David B. Bunnell

Linking Landscapes and Food Webs: Effects of Omnivorous Fish and Watersheds on Reservoir Ecosystems

Abstract Ecologists increasingly recognize the need to understand how landscapes and food webs interact. Reservoir ecosystems are heavily subsidized by nutrients and detritus from surrounding watersheds, and often contain abundant populations of gizzard shad, an omnivorous fish that consumes plankton and detritus. Gizzard shad link terrestrial landscapes and pelagic reservoir food webs by consuming detritus, translocating nutrients from sediment detritus to the water column, and consuming zooplankton. The abundance of gizzard shad increases with watershed agriculturalization, most likely through a variety of mechanisms operating on larval and adult life stages. Gizzard shad have myriad effects on reservoirs, including impacts on nutrients, phytoplankton, zooplankton, and fish, and many of their effects vary with ecosystem productivity (i.e., watershed land use). Interactive feedbacks among watersheds, gizzard shad populations, and reservoir food webs operate to maintain dominance of gizzard shad in highly productive systems. Thus, effective stewardship of reservoir ecosystems must incorporate both watershed and food-web perspectives.

Adverse Effects of Alewives on Laurentian Great Lakes Fish Communities

Abstract The alewife Alosa pseudoharengus, an invader to the Laurentian Great Lakes from the Atlantic Ocean, has been blamed for causing major disruptions of Great Lakes fish communities during the past 50 years. We reviewed the literature and examined long-term data on fish abundances in the Great Lakes to develop a new synthesis on the negative effects of alewives on Great Lakes fish communities. The results indicated that certain fish populations are substantially more vulnerable to the effects of alewives than others. More specifically, the effects of alewives on other fish populations appeared to follow a continuum—from such fishes as slimy sculpin Cottus cognatus, lake whitefish Coregonus clupeaformis, and bloater Coregonus hoyi, which were relatively unsusceptible—to Atlantic salmon Salmo salar, lake trout Salvelinus namaycush, and emerald shiner Notropis atherinoides, which were highly susceptible. Intermediate species in this continuum included yellow perch Perca flavescens, deepwater sculpin Myo...

Long-term changes of the Lake Michigan fish community following the reduction of exotic alewife (Alosa pseudoharengus)

We used our long-term annual bottom trawl survey (1973–2004) in Lake Michigan to reveal the response of the native fish community to the biological control of a dominant exotic fish, alewife (Alosa pseudoharengus), as well as to changes in total phosphorus and salmonine biomass. Through nonmetric multidimensional scaling, we documented a 1970s community largely dominated by alewife, and then a shift to a community dominated by several native species during the 1980s through 1990s, when alewife remained at relatively low levels. We argue that the recovery of burbot (Lota lota), deepwater sculpin (Myoxocephalus thompsonii), and yellow perch (Perca flavescens) was partially or fully aided by the alewife reduction. We argue that changes in phosphorus or salmonines were not directly related to abundance increases of native species. An additional community shift occurred during 1999–2004, which coincided with a reduction in species richness and total fish biomass in our trawl. The mechanisms underlying this lat...

Expansion of Dreissena into offshore waters of Lake Michigan and potential impacts on fish populations

ABSTRACT Lake Michigan was invaded by zebra mussels (Dreissena polymorpha) in the late 1980s and then followed by quagga mussels (D. bugensis) around 1997. Through 2000, both species (herein Dreissena) were largely restricted to depths less than 50 m. Herein, we provide results of an annual lake-wide bottom trawl survey in Lake Michigan that reveal the relative biomass and depth distribution of Dreissena between 1999 and 2007 (although biomass estimates from a bottom trawl are biased low). Lake-wide mean biomass density (g/m2) and mean depth of collection revealed no trend between 1999 and 2003 (mean = 0.7 g/m2 and 37 m, respectively). Between 2004 and 2007, however, mean lake-wide biomass density increased from 0.8 g/m2 to 7.0 g/m2, because of increased density at depths between 30 and 110 m, and mean depth of collection increased from 42 to 77 m. This pattern was confirmed by a generalized additive model. Coincident with the Dreissena expansion that occurred beginning in 2004, fish biomass density (generally planktivores) declined 71% between 2003 and 2007. Current understanding of fish population dynamics, however, indicates that Dreissena expansion is not the primary explanation for the decline of fish, and we provide a species-specific account for more likely underlying factors. Nonetheless, future sampling and research may reveal a better understanding of the potential negative interactions between Dreissena and fish in Lake Michigan and elsewhere.

