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.
Nearshore water clarity, as measured by remotely sensed Kd(490), and stable C and N isotopes of several nearshore fishes differed across the Lake Michigan basin. Values of δ13C of round goby (Neogobius melanstomus), yellow perch (Perca flavescens), and spottail shiner (Notropis hudsonis) were depleted in the southeast where water clarity was low relative to the southwest where water clarity was greater. Bayesian analyses were used to evaluate spatial variation in diet composition and quantify the relationship between water clarity and the proportional importance of pelagic energy in fish diets. Water clarity in nearshore areas is likely related to variable riverine inputs, resuspension, and upwelling processes. While these processes may not directly impact δ13C or δ15N of nearshore fishes, we hypothesize that water clarity differentially affects benthic and pelagic algal production. Lower water clarity in the benthos and subsequently lower benthic productivity may be related to regional diet differences and increased reliance on pelagic energy sources. Mobile fishes such as alewife (Alosa pseudoharengus) may not be in isotopic equilibrium with regional prey sources and depart from spatial patterns observed in other nearshore fishes.
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