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Jonsson, H., Blume-Werry, G., Wackett, A. A., Olofsson, J., Arvidsson, E., Sparrman, T. & Klaminder, J. (2025). Non-native earthworms alter carbon sequestration in arctic tundra ecosystems. Journal of Geophysical Research - Biogeosciences, 130(4), Article ID e2024JG008598.
Open this publication in new window or tab >>Non-native earthworms alter carbon sequestration in arctic tundra ecosystems
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2025 (English)In: Journal of Geophysical Research - Biogeosciences, ISSN 2169-8953, E-ISSN 2169-8961, Vol. 130, no 4, article id e2024JG008598Article in journal (Refereed) Published
Abstract [en]

Earthworms, as detritivores, play a significant role in breaking down soil organic carbon (SOC). The introduction of non-native earthworms to arctic ecosystems has, therefore, raised concerns about the potential impact they may have on one of the world's largest SOC reservoirs. Earthworms could also have considerable effects on plant productivity, and the lack of experimental studies quantifying their impact on carbon (C) reservoirs in both soil and plants makes it difficult to predict the effect of earthworms on ecosystem C storage. Here we experimentally tested how earthworms known to be non-native to arctic ecosystems (Aporrectodea spp. and Lumbricus spp.) affect C reservoirs in soil and plants (above and belowground separately) in two common tundra vegetation types (heath and meadow). Earthworms lowered the mean SOC pool and substantially altered SOC quality in meadow soils by increasing the proportion of aromatic-C compounds. Simultaneously, earthworms increased the C pool stored in plant biomass, which counteracted earthworm-induced SOC losses in meadow ecosystems. A positive earthworm effect on belowground biomass in heath soil facilitated a net ecosystem uptake of ∼0.84 kg C m−2 over the 4-year study period. The higher C uptake into plant biomass in the heath resulted in a notable increase of SOC but lower δ13C values, likely because of recently captured C being sourced from roots or litter. Our observations of vegetation-specific feedbacks between plants, earthworms, and soils advance our understanding of non-native earthworms' impact on SOC dynamics and C budgets in high-latitude ecosystems.

Place, publisher, year, edition, pages
American Geophysical Union (AGU), 2025
Keywords
invasive, lumbricidae, NMR, root, SOC, tundra
National Category
Ecology Soil Science
Identifiers
urn:nbn:se:umu:diva-238228 (URN)10.1029/2024JG008598 (DOI)001466915900001 ()2-s2.0-105002855693 (Scopus ID)
Funder
Swedish Research Council Formas, 2018‐01312
Available from: 2025-04-30 Created: 2025-04-30 Last updated: 2025-04-30Bibliographically approved
Jonsson, H., Olofsson, J., Blume-Werry, G. & Klaminder, J. (2024). Cascading effects of earthworm invasion increase graminoid density and rodent grazing intensities. Ecology, 105(2), Article ID e4212.
Open this publication in new window or tab >>Cascading effects of earthworm invasion increase graminoid density and rodent grazing intensities
2024 (English)In: Ecology, ISSN 0012-9658, E-ISSN 1939-9170, Vol. 105, no 2, article id e4212Article in journal (Refereed) Published
Abstract [en]

Human-mediated dispersal of non-native earthworms can cause substantial changes to the functioning and composition of ecosystems previously earthworm-free. Some of these earthworm species have the potential to “geoengineer” soils and increase plant nitrogen (N) uptake. Yet the possible consequences of increased plant N concentrations on rodent grazing remains poorly understood. In this study, we present findings from a common garden experiment with two tundra communities, meadow (forb dominated) and heath (shrub dominated), half of them subjected to 4 years of earthworm presence (Lumbricus spp. and Aporrectodea spp.). Within four summers, our earthworm treatment changed plant community composition by increasing graminoid density by, on average, 94% in the heath vegetation and by 49% in the meadow. Rodent winter grazing was more intense on plants growing in soils with earthworms, an effect that coincided with higher N concentrations in plants, indicating a higher palatability. Even though earthworms reduced soil moisture, plant community productivity, as indicated by vegetation greenness (normalized difference vegetation index), was not negatively impacted. We conclude that earthworm-induced changes in plant composition and trophic interactions may fundamentally alter the functioning of tundra ecosystems.