Nearshore energy subsidies support Lake Michigan fishes and invertebrates following major changes in food web structure

Aquatic food webs that incorporate multiple energy channels (e.g., nearshore benthic and pelagic) with varying productivity and turnover rates convey stability to biological communities by providing independent energy sources. Within the Lake Michigan food web, invasive dreissenid mussels have caused rapid changes to food web structure and potentially altered the channels through which consumers acquire energy. We used stable C and N isotopes to determine how Lake Michigan food web structure has changed in the past decade, coincident with the expansion of dreissenid mussels, decreased pelagic phytoplankton production, and increased nearshore benthic algal production. Fish and invertebrate samples collected from sites around Lake Michigan were analyzed to determine taxa-specific 13C:12C (delta13C) and 15N:14N (delta15N) ratios. Sampling took place during two distinct periods, 2002-2003 and 2010-2012, that spanned the period of dreissenid expansion, and included nearshore, pelagic and profundal fish and invertebrate taxa. The magnitude and direction of the delta13C shift indicated significantly greater reliance upon nearshore benthic energy sources among nearly all fish taxa as well as profundal invertebrates following dreissenid expansion. Although the mechanisms underlying this delta13C shift likely varied among species, possible causes include the transport of benthic algal production to offshore waters and increased feeding on nearshore prey items by pelagic and profundal species. delta15N shifts were more variable and of smaller magnitude across taxa, although declines in delta15N among some pelagic fishes suggest a shift to alternative prey resources. Lake Michigan fishes and invertebrates appear to have responded to dreissenid-induced changes in nutrient and energy pathways by switching from pelagic to alternative nearshore energy subsidies. Although large shifts in energy allocation (i.e., pelagic to nearshore benthic) resulting from invasive species appear to affect total production at upper trophic levels, changes in trophic structure and utilization of novel energy pathways may help to stabilize food webs following species invasions.

Recent increases in the large glacial-relict calanoid Limnocalanus macrurus in Lake Michigan

ABSTRACT Since 2004, population density of the large hypolimnetic calanoid Limnocalanus macrurus Sars. has increased dramatically in Lake Michigan. The average summer biomass of this species between 2004 and 2006 was roughly three times that of the period 1984–2003, and at levels unprecedented in our 22-year dataset, making L. macrurus the dominant zooplankter in the lake in terms of biomass. These increases have been accentuated by coincident population declines of the main daphnid, Daphnia mendotae, in the lake with the result that in 2006, L. macrurus accounted for 75% and 50% of the large (>0.9 mm) crustacean biomass in the northern and southern basins of Lake Michigan, respectively. The increases in L. macrurus populations have closely coincided with equally dramatic increases in summer water clarity. Recent extinction coefficients are among the lowest recorded for the lake, and deepening light penetration has permitted increases in the size of the deep chlorophyll layer. In addition, planktivorous fish populations have declined coincidently with the increases in L. macrurus. It seems likely that an increase in sub-epilimnetic production has resulted in increased food resources for the deep-living L. macrurus, while low planktivore abundances have reduced predation loss, permitting L. macrurus to respond to these increases in sub-epilimnetic production.

Diet of Lake Trout and Burbot in Northern Lake Michigan During Spring: Evidence of Ecological Interaction

ABSTRACT We used analyses of burbot (Lota lota) and lake trout (Salvelinus namaycush) diets taken during spring gillnet surveys in northern Lake Michigan in 2006–2008 to investigate the potential for competition and predator-prey interactions between these two species. We also compared our results to historical data from 1932. During 2006–2008, lake trout diet consisted mainly of alewives (Alosa pseudoharengus) and rainbow smelt (Osmerus mordax), whereas burbot utilized a much wider prey base including round goby (Neogobius melanostomus), rainbow smelt, alewives, and sculpins. Using the Schoeners diet overlap index, we found a higher potential for interspecific competition in 1932 than in 2006–2008, though diet overlap was not significant in either time period. No evidence of cannibalism by lake trout or lake trout predation on burbot was found in either time period. In 2006–2008, however, lake trout composed 5.4% (by weight) of burbot diet. To determine whether this predation could be having an impact on lake trout rehabilitation efforts in northern Lake Michigan, we developed a bioenergetic-based consumption estimate for burbot on Boulder Reef (a representative reef within the Northern Refuge) and found that burbot alone can consume a considerable proportion of the yearling lake trout stocked annually, depending on burbot density. Overall, we conclude that predation, rather than competition, is the more important ecological interaction between burbot and lake trout, and burbot predation may be contributing to the failed lake trout rehabilitation efforts in Lake Michigan.

