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Ecosystem Engineering by Mussels (Margaritifera margaritifera) Influences the Behaviour of Their Host Fish Brown Trout (Salmo trutta) Under Various Flows
Karlstad University, Faculty of Health, Science and Technology (starting 2013), Department of Environmental and Life Sciences (from 2013). University of Gothenburg, Sweden.ORCID iD: 0000-0001-8026-3457
Karlstad University, Faculty of Health, Science and Technology (starting 2013), Department of Environmental and Life Sciences (from 2013).ORCID iD: 0009-0002-3032-229X
Karlstad University, Faculty of Health, Science and Technology (starting 2013), Department of Environmental and Life Sciences (from 2013).ORCID iD: 0000-0001-6758-5857
2026 (English)In: Freshwater Biology, ISSN 0046-5070, E-ISSN 1365-2427, Vol. 71, no 1, article id e70169Article in journal (Refereed) Published
Abstract [en]

Ecosystem engineers modify habitats in many ways with consequences for other species. Aggregations of autogenic ecosystem engineers in lotic systems, such as unionid mussels, can increase habitat complexity and change water flow, affecting the diversity and abundance of other benthic species. The effects of unionid mussel aggregations on mobile species, such as their host fishes, are less well-studied, but the increase in habitat complexity caused by mussels may provide refuges from water flow and may change fish behaviour. Using stream flumes, we examined how the presence and density (zero/low density/high density) of the freshwater pearl mussel (Margaritifera margaritifera) and water flow (low/high) affected behaviour of three different size classes of young-of-the-year brown trout (Salmo trutta) in May, June and August 2023, respectively. We found that small-sized trout were more likely to reside on the bottom at high than at low water flow but increased the likelihood to swim when mussel density increased. Medium-sized trout were more likely to reside on the bottom during high flow at zero to low mussel densities, while the proportions of swimming individuals increased regardless of flow at high mussel density. Large-sized trout were instead more likely to reside on the bottom when mussel densities were high, and in contrast to the small- and medium-sized trout, positioned themselves within the mussel zone more often. Our results demonstrate that freshwater pearl mussels can influence juvenile trout behaviour, probably by modifying habitat structure and flow conditions in the downstream area nearest to the mussels, particularly under hydrological stress. These findings underscore the role of mussels as ecosystem engineers and highlight potential feedback loops between freshwater pearl mussel presence and the behaviour of juvenile brown trout, which may be highly relevant for conservation strategies targeting both species. 

Place, publisher, year, edition, pages
John Wiley & Sons, 2026. Vol. 71, no 1, article id e70169
Keywords [en]
brown trout early life stage, flow alteration, habitat degradation, habitat use, loss of mussel beds
National Category
Biological Sciences
Research subject
Biology
Identifiers
URN: urn:nbn:se:kau:diva-108247DOI: 10.1111/fwb.70169ISI: 001676294200002Scopus ID: 2-s2.0-105026883657OAI: oai:DiVA.org:kau-108247DiVA, id: diva2:2029975
Note

Artikeln tidigare publicerad som manuskript i Gals (2025) doktorsavhandling Ecosystem Engineering by Freshwater Mussels: Effects on Macroinvertebrate Communities, Decomposition Processes and Fish Behaviour

Available from: 2026-01-19 Created: 2026-01-19 Last updated: 2026-02-23Bibliographically approved
In thesis
1. Ecosystem Engineering by Freshwater Mussels: Effects on Macroinvertebrate Communities, Decomposition Processes and Fish Behaviour
Open this publication in new window or tab >>Ecosystem Engineering by Freshwater Mussels: Effects on Macroinvertebrate Communities, Decomposition Processes and Fish Behaviour
2025 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

Ecosystem engineers are organisms whose activities modify habitats, and thus also resource availability, biodiversity, and behaviour of other species. Freshwater mussels (order Unionida) are a group of ecosystem engineers that encompasses many threatened species. Freshwater mussels modify habitats by removing suspended particles from the water and by depositing particles on the sediment, as well as via bioturbation. By the presence of their shells, they increase substrate complexity and modify near-benthic flows. Our understanding of how engineering by mussels affects different organisms and processes, especially in brown food webs, is limited. Understanding how species, communities and processes are affected by mussels can help us understand how ecosystems might respond to the ongoing decline of many mussel populations.

I evaluated how the presence and density of freshwater mussels affected decomposition of leaf litter, macroinvertebrate communities, and the behaviour of juvenile brown trout, using a combination of field and laboratory experiments. In a field enclosure experiment, mussel presence slowed total decomposition of leaf litter, while mussel biodeposits did not change decomposition rates in microcosms. I found that mussels affected many response variables related to the macroinvertebrate communities by decreasing the effect of flow. For example, community evenness decreased with flow in a field enclosure experiment, but this effect decreased with increasing mussel densities. This is likely because blackfly (Simuliidae) larvae, the most abundant taxon, did not benefit from flow as much at higher mussel densities. In stream mesocosms, mussel beds allowed smaller juvenile trout to swim more at high flows, while larger juvenile trout spent more time in proximity to the mussels beds themselves. This thesis highlights how mussels affect boreal ecosystems, and the importance of conservation and reintroduction of mussels for naturally functioning ecosystems.

