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Abstract The phenology of critical biological events in aquatic ecosystems is rapidly shifting due to climate change. Growing variability in phenological cues can increase the likelihood of trophic mismatches (i.e., mismatches in the timing of peak prey and predator abundances), causing recruitment failures in important fisheries. We assessed changes in the spawning phenology of walleye (Sander vitreus) in 194 Midwest US lakes to investigate factors influencing walleye phenological responses to climate change and associated climate variability, including ice‐off timing, lake physical characteristics, and population stocking history. Ice‐off phenology shifted earlier, about three times faster than walleye spawning phenology over time. Spawning phenology deviations from historic averages increased in magnitude over time, and large deviations were associated with poor offspring survival. Our results foreshadow the risks of increasingly frequent natural recruitment failures due to mismatches between historically tightly coupled spawning and ice‐off phenology. 

The phenology of critical biological events in aquatic ecosystems are rapidly shifting due to climate change. Growing variability in phenological cues can increase the likelihood of trophic mismatches, causing recruitment failures in commercially, culturally, and recreationally important fisheries. We tested for changes in spawning phenology of regionally important walleye (Sander vitreus) populations in 194 Midwest US lakes in Minnesota, Michigan, and Wisconsin spanning 1939-2019 to investigate factors influencing walleye phenological responses to climate change and associated climate variability, including ice-off timing, lake physical characteristics, and population stocking history. Data from Wisconsin and Michigan lakes (185 and 5 out of 194 total lakes, respectively) were collected by the Wisconsin Department of Natural Resources (WDNR) and the Great Lakes Indian Fish and Wildlife Commission (GLIFWC) through standardized spring walleye mark-recapture surveys and spring tribal harvest season records. Standardized spring mark-recapture population estimates are performed shortly after ice-off, where following a marking event, a subsequent recapture sampling event is conducted using nighttime electrofishing (typically AC – WDNR, pulsed-DC – GLIFWC) of the entire shoreline including islands for small lakes and index stations for large lakes (Hansen et al. 2015) that is timed to coincide with peak walleye spawning activity (G. Hatzenbeler, WDNR, personal communication; M. Luehring, GLIFWC, personal communication; Beard et al. 1997). Data for four additional Minnesota lakes were collected by the Minnesota Department of Natural Resources (MNDNR) beginning in 1939 during annual collections of walleye eggs and broodstock (Schneider et al. 2010), where date of peak egg take was used to index peak spawning activity. For lakes where spawning location did not match the lake for which the ice-off data was collected, the spawning location either flowed into (Pike River) or was within 50 km of a lake where ice-off data were available (Pine River) and these ice-off data were used. Following the affirmation of off-reservation Ojibwe tribal fishing rights in the Ceded Territories of Wisconsin and the Upper Peninsula of Michigan in 1987, tribal spearfishers have targeted walleye during spring spawning (Mrnak et al. 2018). Nightly harvests are recorded as part of a compulsory creel survey (US Department of the Interior 1991). Using these records, we calculated the date of peak spawning activity in a given lake-year as the day of maximum tribal harvest. Although we were unable to account for varying effort in these data, a preliminary analysis comparing spawning dates estimated using tribal harvest to those determined from standardized agency surveys in the same lake and year showed that they were highly correlated (Pearson’s correlation: r = 0.91, P < 0.001). For lakes that had walleye spawning data from both agency surveys and tribal harvest, we used the data source with the greatest number of observation years. Ice-off phenology data was collected from two sources – either observed from the Global Lake and River Ice Phenology database (Benson et al. 2000)t, or modeled from a USGS region-wide machine-learning model which used North American Land Data Assimilation System (NLDAS) meteorological inputs combined with lake characteristics (lake position, clarity, size, depth, hypsography, etc.) to predict daily water column temperatures from 1979 - 2022, from which ice-off dates could be derived (https://www.sciencebase.gov/catalog/item/6206d3c2d34ec05caca53071; see Corson-Dosch et al. 2023 for details). Modeled data for our study lakes (see (Read et al. 2021) for modeling details), which performed well in reflecting ice phenology when compared to observed data (i.e., highly significant correlation between observed and modeled ice-off dates when both were available; r = 0.71, p < 0.001). Lake surface area (ha), latitude, and maximum depth (m) were acquired from agency databases and lake reports. Lake class was based on a WDNR lakes classification system (Rypel et al. 2019) that categorized lakes based on temperature, water clarity, depth, and fish community. Walleye stocking history was defined using the walleye stocking classification system developed by the Wisconsin Technical Working Group (see also Sass et al. 2021), which categorized lakes based on relative contributions of naturally-produced and stocked fish to adult recruitment by relying heavily on historic records of age-0 and age-1 catch rates and stocking histories. Wisconsin lakes were divided into three groups: natural recruitment (NR), a combination of stocking and natural recruitment (C-ST), and stocked only (ST). Walleye natural recruitment was indexed as age-0 walleye CPE (number of age-0 walleye captured per km of shoreline electrofished) from WDNR and GLIFWC fall electrofishing surveys (see Hansen et al. 2015 for details). We excluded lake-years where stocking of age-0 fish occurred before age-0 surveys to only include measurements of naturally-reproduced fish. 

Climate change is leading to shifts in not only the average timing of phenological events, but also their variance and predictability. Increasing phenological variability creates a stochastic environment that is critically understudied, particularly in aquatic ecosystems. We provide a perspective on the possible implications for increasingly unpredictable aquatic habitats, including more frequent trophic asynchronies and altered hydrologic regimes, focusing on ice-off phenology in lakes. Increasingly frequent phenological extremes may limit the ability of organisms to optimize traits required to adapt to a warming environment. Using a unique, long-term ecological dataset on Escanaba Lake, WI, USA, as a case study, we show that the average date of ice-off is shifting earlier and becoming more variable, thus altering limnological conditions and yielding uncoupled food web responses with ramifications for fish spawn timing and recruitment success. A genes-to-ecosystems understanding of the responses of aquatic communities to increasingly variable phenology is needed. Our perspective suggests that management for diversity, at the intra- and interspecific levels, will become paramount for conserving resilient aquatic ecosystems. 

Freshwater recreational fisheries regulations are a vital tool for achieving social and ecological fisheries objectives. However, angler behavior and fish biology may interact to influence regulation efficacy in unexpected ways. We combined models of fish growth and angler behavior to explore how angler behavior interacts with fish life history to shape the probability of fish harvest given capture across ages, life-stages, and sexes of walleye (Sander vitreus). Compared to females, males grew more quickly as juveniles, matured earlier, and reached smaller maximum sizes. Male walleye were therefore vulnerable to harvest for more of their reproductive lives than females because males spent more time at sizes where anglers were very likely to harvest them. We suggest that restricting harvest of large individuals in sexually-dimorphic species may favor the survival of large, reproductive-aged females. Moreover, we show that combining models of fish growth and harvester behavior can provide insights into how harvest affects fish with complex life histories over the course of their lives.