Place, publisher, year, edition, pages
The Ecological Society of America, 2024
Keywords
earthworms, grazing, Lumbricidae, non-native, plant community, soil moisture, tundra
National Category
Ecology
Identifiers
urn:nbn:se:umu:diva-218292 (URN)10.1002/ecy.4212 (DOI)001121395900001 ()37996966 (PubMedID)2-s2.0-85179362361 (Scopus ID)
Available from: 2023-12-22 Created: 2023-12-22 Last updated: 2024-09-03Bibliographically approved
Leander, J., Hellström, G., Nordin, J., Jonsson, M. & Klaminder, J. (2024). Guiding downstream migrating Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) of different life stages in a large river using bubbles. Rivers Research and Applications: an international journal devoted to river research and management, 40(1), 107-115
Open this publication in new window or tab >>Guiding downstream migrating Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) of different life stages in a large river using bubbles
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2024 (English)In: Rivers Research and Applications: an international journal devoted to river research and management, ISSN 1535-1459, E-ISSN 1535-1467, Vol. 40, no 1, p. 107-115Article in journal (Refereed) Published
Abstract [en]

Salmonid repeat spawners are precious individuals for wild populations due to their high fecundity and previous spawning experience, making them important in environmental policy. However, repeat spawners rarely exist above hydropower dams in regulated rivers as the mortality of post-spawners (kelts) when passing through turbines during downstream migration is very high. To mitigate this problem, there are different technical solutions that potentially guide fish toward available fishways. Bubble barriers represent one alternative to costly physical guiding structures, but the efficiency of bubbles for guiding downstream migrating kelts has not been tested. In this study, we evaluate a 100 m long bubble barrier in guiding salmonids—both smolts and kelts—away from the main current and toward an alternative fishway in Ume River, a large regulated river in northern Sweden. We used both acoustic telemetry and sonar to measure the guiding effect of the bubble barrier for downstream migrating fish. We found that more than twice as many salmonids chose the alternative fishway when the bubble barrier was turned on. This was true both for smolts and kelts, suggesting that bubble barriers can be used to guide salmonids of different life stages in rivers with flow rates over 500 m3 s−1. Indeed, our study indicates that bubble barriers are low-cost structures that could be rapidly applied in many regulated rivers to support salmonid migration. 

Place, publisher, year, edition, pages
John Wiley & Sons, 2024
Keywords
Atlantic salmon, Brown trout, bubble curtain, fish migration, fish passage, hydropower
National Category
Ecology
Identifiers
urn:nbn:se:umu:diva-186819 (URN)10.1002/rra.4209 (DOI)001077498200001 ()2-s2.0-85170712321 (Scopus ID)
Funder
Swedish Energy Agency, 48412-1Vattenfall ABUmeå University
Note

Originally included in thesis in manuscript form. 

Available from: 2021-08-24 Created: 2021-08-24 Last updated: 2024-01-12Bibliographically approved
Klaminder, J., Fassl, M., Baudet, M., Östlund, L., Linderholm, J. & Zale, R. (2024). Landscape of ice and fire: uniquely well-preserved scots pine trunks reveal forest fires near the retreating weichselian ice margin. Vegetation History and Archaeobotany, 33, 519-527
Open this publication in new window or tab >>Landscape of ice and fire: uniquely well-preserved scots pine trunks reveal forest fires near the retreating weichselian ice margin
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2024 (English)In: Vegetation History and Archaeobotany, ISSN 0939-6314, E-ISSN 1617-6278, Vol. 33, p. 519-527Article in journal (Refereed) Published
Abstract [en]

