title: Ecology under lake ice.
abstract: Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer 'growing seasons'. We executed the first global quantitative synthesis on under-ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter-summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake-specific, species-specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass.
authors.affiliation: Center for Environmental Research, Education and Outreach, Washington State University, Pullman, WA, USA.
authors.forename: Stephanie E
authors.initials: SE
authors.lastname: Hampton
authors.affiliation: Oregon Institute of Marine Biology, University of Oregon, Charleston, OR, USA.
authors.forename: Aaron W E
authors.initials: AW
authors.lastname: Galloway
authors.affiliation: Center for Environmental Research, Education and Outreach, Washington State University, Pullman, WA, USA.
authors.forename: Stephen M
authors.initials: SM
authors.lastname: Powers
authors.affiliation: Department of Biology & Large Lakes Observatory, University of Minnesota Duluth, Duluth, MN, USA.
authors.forename: Ted
authors.initials: T
authors.lastname: Ozersky
authors.affiliation: Center for Environmental Research, Education and Outreach, Washington State University, Pullman, WA, USA.
authors.forename: Kara H
authors.initials: KH
authors.lastname: Woo
authors.affiliation: Department of Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, NJ, USA.
authors.forename: Ryan D
authors.initials: RD
authors.lastname: Batt
authors.affiliation: Center for Environmental Research, Education and Outreach, Washington State University, Pullman, WA, USA.
authors.forename: Stephanie G
authors.initials: SG
authors.lastname: Labou
authors.affiliation: Department of Geography-Geology, Illinois State University, Normal, IL, USA.
authors.forename: Catherine M
authors.initials: CM
authors.lastname: O'Reilly
authors.affiliation: Department of Biology, York University, Toronto, ON, Canada.
authors.forename: Sapna
authors.initials: S
authors.lastname: Sharma
authors.affiliation: Center for Limnology, University of Wisconsin, Boulder Junction, WI, USA.
authors.forename: Noah R
authors.initials: NR
authors.lastname: Lottig
authors.affiliation: Center for Limnology, University of Wisconsin, Madison, WI, USA.
authors.forename: Emily H
authors.initials: EH
authors.lastname: Stanley
authors.affiliation: Global Institute for Water Security, University of Saskatchewan, Saskatoon, SK, Canada.
authors.forename: Rebecca L
authors.initials: RL
authors.lastname: North
authors.affiliation: Rubenstein Ecosystem Science Laboratory, University of Vermont, Burlington, VT, USA.
authors.forename: Jason D
authors.initials: JD
authors.lastname: Stockwell
authors.affiliation: Department of Ecosystem Research, Leibniz Institute of Freshwater Ecology and Inland Fisheries, Berlin, Germany.
authors.forename: Rita
authors.initials: R
authors.lastname: Adrian
authors.affiliation: Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
authors.forename: Gesa A
authors.initials: GA
authors.lastname: Weyhenmeyer
authors.affiliation: Lammi Biological Station, University of Helsinki, Lammi, Finland.
authors.forename: Lauri
authors.initials: L
authors.lastname: Arvola
authors.affiliation: Global Institute for Water Security, University of Saskatchewan, Saskatoon, SK, Canada.|School of Environment and Sustainability, University of Saskatchewan, Saskatoon, SK, Canada.
authors.forename: Helen M
authors.initials: HM
authors.lastname: Baulch
authors.affiliation: Water Center, Graham Sustainability Institute, University of Michigan, Ann Arbor, MI, USA.
authors.forename: Isabella
authors.initials: I
authors.lastname: Bertani
authors.affiliation: Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT, USA.
authors.forename: Larry L
authors.initials: LL
authors.lastname: Bowman
authors.affiliation: Department of Biological Sciences, Virginia Tech, Blacksburg, VA, USA.
authors.forename: Cayelan C
authors.initials: CC
authors.lastname: Carey
authors.affiliation: CREAF, Consejo Superior de Investigaciones Científicas, Cerdanyola, Spain.
