Research Themes in NTL-8

Our vision is to gain a predictive understanding of the ecology of lakes at longer and broader scales than has been traditional in limnology. Our conceptual framework uses a nested set of spatial scales, from individual lakes and their watersheds, to hydrologically-linked sets of lakes, entire lake districts, multiple lake districts within the Great Lakes region, and comparative studies of lakes and lake districts around the globe. Our research program is interdisciplinary and aims to understand the ecology of lakes in relation to relevant atmospheric, geochemical, landscape and human processes. Within the NTL domain, we have observed shifts in driver and response variables that are both gradual and abrupt on time scales of years to decades. Examples of rapid changes in lake responses that have been particularly conspicuous to both researchers and many Wisconsin residents include the collapse of valuable Walleye fisheries; increasing fluctuations in lake levels; arrival of new aquatic invaders; and sudden declines in lake water clarity, among others. At the same time, we are also seeing changes in drivers that can cause abrupt change, but abrupt changes have not followed, leading us to ask why. Our research activities are inspired by these observations and are organized around the overarching question: What are the causes and consequences of abrupt ecological change in lakes and their surrounding landscapes?
What are lake phenological responses to a warmer and more variable climate that may lead to abrupt ecological change?
We’re investigating how increasing climate variability shifts winter and spring phenology, triggering abrupt ecological responses (Dugan 2021; Feiner et al. 2022). Analyses of long-term data showed that increasing variability in ice-off timing disrupts walleye recruitment due to photoperiod spawning cues, limiting adaptation to changing spring conditions and contributing to population declines (Barta et al. 2024; Feiner et al. 2025). Complementary work revealed how species invasions can trigger abrupt responses in organic matter cycling (Rohwer et al. 2023, 2025), as the timing of stratification, primary production, and invasive species interact to drive the spring clearwater phase and summer anoxia (Ladwig et al. 2021; Rohwer et al. 2024). Bacteria, viruses, and protists in Lake Mendota undergo predictable phenological cycles along with abrupt shifts in species composition (Rohwer et al. 2023, 2025; Zhou et al. 2025). These findings demonstrate that variability in winter and spring weather propagates through food webs and biogeochemical processes.
Representative Publications from this Research Theme
- Barta, M.E., G.G. Sass, J.R. Reed, T.A. Cichosz, A.D. Shultz, M. Luehring, and Z.S. Feiner (2024) Lagging spawning and increasing phenological extremes jeopardize walleye (Sander vitreus) in north‐temperate lakes. Limnology and Oceanography Letters 9: 229–236.
- Dugan, H A (2021) A comparison of ecological memory of lake ice‐off in eight north‐temperate lakes. Journal of Geophysical Research: Biogeosciences 126: e2020JG006232.
- Feiner, ZS, Dugan, HA, Lottig, NR, Sass, GG, & Gerrish, GA (2022) A perspective on the ecological and evolutionary consequences of phenological variability in lake ice on north-temperate lakes. Canadian Journal of Fisheries and Aquatic Sciences, 79 (9), 1590-1604.
- Feiner, Z.S., S.L. Shaw, and G.G. Sass (2025) Understanding shifting cues for walleye spawning phenology and recruitment in a changing climate. Canadian Journal of Fisheries and Aquatic Sciences 82: 1-10.
- Ladwig, R, Hanson, PC, Dugan, HA, Carey, CC, Zhang, Y, Shu, L, …& Cobourn, KM (2021) Lake thermal structure drives interannual variability in summer anoxia dynamics in a eutrophic lake over 37 years. Hydrology and Earth System Sciences, 25 (2), 1009-1032.
- Ladwig, R, Appling, AP, Delany, A, Dugan, HA, Gao, Q, Lottig, N, …& Hanson, PC (2022) Long‐term change in metabolism phenology in north temperate lakes. Limnology and Oceanography, 67 (7), 1502-1521.
- Rohwer, RR, Hale, RJ, Vander Zanden, MJ, Miller, TR, & McMahon, KD. (2023) Species invasions shift microbial phenology in a two-decade freshwater time series. Proceedings of the National Academy of Sciences, 120 (11), e2211796120.
- Rohwer, R.R., R. Ladwig, P.C. Hanson, J.R. Walsh, M.J. Vander Zanden, and H.A. Dugan (2024) Increased anoxia following species invasion of a eutrophic lake. Limnology and Oceanography Letters 9: 33-42.