Physiological Basis of Climate Change Impacts on North American Inland Fishes

Global climate change is altering freshwater ecosystems and affecting fish populations and communities. Underpinning changes in fish distribution and assemblage-level responses to climate change are individual-level physiological constraints. In this review, we synthesize the mechanistic effects of climate change on neuroendocrine, cardiorespiratory, immune, osmoregulatory, and reproductive systems of freshwater and diadromous fishes. Observed climate change effects on physiological systems are varied and numerous, including exceedance of critical thermal tolerances, decreased cardiorespiratory performance, compromised immune function, and altered patterns of individual reproductive investment. However, effects vary widely among and within species because of species, population, and even sex-specific differences in sensitivity and resilience and because of habitat-specific variation in the magnitude of climate-related environmental change. Research on the interactive effects of climate change with other e...

Chinook Salmon Foraging Patterns in a Changing Lake Michigan

Abstract Since Pacific salmon stocking began in Lake Michigan, managers have attempted to maintain salmon abundance at high levels within what can be sustained by available prey fishes, primarily Alewife Alosa pseudoharengus. Chinook Salmon Oncorhynchus tshawytscha are the primary apex predators in pelagic Lake Michigan and patterns in their prey selection (by species and size) may strongly influence pelagic prey fish communities in any given year. In 1994–1996, there were larger Alewives, relatively more abundant alternative prey species, fewer Chinook Salmon, and fewer invasive species in Lake Michigan than in 2009–2010. The years 2009–2010 were instead characterized by smaller, leaner Alewives, fewer alternative prey species, higher abundance of Chinook Salmon, a firmly established nonnative benthic community, and reduced abundance of Diporeia, an important food of Lake Michigan prey fish. We characterized Chinook Salmon diets, prey species selectivity, and prey size selectivity between 1994–1996 and 2...

Context‐dependent planktivory: interacting effects of turbidity and predation risk on adaptive foraging

By shaping species interactions, adaptive phenotypic plasticity can profoundly influence ecosystems. Predicting such outcomes has proven difficult, however, owing in part to the dependence of plasticity on the environmental context. Of particular relevance are environmental factors that affect sensory performance in organisms in ways that alter the tradeoffs associated with adaptive phenotypic responses. We explored the influence of turbidity, which simultaneously and differentially affects the sensory performance of consumers at multiple trophic levels, on the indirect effect of a top predator (piscivorous fish) on a basal prey resource (zooplankton) that is mediated through changes in the plastic foraging behavior of an intermediate consumer (zooplanktivorous fish). We first generated theoretical predictions of the adaptive foraging response of a zooplanktivore across wide gradients of turbidity and predation risk by a piscivore. Our model predicted that predation risk can change the negative relationship between intermediate consumer foraging and turbidity into a humped-shaped (unimodal) one in which foraging is low in both clear and highly turbid conditions due to foraging-related risk and visual constraints, respectively. Consequently, the positive trait-mediated indirect effect (TMIE) of the top predator on the basal resource is predicted to peak at low turbidity and decline thereafter until it reaches an asymptote of zero at intermediate turbidity levels (when foraging equals that which is predicted when the top predator is absent). We used field observations and a laboratory experiment to test our model predictions. In support, we found humped-shaped relationships between planktivory and turbidity for several zooplanktivorous fishes from diverse freshwater ecosystems with predation risk. Further, our experiment demonstrated that predation risk reduced zooplanktivory by yellow perch (Perca flavescens) at a low turbidity, but had no effect on consumption at an intermediate turbidity. Together, our theoretical and empirical findings show how the environmental context can govern the strength of TMIEs by influencing consumer sensory performance and how these effects can become realized in nature over wide environmental gradients. Additionally, our hump-shaped foraging curve represents an important departure from the conventional view of turbiditys effect on planktivorous fishes, thus potentially requiring a reconceptualization of turbiditys impact on aquatic food-web interactions.