authors.forename: Jordi
authors.initials: J
authors.lastname: Catalan
authors.affiliation: Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
authors.forename: William
authors.initials: W
authors.lastname: Colom-Montero
authors.affiliation: Department of Biology, University of St. Thomas, St. Paul MN, USA.
authors.forename: Leah M
authors.initials: LM
authors.lastname: Domine
authors.affiliation: Department of Ecology, Universitat de Barcelona, Barcelona, Spain.
authors.forename: Marisol
authors.initials: M
authors.lastname: Felip
authors.affiliation: Centre for Research, Monitoring and Evaluation, Sierra de Guadarrama National Park, Rascafría, Spain.
authors.forename: Ignacio
authors.initials: I
authors.lastname: Granados
authors.affiliation: Center for Limnology, University of Wisconsin, Madison, WI, USA.
authors.forename: Corinna
authors.initials: C
authors.lastname: Gries
authors.affiliation: Department of Experimental Limnology, Leibniz Institute of Freshwater Ecology and Inland Fisheries, Stechlin, Germany.|Institute for Biochemistry and Biology, Potsdam University, Potsdam, Germany.
authors.forename: Hans-Peter
authors.initials: HP
authors.lastname: Grossart
authors.affiliation: Centre for Limnology, Estonian University of Life Sciences, Tartu, Estonia.
authors.forename: Juta
authors.initials: J
authors.lastname: Haberman
authors.affiliation: Centre for Limnology, Estonian University of Life Sciences, Tartu, Estonia.
authors.forename: Marina
authors.initials: M
authors.lastname: Haldna
authors.affiliation: Department of Biology, University of New Brunswick, Fredericton, NB, Canada.
authors.forename: Brian
authors.initials: B
authors.lastname: Hayden
authors.affiliation: IISD Experimental Lakes Area, Winnipeg, MB, Canada.
authors.forename: Scott N
authors.initials: SN
authors.lastname: Higgins
authors.affiliation: Columbia River Fisheries Program Office, U.S. Fish & Wildlife Service, Vancouver, WA, USA.
authors.forename: Jeff C
authors.initials: JC
authors.lastname: Jolley
authors.affiliation: Department of Environmental Sciences, University of Helsinki, Helsinki, Finland.
authors.forename: Kimmo K
authors.initials: KK
authors.lastname: Kahilainen
authors.affiliation: Institute of Geology, Department of Isotope Paleoclimatology, Tallinn University of Technology, Tallinn, Estonia.
authors.forename: Enn
authors.initials: E
authors.lastname: Kaup
authors.affiliation: Global Institute for Water Security, University of Saskatchewan, Saskatoon, SK, Canada.|School of Environment and Sustainability, University of Saskatchewan, Saskatoon, SK, Canada.
authors.forename: Michael J
authors.initials: MJ
authors.lastname: Kehoe
authors.affiliation: Department of Ecology, Evolution and Marine Biology, University of California Santa Barbara, Santa Barbara, CA, USA.
authors.forename: Sally
authors.initials: S
authors.lastname: MacIntyre
authors.affiliation: Department of Geography, University College London, London, UK.
authors.forename: Anson W
authors.initials: AW
authors.lastname: Mackay
authors.affiliation: National Wildlife Research Centre, Science and Technology Division, Environment and Climate Change Canada, Ottawa, ON, Canada.
authors.forename: Heather L
authors.initials: HL
authors.lastname: Mariash
authors.affiliation: Department of Biological Sciences, Bowling Green State University, Bowling Green, OH, USA.
authors.forename: Robert M
authors.initials: RM
authors.lastname: McKay
authors.affiliation: Department of Freshwater Conservation, Brandenburg University of Technology Cottbus - Senftenberg, Bad Saarow, Germany.
authors.forename: Brigitte
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authors.lastname: Nixdorf
authors.affiliation: Centre for Limnology, Estonian University of Life Sciences, Tartu, Estonia.
authors.forename: Peeter
authors.initials: P
authors.lastname: Nõges
authors.affiliation: Centre for Limnology, Estonian University of Life Sciences, Tartu, Estonia.