Environmental conditions for trees that established in central Fennoscandia shortly after the final retreat of the Weichselian ice sheet remain poorly understood. In this study we examine tree rings of five well-preserved Pinus sylvestris (Scots pines) that grew in the area in front of the retreating ice sheet in northern Sweden. They became buried in flood sediments deposited by a glacial outburst flood (jökulhlaup) about 9.5–9.9 kyr cal bp and the aim of our study was to search for information regarding damage from fires and bioclimatic conditions in their ancient tree ring records. Our analysis, providing a glimpse into the local early Holocene environment in north-central Sweden, suggests that: 1, there were repeated fires (four fire events detected) during the early Holocene; and 2, bioclimatic conditions when the ancient pines were growing resembled those of modern sub-alpine pine woods. The latter is indicated by their patterns of tree ring growth (growth rate and variation), which were statistically similar to those of pines growing in sub-alpine woods with an open canopy, but different from pines in protected and managed boreal forests. Lower δ13C for the ancient latewood in comparison to pine wood from trees growing near the Scandinavian mountains before the 1850s were probably caused both by stomata fractionation due to lower atmospheric CO2 during the early Holocene and by the moist local environment created by the nearby ancient Ancylus lake, which preceded the Baltic Sea. Periods with cloudy and cold summers were also indicated by the occurrence of ‘false rings’. Finds of charred fragments of Calluna vulgaris (heather, ling), an understory shrub that can burn even with a relatively high moisture content, suggest that heath vegetation was crucial to make fire a reoccurring ecological factor in the area during the early Holocene.

Place, publisher, year, edition, pages
Springer, 2024
Keywords
Climate, Early Holocene, Forest fires, Forest history, Lycksele, Pinus sylvestris
National Category
Forest Science
Identifiers
urn:nbn:se:umu:diva-218105 (URN)10.1007/s00334-023-00974-6 (DOI)001118670600002 ()2-s2.0-85178969112 (Scopus ID)
Funder
Swedish Research Council, 2017–04548Wallenberg Foundations, WAF
Available from: 2023-12-15 Created: 2023-12-15 Last updated: 2024-06-25Bibliographically approved
Larsbo, M., Koestel, J., Krab, E. J. & Klaminder, J. (2024). Quantifying earthworm soil ingestion from changes in vertical bulk density profiles. European journal of soil biology, 120, Article ID 103574.
Open this publication in new window or tab >>Quantifying earthworm soil ingestion from changes in vertical bulk density profiles
2024 (English)In: European journal of soil biology, ISSN 1164-5563, E-ISSN 1778-3615, Vol. 120, article id 103574Article in journal (Refereed) Published
Abstract [en]

Soil mixing by earthworms can have a large impact on the fate of nutrients and pollutants and on the soil's ability to sequester carbon. Nevertheless, methods to quantify earthworm ingestion and egestion under field conditions are largely lacking. Soils of the Fennoscandian tundra offer a special possibility for such quantifications, as these soils commonly lack burrowing macrofauna and exhibit a well-defined O horizon with low bulk density on top of a mineral soil with higher density. Since ingestion-egestion mixes the two soil layers, the temporal changes in the bulk density profile of such soils may be useful for estimating field ingestion rates. In this study, we applied a model for earthworm burrowing through soil ingestion to observed changes in soil densities occurring in a mesocosm experiment carried out in the arctic during four summers with intact soil. The earthworms present in the mesocosms were Aporrectodea trapezoides, Aporrectodea tuberculata, Aporrectodea rosea, Lumbricus rubellus and Lumbricus Terrestris (fourth season only). We show that changes in soil density profiles can indeed be used to infer earthworm ingestion rates that are realistic in comparison to literature values. Although uncertainties in parameter values were sometimes large, the results from this study suggest that soil turnover rates and endogeic earthworm soil ingestion rates in tundra heath and meadow soils may be as high as those reported for temperate conditions. Such large ingestion rates can explain observed large morphological changes in arctic soils where dispersing earthworms have resulted in complete inmixing of the organic layer into the mineral soil. Our approach is applicable to soil profiles with marked vertical differences in bulk density such as the soils of the Fennoscandian tundra where earthworms are currently dispersing into new areas and to layered repacked soil samples that are incubated in the field.

