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Lake food webs under changing winter conditions
Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Limnology.ORCID iD: 0000-0002-5836-0602
2025 (English)Doctoral thesis, comprehensive summary (Other academic)
Description
Abstract [en]

At northern latitudes, lakes are ice-covered for several months every year, and winter has long been viewed as an ecologically dormant period. However, biological processes continue beneath the ice, and the conditions during ice cover can shape lake food webs into spring. This thesis examines how ice cover and winter conditions structure lake food webs, with a particular focus on the drivers of phyto- and zooplankton growth under ice. Through a combination of literature synthesis, long-term monitoring, and experimental and field studies, the work investigates how ice cover, winter precipitation and runoff, and habitat-specific processes. influence productivity and trophic interactions below ice. One main finding was that snow cover and ice quality (black versus white ice) are critical predictors of under-ice phytoplankton growth (paper I-II). Reduced light transmission through snow limited phytoplankton biomass and shifted community composition toward mixotrophic taxa. Further, the duration of ice cover influenced spring dynamics, where shorter ice-covered periods reduced the magnitude of the spring phytoplankton bloom (paper I-II). These findings highlight that both the duration and composition of the ice cover set the conditions for winter and spring productivity in lakes. Furthermore, the timing and magnitude of runoff were found to shape under-ice food webs. Experimental manipulations showed that early nutrient inputs under low-light conditions promoted mixotrophs, while later inputs supported stronger spring phytoplankton blooms (paper III). Field observations indicated that terrestrial organic matter inputs reduced the nutritional quality of basal resources during winter. In lakes with high terrestrial organic matter availability, copepods incorporated more terrestrial and bacterial resources but maintained nutritional quality (paper IV). Finally, spatial analyses revealed pronounced habitat heterogeneity in winter resources, with littoral habitats enriched in terrestrial and bacterial organic matter, and pelagic habitats dominated by algal-derived organic matter. These differences were reflected in copepods (paper IV), demonstrating that resource heterogeneity under ice is transferred to higher trophic levels. Together, the findings of this thesis highlight winter as a dynamic period in lake ecosystems. Climate-driven shifts in ice phenology, ice quality, and winter runoff may increase heterotrophy and reduce resource quality, weakening energy transfer to higher trophic levels and altering lake carbon cycling.

Place, publisher, year, edition, pages
Uppsala: Acta Universitatis Upsaliensis, 2025. , p. 57
Series
Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology, ISSN 1651-6214 ; 2613
Keywords [en]
Ice cover, bacteria, phytoplankton, zooplankton, light, nutrients, runoff
National Category
Ecology
Research subject
Biology with specialization in Limnology
Identifiers
URN: urn:nbn:se:uu:diva-571112ISBN: 978-91-513-2670-2 (print)OAI: oai:DiVA.org:uu-571112DiVA, id: diva2:2013965
Public defence
2026-01-20, Ekmansalen, Norbyvägen 14, Uppsala, 09:00 (English)
Opponent
Supervisors
Available from: 2025-12-17 Created: 2025-11-14 Last updated: 2025-12-17
List of papers
1. Lake ice quality in a warming world
Open this publication in new window or tab >>Lake ice quality in a warming world
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2024 (English)In: Nature Reviews Earth & Environment, E-ISSN 2662-138X, Vol. 5, no 10, p. 671-685Article, review/survey (Refereed) Published
Abstract [en]

Ice phenology has shifted with anthropogenic warming such that many lakes are experiencing a shorter ice season. However, changes to ice quality - the ratio of black and white ice layers - remain little explored, despite relevance to lake physics, ecological function, human recreation and transportation. In this Review, we outline how ice quality is changing and discuss knock-on ecosystem service impacts. Although direct evidence is sparse, there are suggestions that ice quality is diminishing across the Northern Hemisphere, encompassing declining ice thickness, decreasing black ice and increasing white ice. These changes are projected to continue in the future, scaling with global temperature increases, and driving considerable impacts to related ecosystem services. Rising proportions of white ice will markedly reduce bearing strength, implying more dangerous conditions for transportation (limiting operational use of many winter roads) and recreation (increasing the risk of fatal spring-time drownings). Shifts from black to white ice conditions will further reduce the amount of light reaching the water column, minimizing primary production, and altering community composition to favour motile and mixotrophic species; these changes will affect higher trophic levels, including diminished food quantity for zooplankton and fish, with potential developmental consequences. Reliable and translatable in situ sampling methods to assess and predict spatiotemporal variations in ice quality are urgently needed. Lake ice has witnessed considerable changes in its phenology, but less is known about ice quality - the ratio of black ice to white ice. This Review assesses the changes in lake ice quality and its ecosystem services, noting diminished ice quality in observations and projections.