Abstract [sv]

Förlust av biodiversitet, inte minst i sötvatten, hotar att påverka många ekosystemsprocesser negativt. Vissa organismer har en större effekt på hur ekosystem ser ut och fungerar, t.ex. genom att kontrollera hur resten av systemet eller artsamhället ser ut. Ekosystemsingenjörer är en sådan kategori. Ekosystemsingenjörer är organismer som, genom sitt beteende eller struktur, påverkar den fysiska miljön samt fördelningen av resurser i systemet. Sötvattensmusslorna av ordningen Unionida, som i Sverige representeras av flodpärlmusslan, målarmusslorna och dammusslorna, är en grupp av viktiga ekosystemsingenjörer i sötvatten. Musslorna filtrerar partiklar ur vattnet, deponerar partiklar på sjö- och vattendragsbotten och syresätter sediment genom att gräva. Dessutom kan täta musselbestånd minska flödeshastigheten närmast botten i vattendrag. Populationerna för en stor andel av arterna i gruppen har minskat kraftigt på grund av människans inverkan, och många riskerar att dö ut. Därför är det viktigt att förstå hur deras närvaro och försvinnande kan påverka andra arter och processer.

Jag har undersökt hur närvaron och tätheten av musslor påverkar lövnedbrytning, olika aspekter av akvatiska evertebratsamhällen i vattendrag och fiskbeteende. I en fältstudie i en bäck i Örebro län under sen-höst och vinter har jag satt ut burar med olika tätheter av flodpärlmusslor, och såg att löv bröts ner långsammare när musslor var närvarande. I ett laboratorieexperiment, hade inte musslors biodepositioner (avföring samt pseudofekalier, partiklar som musslorna valde att inte äta) inte påverkade lövnedbrytningen. Musslorna i fältstudien hade dessutom påverkat evertebrat-samhällen som levde i burarna, främst genom att motverka strömhastighetens effekter på olika djuren. Knott-larver var den vanligaste gruppen i burarna. Knott gynnades av högre strömhastigheter, medan andra fanns i högre antal vid lägre flöden. Ökad musseltäthet ledde till samhällen med relativt mindre knott, relativt fler andra arter och där antalet individer av olika arter var jämnare. Förändring av flöde nära botten visade sig också vara en viktig mekanism för hur musslor kan påverka fiskbeteende. I ett laboratorieexperiment i strömakvarier visade det sig att små öringar (3 – 4 cm i längd) simmade oftare när musslor var närvarande, och att större unga öringar (ca 8 cm) befann sig då oftare högst uppströms i närheten till musslorna. Denna avhandling visar en del av sätten som sötvattensmusslor påverkar sötvattensekosystem i Norden. Den dramatiska minskningen av vissa sötvattensmusslors populationer lär redan ha förändrat många sötvattenssystem, men bevarande och återintroduktion av hotade musslor kan bidra att återskapa den naturliga dynamiken i våra vattendrag.

Abstract [en]

Ecosystem engineers are organisms whose activities modify habitats, and thus also resource availability, biodiversity, and the behaviour of other species. Freshwater mussels modify habitats by removing suspended particles from water, depositing particles on the bottom, and by modification of near-benthic flows. Understanding how ecosystems are affected by mussels can help us understand how systems might be responding to ongoing declines of many mussel populations.

I examined how freshwater mussels affected leaf decomposition, macroinvertebrate communities, and the behaviour of juvenile brown trout. In the field, mussel presence slowed total decomposition of leaf litter, while mussel biodeposits did not change decomposition rates in the laboratory. Mussels decreased near-benthic flow, an effect that in the field decreased the otherwise dominant blackfly larvae and benefitted other taxa, increasing community evenness and shifting community composition. In stream aquaria, mussel beds allowed smaller juvenile trout to swim more at high flows, while larger juvenile trout spent more time upstream near the mussel beds. This thesis highlights how mussels affect boreal ecosystems, and the importance of conservation and reintroduction of mussels for naturally functioning ecosystems.

Place, publisher, year, edition, pages
Karlstad: Karlstads universitet, 2025. p. 58
Series
Karlstad University Studies, ISSN 1403-8099 ; 2025:7
Keywords
Mussels, Unionida, Decomposition, Freshwater ecology, Macroinvertebrates, Simuliidae, Ecosystem engineers, Fish, Trout, Substrate complexity, Near-benthic flow, Stream ecology, צדפות נחלים, רקבון, אקולוגיה, מים מתוקים, חרקים, ישחוריים, מין מהנדס סביבה, Musslor, Sötvattensmusslor, Nedbrytning, Sötvattensekologi, Makroevertebrater, Knott, Ekosystemsingenjörer, Fisk, Öring, Vattendrag
National Category
Ecology
Research subject
Biology
Identifiers
urn:nbn:se:kau:diva-102942 (URN)10.59217/wzbf1566 (DOI)978-91-7867-540-1 (ISBN)978-91-7867-541-8 (ISBN)
Public defence
2025-03-14, Sjöströmsalen, 1B 309, Universitetsgatan 2, Karlstad, 10:00 (English)
Opponent
Supervisors
Available from: 2025-02-21 Created: 2025-01-31 Last updated: 2026-02-12Bibliographically approved

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Gal, RavivÖsterling, Martin

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