase in population abundance with harvest was most likely to occur when either (i) per capita recruitment at low levels of spawner abundance was large, juvenile survivorship was high, and maturation of age-4 and older juveniles was moderately high or (ii) per capita recruitment at low levels of spawner abundance was slightly lower, yet the maturation rate of age-3 juveniles and adult survivorship were high. Our modeling results together with empirical evidence further demonstrate the importance of overcompensation as a substantial factor to consider in efforts to regulate population abundance through harvest.

Littoral Fish Community Response to Smallmouth Bass Removal from an Adirondack Lake

Large-scale observational studies in eastern Canada and the northeastern USA have concluded that introduced littoral predators are responsible for reductions in native fish diversity and abundance. To determine whether nonnative predator removal could increase native littoral fish abundance, we removed 47,682 smallmouth bass Micropterus dolomieu from a 271-ha Adirondack lake during a 6-year period. Two years after removal began, habitat-stratified snorkel surveys indicated a greater than 90% reduction in smallmouth bass abundance. The relative abundances of six native littoral species increased (4-90 times preremoval abundances) within 2 years of smallmouth bass removal. Decreased relative predation risk during the experiment reflected the reduction in littoral predators and identified seasonal differences in nearshore predation risk. The smallmouth bass population was resilient to removal, producing strong year-classes throughout the experiment. Mechanical removal was successful at decreasing smallmouth bass abundance and increasing native fish abundance, but removal must be conducted on a yearly basis to maintain low smallmouth bass population abundance. Our results provide experimental evidence regarding the need to prevent littoral predator introductions in Adirondack waters and offer support for nonnative control wherever native fish species conservation is a management priority.

Influence of Variable Interannual Summer Water Temperatures on Brook Trout Growth, Consumption, Reproduction, and Mortality in an Unstratified Adirondack Lake

Abstract Stressful water temperatures negatively affect physiological processes in fishes, yet evidence for how elevated temperatures influence population-level characteristics is rare. An 8-year field study of brook trout Salvelinus fontinalis in an unstratified Adirondack lake revealed that an aggregate measure of chronically stressful summer water temperatures strongly influenced brook trout population-level characteristics. We quantified chronic thermal stress using the cumulative degree-days over which bottom temperatures exceeded a range of thresholds (18–22°C) and found that the strongest relationships were with cumulative degree-days over 20°C (DD>20). Across years with similar brook trout densities, warmer summer water temperatures resulted in decreased brook trout growth, but growth was not reduced in a year with high water temperatures and low density. Maximum stomach fullness was negatively related to water temperature. Reproductive activity was negatively correlated with stressful summer ther...