- Rohwer, R.R., M. Kirkpatrick, S.L. Garcia, M. Kellom, K.D. McMahon, and B.J. Baker (2025) Two decades of bacterial ecology and evolution in a freshwater lake. Nature Microbiology 10: 246–257.
- Zhou, Z., P. Q. Tran, …. K.D. McMahon (2025) Unravelling viral ecology and evolution over 20 years in a freshwater lake. Nature Microbiology 10: 231–45.
How do interactions of land use/land cover and climate affect urban aquatic ecosystems?
We’re examining how climate variability interacts with spatial gradients in urban areas to drive ecosystem change. In the Yahara watershed, abrupt canopy loss from emerald ash borer infestation did not uniformly intensify the urban heat island (Berg and Kucharik 2022, 2024); however, tree canopy cover does regulate runoff through plant water use and interception, with implications for urban flooding (Voter and Loheide 2021). High-frequency measurements in stormwater ponds showed that precipitation can trigger abrupt increases in water level and subsequently primary production (Briggs 2025; Curtis 2025) whereas summer productivity legacies control winter oxygen and greenhouse gases flux (Gorsky et al. 2024). Surveys across land-use gradients revealed lower species richness but high beta diversity in urban ponds and strong effects of habitat complexity and connectivity (Sauer et al. 2022; Trovillion et al. 2023). Together, these studies show how climate and land-use interact to generate diverse, and sometimes abrupt, ecological responses in urban waterbodies.
Representative Publications from this Research Theme
- Berg E, C Kucharik (2022) The dynamic relationship between air and land surface temperature within the Madison, Wisconsin urban heat island. Remote Sensing 14:165.
- Berg, E., and C. Kucharik (2024) Impacts of ash tree removals on summer daytime temperatures in Madison, Wisconsin. Environmental Research Communications 6:085001.
- Gorsky, A.L., H.A. Dugan, G.M. Wilkinson, and E.H. Stanley (2024) Under-ice oxygen depletion and greenhouse gas supersaturation in north temperate urban ponds. Journal of Geophysical Research: Biogeosciences 129: e2024JG008120.
- Sauer, EL, J Cruz, E Crone, C Lewis, E Plumier, B Cwynar, D Drake, BM Herrick, DL Preston (2022) Multiscale drivers of amphibian community occupancy in urban ponds. Urban Ecosystems 25:1469–1479
- Trovillion, DC, EL Sauer, G Shay, ER Crone, DL Preston. (2023) Habitat complexity, connectivity, and introduced fish drive pond community structure along an urban to rural gradient. Ecological Applications 33(4): e2828
- Voter, C.B. and S.P. Loheide II (2021) Climatic controls on the hydrologic effects of urban low impact development practices. Environmental Research Letters 16: 064021
How do external drivers interact with aquatic invasive species to regulate water quality?
We’re examining how external drivers including extreme precipitation and spatial variation in watershed inputs interact to regulate water quality in Lake Mendota. Analysis of long-term high frequency sensor data showed that cyanobacterial blooms lag storms by 1–60 days due to internal processes of grazing and nutrient recycling (Carpenter et al. 2022). Modeling further demonstrated the effects of nutrient loading can persist for decades even if external P loading was abruptly eliminated (Hanson et al. 2023). Spatial surveys of water quality after storms revealed that moderate precipitation produces only weak immediate changes in nutrients and phytoplankton (Kibler 2026; Ortiz 2026), but storms may contribute to sediment nutrient pools that are later mobilized by wind-driven mixing, generating lagged effects, while wind-driven waves influence bloom initiation and senescence (Gushulak et al. 2025). These studies of Mendota demonstrate that abrupt shifts in water quality of large lakes arise from lagged interactions among watershed forcing and climate variability.
Representative Publications from this Research Theme
- Carpenter, SR, MR Gahler, CJ Kucharik, EH Stanley (2022) Long-range dependence and extreme values of precipitation, phosphorus load, and cyanobacteria. Proceedings of the National Academy of Sciences 119 (48) e2214343119o
- Carpenter, S. R., and W. A. Brock (2024) Stochastic dynamics of phycocyanin in years of contrasting phosphorus load. Ecosphere 15: e4903.