Place, publisher, year, edition, pages
Elsevier, 2024
National Category
Soil Science Ecology
Identifiers
urn:nbn:se:umu:diva-218630 (URN)10.1016/j.ejsobi.2023.103574 (DOI)001141525900001 ()2-s2.0-85179614919 (Scopus ID)
Funder
Swedish Research Council Formas, 201-01312
Available from: 2023-12-27 Created: 2023-12-27 Last updated: 2025-04-24Bibliographically approved
Morlock, M. A., Rodriguez-Martinez, S., Huang, D.-T. Y. & Klaminder, J. (2023). Erosion regime controls sediment environmental DNA-based community reconstruction. Environmental DNA, 5(6), 1393-1404
Open this publication in new window or tab >>Erosion regime controls sediment environmental DNA-based community reconstruction
2023 (English)In: Environmental DNA, E-ISSN 2637-4943, Vol. 5, no 6, p. 1393-1404Article in journal (Refereed) Published
Abstract [en]

Analysis of environmental DNA detected in lake sediments shows promise to become a great paleoecological technique that can provide detailed information about organism communities living in past environments. However, when interpreting sedimentary environmental DNA records, it is of crucial importance to separate ecosystem responses to large-scale environmental change from “noise” caused by changes in sediment provenance or potential post-depositional DNA mobility. In this study, we show that plant and mammalian communities reconstructed from sediments are strongly affected by sediment provenance, but unaffected by vertical mobility of DNA after sediment deposition. We observe that DNA from aquatic plants was abundant in background sediment, while embedded detrital event layers (sediment deposited under erosion events) primarily contained terrestrial plants; hence, vertical mobility of aquatic plant DNA across sediment layers was negligible within our studied lakes. About 33% of the identified terrestrial plant genera were only found in detrital sediment, suggesting that sediment origin had a strong impact on the reconstructed plant community. Similarly, DNA of some mammalian taxa (Capra hircus, Ursus arctos, Lepus, and Felis) were only or preferentially found in detrital event layers. Temporal changes across the Holocene were the main drivers of change for reconstructed plant communities, but sediment type was the second most important factor of variance. Our results highlight that erosion and sediment provenance need to be considered when reconstructing past mammalian and plant communities using environmental DNA from lake sediments.

Place, publisher, year, edition, pages
John Wiley & Sons, 2023
Keywords
DNA taphonomy, erosion, Lake Grosssee, lake sediment, paleoecology, sedaDNA, Switzerland
National Category
Environmental Sciences Ecology
Identifiers
urn:nbn:se:umu:diva-212991 (URN)10.1002/edn3.458 (DOI)001306414300022 ()2-s2.0-85167338602 (Scopus ID)
Funder
Swedish Research Council, 2017-04548The Kempe Foundations
Available from: 2023-08-21 Created: 2023-08-21 Last updated: 2025-04-24Bibliographically approved
Klaminder, J., Krab, E. J., Larsbo, M., Jonsson, H., Fransson, J. & Koestel, J. (2023). Holes in the tundra: Invasive earthworms alter soil structure and moisture in tundra soils. Science of the Total Environment, 859, Article ID 160125.
Open this publication in new window or tab >>Holes in the tundra: Invasive earthworms alter soil structure and moisture in tundra soils
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2023 (English)In: Science of the Total Environment, ISSN 0048-9697, E-ISSN 1879-1026, Vol. 859, article id 160125Article in journal (Refereed) Published
Abstract [en]