authors.forename: Tiina
authors.initials: T
authors.lastname: Nõges
authors.affiliation: Environmental Monitoring and Reporting Branch, Ontario Ministry of the Environment and Climate Change, Toronto, ON, Canada.
authors.forename: Michelle
authors.initials: M
authors.lastname: Palmer
authors.affiliation: Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
authors.forename: Don C
authors.initials: DC
authors.lastname: Pierson
authors.affiliation: Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT, USA.
authors.forename: David M
authors.initials: DM
authors.lastname: Post
authors.affiliation: Center for Environmental Research, Education and Outreach, Washington State University, Pullman, WA, USA.
authors.forename: Matthew J
authors.initials: MJ
authors.lastname: Pruett
authors.affiliation: Department of Fundamental Sciences, Université du Québec à Chicoutimi, Chicoutimi, QC, Canada.
authors.forename: Milla
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authors.lastname: Rautio
authors.affiliation: Office of Water Information, U.S. Geological Survey, Middleton, WI, USA.
authors.forename: Jordan S
authors.initials: JS
authors.lastname: Read
authors.affiliation: School of Geography, University of Nottingham, Nottingham, UK.
authors.forename: Sarah L
authors.initials: SL
authors.lastname: Roberts
authors.affiliation: Department of Freshwater Conservation, Brandenburg University of Technology Cottbus - Senftenberg, Bad Saarow, Germany.
authors.forename: Jacqueline
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authors.lastname: Rücker
authors.affiliation: Department of Environmental Science and Policy, University of California Davis, Davis, CA, USA.
authors.forename: Steven
authors.initials: S
authors.lastname: Sadro
authors.affiliation: Institute of Biology, Irkutsk State University, Irkutsk, Russian Federation.
authors.forename: Eugene A
authors.initials: EA
authors.lastname: Silow
authors.affiliation: Department of Biostatistics and Informatics, Colorado School of Public Health, Aurora, CO, USA.
authors.forename: Derek E
authors.initials: DE
authors.lastname: Smith
authors.affiliation: Department of Biology & Large Lakes Observatory, University of Minnesota Duluth, Duluth, MN, USA.
authors.forename: Robert W
authors.initials: RW
authors.lastname: Sterner
authors.affiliation: School of Geography, University of Nottingham, Nottingham, UK.
authors.forename: George E A
authors.initials: GE
authors.lastname: Swann
authors.affiliation: Institute of Biology, Irkutsk State University, Irkutsk, Russian Federation.
authors.forename: Maxim A
authors.initials: MA
authors.lastname: Timofeyev
authors.affiliation: Department of Aquatic Environment, Centre for Hydrographic Studies CEDEX, Madrid, Spain.
authors.forename: Manuel
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authors.lastname: Toro
authors.affiliation: Department of Biology, Clarkson University, Potsdam, NY, USA.
authors.forename: Michael R
authors.initials: MR
authors.lastname: Twiss
authors.affiliation: Department of Biology, Trent University, Peterborough, ON, Canada.
authors.forename: Richard J
authors.initials: RJ
authors.lastname: Vogt
authors.affiliation: Canada Centre for Inland Waters, Environment and Climate Change Canada, Burlington, ON, Canada.
authors.forename: Susan B
authors.initials: SB
authors.lastname: Watson
authors.affiliation: Department of Geography, Loughborough University, Loughborough, UK.
authors.forename: Erika J
authors.initials: EJ
authors.lastname: Whiteford
authors.affiliation: Department of Biology, Trent University, Peterborough, ON, Canada.
authors.forename: Marguerite A
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authors.lastname: Xenopoulos
mesh_terms: D017753:Ecosystem; D046448:Ice Cover; D060106:Lakes; D010933:Plankton / Q000502:physiology*; D012621:Seasons
keywords: Aquatic ecosystem; data synthesis; freshwater; lake; limnology; long-term; plankton; seasonal; time series; winter ecology