Global climate warming is causing the loss of freshwater ice around the Northern Hemisphere. Although the timing and duration of ice covers are known to regulate ecological processes in seasonally ice-covered ecosystems, the consequences of shortening winters for freshwater biota are poorly understood owing to the scarcity of under-ice research. Here, we present one of the first in-lake experiments to postpone ice-cover onset (by ≤21 d), thereby extending light availability (by ≤40 d) in early winter, and explicitly demonstrate cascading effects on pelagic food web processes and phenologies. Delaying ice-on elicited a sequence of events from winter to spring: 1) relatively greater densities of algal resources and primary consumers in early winter; 2) an enhanced prevalence of winter-active (overwintering) consumers throughout the ice-covered period, associated with augmented storage of high-quality fats likely due to a longer access to algal resources in early winter; and 3) an altered trophic structure after ice-off, with greater initial springtime densities of overwintering consumers driving stronger, earlier top-down regulation, effectively reducing the spring algal bloom. Increasingly later ice onset may thus promote consumer overwintering, which can confer a competitive advantage on taxa capable of surviving winters upon ice-off; a process that may diminish spring food availability for other consumers, potentially disrupting trophic linkages and energy flow pathways over the subsequent open-water season. In considering a future with warmer winters, these results provide empirical evidence that may help anticipate phenological responses to freshwater ice loss and, more broadly, constitute a case of climate-induced cross-seasonal cascade on realized food web processes.Global climate warming increases surface water temperature (
1,
2), causing widespread loss of freshwater ice cover across the Northern Hemisphere (
3,
4). Long-term records indicate that freshwater ecosystems at higher latitudes experience accelerating rates of warming, with strong trends of increasingly later ice-on, earlier ice-off, and shorter duration of seasonal ice covers (
5–
8). Pronounced changes in the timing and duration of ice-covered seasons are bound to have far-reaching ecological consequences (
9). Yet, freshwater ecology remains relatively understudied in winter, especially when compared with marine research (
10–
13). Although ice phenology and some remotely sensed abiotic measures have been well-documented in lakes and rivers (
3,
8,
10), under-ice assessments of biota are considerably limited. As a result, the effects of rapidly changing winters on freshwater food webs and biological processes remain largely unknown, impeding our ability to anticipate the implications of ice loss.The ice-covered period has long been perceived as a dormant season for freshwater organisms, especially in lakes (
12). Recent studies challenged this prevailing view, suggesting that critical processes can occur under ice (
14,
15), with the timing and physical features of ice covers driving dynamics in winter, spring, and possibly over the summer (
12,
16,
17). Indeed, while several freshwater taxa may favor dormancy in winter, certain environmental conditions (e.g., light, food availability) and organismal traits (e.g., thermal tolerance) may allow communities to remain active under ice. For example, motile phytoplankton with flexible forms of nourishment—such as mixotrophic phytoflagellates—may prevail in winter (
18,
19), remaining in suspension despite the absence of water mixing under ice and shifting their nutritional mode from photosynthesis to partially or fully utilizing organic sources of carbon when light availability is reduced. Pelagic primary consumers may also benefit from relaxed predation in winter, with reports of active, abundant, and, even, reproducing zooplankton populations (
20–
22). Active overwintering, however, may impose a long period of limited resources on consumers. For instance, energetically rewarding, photosynthetic prey can become rare in winter, compelling zooplankton to rely on other survival strategies to overcome nutritional shortage, such as using previously accumulated high-quality fat reserves (
20,
23,
24) or shifting their diet to incorporate alternative, lower-quality resources of bacterial or terrestrial origin (
19,
25).As ice-on dates trend later with climate warming, prolonged light exposure in early winter may sustain photosynthesis and energy-rich resource availability longer, permitting greater accumulation of algal-derived fat reserves in primary consumers. Extending the time period during which zooplankton can store high-quality fats may in turn enhance active overwintering. Shorter seasonal duration of ice cover could also facilitate winter survival, reducing the period during which consumers may rely on previously accumulated fat storage. A greater prevalence of winter-active consumers could, however, trigger strong cascading effects on trophic interactions upon ice breakup, including reduced spring algal blooms or greater mismatches in the timing of resource and consumer growth [phenological decoupling (
26–
30)]. Enhanced winter survival in consumers could also promote the co-occurrence of competing taxa in spring, resulting in earlier and possibly greater competition for limited resources (
31). Thus, the timing of ice-cover onset may represent a key determinant of consumer winter strategies and food web interactions at the beginning of the open-water season, a question that remains hitherto unanswered owing to the paucity of empirical data on under-ice freshwater food webs.To explicitly test how later ice-on may affect winter survival in pelagic primary consumers and springtime planktonic food webs, we performed an in situ experimental study to manipulate the timing of ice-cover onset using in-lake enclosures. Exploiting a floating facility deployed on a temperate lake in eastern Canada (45°32′N, 73°08′W), we filled 16 pelagic enclosures with
ca. 3,400 L of lake water and planktonic organisms at the beginning of the ice-covered period () and postponed the onset of naturally forming ice cover by 0, 7, 14, and 21 d (), resulting in contrasting light-incidence patterns over a period up to 40 d (
SI Appendix, Fig. S1). We recorded a variety of biotic responses over 183 d through winter and spring () to address whether delayed ice-on dates and prolonged light exposure can: 1) sustain photosynthesis longer and allow pelagic multitrophic communities (algal resources and zooplankton consumers) to maintain high densities in early winter; 2) enhance the prevalence of overwintering consumers over the winter months, and whether winter survival is linked to early-winter fat accumulation and/or shifts in dietary supply sources; and 3) elicit cascading effects on the structure of springtime planktonic food webs, both across and within trophic levels (i.e., consumer–resource and consumer–consumer relationships).
Open in a separate windowIn-lake experimental setup, under-ice organisms, design, and timeline. (
A) The floating platform deployed on Lac Hertel (Mont Saint-Hilaire, Québec), accommodating 16 enclosures (depth
ca. 4.2 m), each containing
ca. 3,400 L of lake water and planktonic organisms. (
B and
C) Microscope photographs of under-ice plankton in early winter: (
B) phototrophic nanophytoflagellates (
Cryptophyceae; epifluorescence microscopy) and (
C) crustacean zooplankton (
Bosmina). (
D and
E) Schematic representation of ice-cover treatments and timeline. Colors refer to treatment levels of ice-cover manipulation, indicating the (
D) timing of ice-cover formation onset across enclosures and (
E) resulting ice-covered periods. Lac Hertel’s ice-covered period is illustrated as a frame of reference. (
E) Symbols indicate sampling days over the course of the 183-d experiment. Biotic response variables include total and group-specific chl-
a concentrations, crustacean and rotifer zooplankton densities, fully or partially phototrophic (PNF) and heterotrophic/phagotrophic (HNF) nanoflagellates, and characterization of FAs in zooplankton and seston. Upper and lower horizontal axes indicate time of the year and day of experiment, respectively. Temporal dynamics of light incidence and other baseline measurements are provided in
SI Appendix, with additional details in
Materials and Methods. *FAs were characterized on six occasions for seston (filled symbols) and on three occasions for zooplankton (framed symbols) (see details in
Materials and Methods).
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