Human introductions have resulted in earthworms establishing in the Arctic, species known to cause cascading ecosystem change. However, few quantitative outdoor experiments have been performed that describe how these soil modifying earthworms are reshaping structures in tundra soils. In this study, we used three-dimensional (3-D) X-ray images of soil cores (approximately 10 cm diameter, 20 cm height, N = 48) to assess how earthworms (Aporrectodea sp. and Lumbricus sp.) affect soil structure and macropore networks in an outdoor mesocosm experiment that lasted four summers. Effects were assessed in both shrub-dominated (heath) and herb-dominated (meadow) tundra. Earthworms almost doubled the macroporosity in meadow soils and tripled macroporosity in heath. Interestingly, the fractal dimension of macropores decreased in response to earthworm burrowing in both systems, indicating that the presence of earthworms reduced the geometric complexity in comparison to other pore-generating processes active in the tundra. Observed effects on soil structure occurred along with a dramatically reduced soil moisture content, which was observed the first winter after earthworm introduction in the meadow. Our findings suggest that predictions of future changes in vegetation and soil carbon pools in the Arctic should include major impacts on soil properties that earthworms induce.

Place, publisher, year, edition, pages
Elsevier, 2023
Keywords
3D, Abisko, Bioturbation, Heath, Long-term, Meadow, Soil-mixing
National Category
Soil Science Ecology
Identifiers
urn:nbn:se:umu:diva-201338 (URN)10.1016/j.scitotenv.2022.160125 (DOI)000898837900015 ()36379337 (PubMedID)2-s2.0-85142356755 (Scopus ID)
Funder
Swedish Research CouncilSwedish Research Council Formas
Available from: 2022-12-14 Created: 2022-12-14 Last updated: 2024-07-02Bibliographically approved
Blume-Werry, G., Klaminder, J., Krab, E. J. & Onteux, S. (2023). Ideas and perspectives: Alleviation of functional limitations by soil organisms is key to climate feedbacks from arctic soils. Biogeosciences, 20(10), 1979-1990
Open this publication in new window or tab >>Ideas and perspectives: Alleviation of functional limitations by soil organisms is key to climate feedbacks from arctic soils
2023 (English)In: Biogeosciences, ISSN 1726-4170, E-ISSN 1726-4189, Vol. 20, no 10, p. 1979-1990Article in journal (Refereed) Published
Abstract [en]

Arctic soils play an important role in Earth's climate system, as they store large amounts of carbon that, if released, could strongly increase greenhouse gas levels in our atmosphere. Most research to date has focused on how the turnover of organic matter in these soils is regulated by abiotic factors, and few studies have considered the potential role of biotic regulation. However, arctic soils are currently missing important groups of soil organisms, and here, we highlight recent empirical evidence that soil organisms' presence or absence is key to understanding and predicting future climate feedbacks from arctic soils. We propose that the arrival of soil organisms into arctic soils may introduce "novel functions", resulting in increased rates of, for example, nitrification, methanogenesis, litter fragmentation, or bioturbation, and thereby alleviate functional limitations of the current community. This alleviation can greatly enhance decomposition rates, in parity with effects predicted due to increasing temperatures. We base this argument on a series of emerging experimental evidence suggesting that the dispersal of until-then absent micro-, meso-, and macroorganisms (i.e. from bacteria to earthworms) into new regions and newly thawed soil layers can drastically affect soil functioning. These new observations make us question the current view that neglects organism-driven "alleviation effects"when predicting future feedbacks between arctic ecosystems and our planet's climate. We therefore advocate for an updated framework in which soil biota and the functions by which they influence ecosystem processes become essential when predicting the fate of soil functions in warming arctic ecosystems.