Place, publisher, year, edition, pages
Springer Nature, 2024
National Category
Ecology Oceanography, Hydrology and Water Resources
Identifiers
urn:nbn:se:uu:diva-549201 (URN)10.1038/s43017-024-00590-6 (DOI)001315053200001 ()2-s2.0-85204205001 (Scopus ID)
Funder
Swedish Research Council, 2020-03222Swedish Research Council Formas, 2020-01091
Note

Correction in: Nature Reviews Earth & Environment, vol. 5, article id 906, DOI: 10.1038/s43017-024-00602-5

Available from: 2025-02-04 Created: 2025-02-04 Last updated: 2025-11-14Bibliographically approved
2. Effects of Changing Snow and Ice Cover Conditions on Phytoplankton Chlorophyll-a and Community Composition in a Mesotrophic Lake
Open this publication in new window or tab >>Effects of Changing Snow and Ice Cover Conditions on Phytoplankton Chlorophyll-a and Community Composition in a Mesotrophic Lake
2025 (English)In: Freshwater Biology, ISSN 0046-5070, E-ISSN 1365-2427, Vol. 70, no 3, article id e70012Article in journal (Refereed) Published
Abstract [en]

Ice and snow cover on lakes plays a fundamental role for under-ice ecology by reducing water column mixing and light availability. Previous studies have shown that such reductions can significantly influence the growth and reproduction of phytoplankton, primarily focusing on changes in ice-on and ice-off dates in a warming climate. This study goes beyond studying the effects of ice phenology on phytoplankton by addressing two fundamental questions: (1) how does a snow cover on ice influence below-ice phytoplankton chlorophyll-a and community composition and (2) how do variations in ice phenology influence spring phytoplankton chlorophyll-a and community composition after ice-off? To address these two questions, we assessed long-term monitoring data collected at least monthly on phytoplankton chlorophyll-a and community composition. We combined the phytoplankton data with annual ice phenology and nearby meteorological data on daily snow depth between 1997 and 2019 in a mesotrophic lake (Erken) in Sweden. Snow cover resulted in an exponential decrease of phytoplankton chlorophyll-a, with detectable effects during all 3 months studied (January-March). Deeper snow cover changed the community dominance from autotrophs to mixotrophs in two of the months studied (January and March), which we explain by reduced light availability caused by snow cover. In spring, phytoplankton chlorophyll-a increased with longer ice periods and delayed ice-off dates. A wide range of taxa in the spring community increased with delayed ice-off dates, while delayed ice-on dates mainly promoted diatoms. Convective mixing is important to keep non-motile taxa in the photic zone and could explain the increased phytoplankton growth with longer ice duration. Our results highlight seasonal ice and snow cover as key regulators for the timing of growth and reproduction of primary producers below ice, with effects of the ice cover period lasting after ice-off. Snow on ice causes light constraints, which commonly result in reduced under-ice primary production and a higher proportion of mixotrophs in the phytoplankton community. Losing high nutritional phytoplankton groups such as mixotrophs following changes in ice phenology and snow cover can have consequences for the trophic transfer and the biogeochemical cycling in lakes.

Place, publisher, year, edition, pages
John Wiley & Sons, 2025
Keywords
ice phenology, ice quality, mixotrophy, primary producers, winter limnology
National Category
Ecology
Identifiers
urn:nbn:se:uu:diva-553126 (URN)10.1111/fwb.70012 (DOI)001438944400001 ()
Funder
Swedish Research Council, 2020-01091Swedish Research Council Formas
Available from: 2025-03-26 Created: 2025-03-26 Last updated: 2025-12-05Bibliographically approved
3. Timing of nutrient input pulses during ice cover regulates lake spring plankton dynamics
Open this publication in new window or tab >>Timing of nutrient input pulses during ice cover regulates lake spring plankton dynamics
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(English)Manuscript (preprint) (Other academic)
National Category
Ecology
Identifiers
urn:nbn:se:uu:diva-571108 (URN)
Available from: 2025-11-06 Created: 2025-11-06 Last updated: 2025-11-14
4. Spatial heterogeneity in copepod resource use under ice
Open this publication in new window or tab >>Spatial heterogeneity in copepod resource use under ice
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(English)Manuscript (preprint) (Other academic)
National Category
Ecology
Identifiers
urn:nbn:se:uu:diva-571111 (URN)
Available from: 2025-11-07 Created: 2025-11-07 Last updated: 2025-11-14

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