- Gushulak, C.A.C., T.H. Bertram, H.A. Dugan, J.A. Franck, M.N. Rogers, S.T. Salemink‐Harry, B.J. Smith, T.J.W. Wagner, L.K. Zoet, N. Pujara, and G.M. Wilkinson (2025) The role of surface water waves on cyanobacterial blooms in lakes. Limnology and Oceanography Letters 10: 602-618.
- Hanson, P.C. R. Ladwig, C. Buelo, E.A. Albright, A.D. Delany, and C.C. Carey (2023) Legacy phosphorus and ecosystem memory control future water quality in a eutrophic lake. Journal of Geophysical Research: Biogeosciences 128: e2023JG007620.
- Kibler, K. 2026. Freshwater cyanobacterial population diversity through time and space in a eutrophic lake. Ph.D., University of Wisconsin-Madison.
- Martin, BE, JR Walsh, MJ Vander Zanden (2022) Rise of a native apex predator and an invasive zooplankton cause successive ecological regime shifts in a North Temperate Lake. Limnology and Oceanography 67 (S1)
- Ortiz, D. A. 2026. Spatial-temporal heterogeneity of lakes across varying scales and trophic status. PhD, University of Wisconsin-Madison, Madison.
- Rohwer, RR, RJ Hale, MJ Vander Zanden, TR Miller, KD McMahon (2023) Species invasions shift microbial phenology in a two-decade freshwater time series. Proceedings of the National Academy of Sciences 120: e2211796120.
- Spear, MJ, JR Walsh, A Ricciardi, MJ Vander Zanden (2021) The invasion ecology of sleeper populations: prevalence, persistence, and abrupt shifts. BioScience 71: 357-369
What causes intentional ecosystem manipulations to persist, revert, or lead to novel states?
We used invasive species management as ecosystem-scale experiments to examine how induced abrupt changes persist, revert, or lead to novel states (Perales et al. 2021; Carpenter 2026). Removal of common carp in Lake Wingra triggered a rapid shift from turbid water to dense macrophytes and filamentous algae that has persisted for over 15 years (Gorsky et al. 2026). Parallel fish manipulations in Crystal and Sparkling Lakes tested whether restructuring food webs could suppress invasive rainbow smelt and promote native fish recovery. After concurrent smelt removal and cisco reintroduction, smelt remain rare while native fishes have increased (Mrnak et al. 2023, 2025). These experiments show how management can trigger abrupt shifts while hinting at the feedbacks that influence persistence. A synthesis of decades of NTL research further articulated general principles of aquatic invasions (Vander Zanden et al. 2024).
Representative Publications from this Research Theme
- Carpenter, S. R (2026) Regime shifts in lakes: Observations, models, and experiments. Current Opinion in Systems Biology 44: 100576.
- Gorsky, A.L., H.A. Dugan, G.M. Wilkinson, E.H. Stanley, M.J. Vander Zanden, and R.C. Lathrop (2026) Long-term stability of macrophyte dominance triggered by common carp removal from a temperate shallow lake. Ecosystems 29:21.
- Mrnak, JT, LW Sikora, MJ Vander Zanden, GG Sass (2023) Applying Panarchy Theory to Aquatic Invasive Species Management: A Case Study on Invasive Rainbow Smelt Osmerus mordax. Reviews in Fisheries Science & Aquaculture 31 (1): 66-85.
- Mrnak, J. T., M. V. Wilkinson, L. W. Sikora, L. M. Feucht, A. M. Mrnak, M. J. Vander Zanden, and G. G. Sass (2025) Invasive control and native restoration: Directing ecosystem transformation through purposeful food web manipulations. Fisheries 50: 194–208.
- Perales, K.M., G.J.A. Hansen, C.L. Hein, J.T. Mrnak, B.M. Roth, J.R. Walsh, and M.J. Vander Zanden, M.J. (2021) Spatial and temporal patterns in native and invasive crayfishes during a 19‐year whole‐lake invasive crayfish removal experiment. Freshwater Biology 66: 2105–2117.
- Vander Zanden, M. J., A Gorsky, GJA Hansen, PTJ Johnson, AW Latzka, A Mikulyuk, RR Rowher, MJ Spear, JR Walsh (2024) Nine Lessons about Aquatic Invasive Species from the North Temperate Lakes Long-Term Ecological Research (NTL-LTER) Program. BioScience 74: 509–523.