Citation: Filipsson, K.; Åsman, V.;
Greenberg, L.; Österling, M.; Watz, J.;
Bergman, E. Winter Behavior of
Juvenile Brown Trout in a Changing
Climate: How Do Light and Ice
Cover Affect Encounters with
Instream Predators? Fishes 2023, 8,
521. https://doi.org/10.3390/
fishes8100521
Academic Editor: Assaf Barki
Received: 14 September 2023
Revised: 13 October 2023
Accepted: 16 October 2023
Published: 20 October 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
fishes
Article
Winter Behavior of Juvenile Brown Trout in a Changing
Climate: How Do Light and Ice Cover Affect Encounters with
Instream Predators?
Karl Filipsson
1,2
, Veronika Åsman
1
, Larry Greenberg
1,
* , Martin Österling
1
, Johan Watz
1
and Eva Bergman
1
1
River Ecology and Management, Department of Environmental and Life Sciences, Karlstad University,
Universitetsgatan 2, SE-651 88 Karlstad, Sweden; martin.osterling@kau.se (M.Ö.); johan.watz@kau.se (J.W.);
eva.bergman.1868@kau.se (E.B.)
2
Jakobi Sustainablility AB, SE-412 58 Gothenburg, Sweden
* Correspondence: larry.greenberg@kau.se; Tel.: +46-54-7001543
Abstract: During winter, stream fishes are vulnerable to semi-aquatic predators like mammals and
birds and reduce encounters by being active in darkness or under surface ice. Less is known about
the behavior of fishes towards instream piscivorous fishes. Here, we examined how surface ice and
light affected the anti-predator behavior of juvenile brown trout (Salmo trutta Linnaeus, 1758) in
relation to piscivorous burbot (Lota lota Linnaeus, 1758) and northern pike (Esox lucius Linnaeus,
1758) at 4
◦
C in experimental flumes. Trout had lower foraging and swimming activity and spent
more time sheltering when predators were present than when absent. In daylight, trout’s swimming
activity was not affected by predators, whereas in darkness trout were less active when predators
were present. Trout consumed more drifting prey during the day when ice was present, and they
positioned themselves further upstream when under ice cover, regardless of light conditions. Trout
stayed closer to conspecifics under ice, but only in the presence of pike. Piscivorous fishes thus
constitute an essential part of the predatory landscape of juvenile trout in winter, and thus loss of ice
cover caused by climate warming will likely affect trout’s interactions with predators.
Keywords: anti-predator; global changer; diel behavior; foraging; piscivores; predators
Key Contribution: Most studies involving predation on fishes in northern temperate streams in
winter have focused on prey behavior in relation to semi-aquatic predators like mammals and birds.
Here, we show that the prey also modify their behavior in the presence of instream piscivorous fishes
and how this interaction is affected by surface ice cover and light.
1. Introduction
The global climate is gradually becoming warmer [1], with the magnitude of the effect
dependent on location and time of year. In most northern latitudes, it is expected that
temperature increases will be more pronounced during winter than during summer [2].
Hence, the effects of global warming during winter may be profound and have direct and
indirect effects on organisms [3–5]. In aquatic systems situated in northern latitudes, the
effects of warmer winters on the physical environment have already been documented.
Warmer winters have resulted in reduced snow and ice cover, both temporally and spatially,
as well as erratic ice cover formation and breakup throughout winter [6–10]. Surface ice
cover is thought to protect ectothermic fish from many endothermic predators, and thus
ongoing and expected further reductions of the period with intact surface ice cover may
have negative effects for fish.
Many juvenile salmonids overwinter in streams and rivers, where they are vulner-
able to predation [11–13]. Semi-aquatic predators, such as mammals and birds, both
Fishes 2023, 8, 521. https://doi.org/10.3390/fishes8100521 https://www.mdpi.com/journal/fishes