Place, publisher, year, edition, pages
Copernicus Publications, 2023
National Category
Soil Science
Identifiers
urn:nbn:se:umu:diva-212060 (URN)10.5194/bg-20-1979-2023 (DOI)000998715900001 ()2-s2.0-85163878232 (Scopus ID)
Funder
Swedish Research Council, 2021-04458Swedish Research Council Formas, 2018-01312
Available from: 2023-07-18 Created: 2023-07-18 Last updated: 2023-07-18Bibliographically approved
Jerand, P., Klaminder, J. & Linderholm, J. (2023). The legacy of ecological imperialism in the Scandes: earthworms and their implications for Arctic research. Arctic, Antarctic and Alpine research, 55(1), Article ID 2274650.
Open this publication in new window or tab >>The legacy of ecological imperialism in the Scandes: earthworms and their implications for Arctic research
2023 (English)In: Arctic, Antarctic and Alpine research, ISSN 1523-0430, E-ISSN 1938-4246, Vol. 55, no 1, article id 2274650Article in journal (Refereed) Published
Abstract [en]

In the nineteenth century, numerous settlements were established in the alpine region of Fennoscandia (the Scandes), an area that later became a major international scene for Arctic research. Here we raise awareness of this era and show that earthworm-driven bioturbation in “pristine” soils around contemporary Arctic research infrastructure is caused by soil fauna left behind during early land use. We use soil preserved under an alpine settlement to highlight that soils were not bioturbated when the first house was built at a site where bioturbation is now widespread. A review of archived material with unique site-specific chronology constrained the onset of bioturbation to the post-1871 era. Our results suggest that small-scale land use introduced earthworms that now thrive far beyond the realms of former cultivated fields. The legacy of soil fauna from this example of “ecological imperialism” still lingers and should be considered when studying soils of the Scandes.

Place, publisher, year, edition, pages
Taylor & Francis, 2023
Keywords
Archaeology, bioturbation, historical sources, phosphate analysis, soil classification
National Category
Ecology Soil Science
Research subject
Archaeology; environmental archaeology
Identifiers
urn:nbn:se:umu:diva-217053 (URN)10.1080/15230430.2023.2274650 (DOI)001105759300001 ()2-s2.0-85177559253 (Scopus ID)
Available from: 2023-11-27 Created: 2023-11-27 Last updated: 2025-04-24Bibliographically approved
Rodriguez-Martinez, S., Klaminder, J., Morlock, M. A., Dalén, L. & Huang, D.-T. Y. (2023). The topological nature of tag jumping in environmental DNA metabarcoding studies. Molecular Ecology Resources, 23(3), 621-631
Open this publication in new window or tab >>The topological nature of tag jumping in environmental DNA metabarcoding studies
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2023 (English)In: Molecular Ecology Resources, ISSN 1755-098X, E-ISSN 1755-0998, Vol. 23, no 3, p. 621-631Article in journal (Refereed) Published
Abstract [en]

Metabarcoding of environmental DNA constitutes a state-of-the-art tool for environmental studies. One fundamental principle implicit in most metabarcoding studies is that individual sample amplicons can still be identified after being pooled with others—based on their unique combinations of tags—during the so-called demultiplexing step that follows sequencing. Nevertheless, it has been recognized that tags can sometimes be changed (i.e., tag jumping), which ultimately leads to sample crosstalk. Here, using four DNA metabarcoding data sets derived from the analysis of soils and sediments, we show that tag jumping follows very specific and systematic patterns. Specifically, we find a strong correlation between the number of reads in blank samples and their topological position in the tag matrix (described by vertical and horizontal vectors). This observed spatial pattern of artefactual sequences could be explained by polymerase activity, which leads to the exchange of the 3′ tag of single stranded tagged sequences through the formation of heteroduplexes with mixed barcodes. Importantly, tag jumping substantially distorted our data sets—despite our use of methods suggested to minimize this error. We developed a topological model to estimate the noise based on the counts in our blanks, which suggested that 40%–80% of the taxa in our soil and sedimentary samples were likely false positives introduced through tag jumping. We highlight that the amount of false positive detections caused by tag jumping strongly biased our community analyses.

Place, publisher, year, edition, pages
John Wiley & Sons, 2023
Keywords
a-DNA, detection limits, e-DNA, false positive, index hopping, sample crosstalk
National Category
Ecology
Identifiers
urn:nbn:se:umu:diva-203577 (URN)10.1111/1755-0998.13745 (DOI)000908019900001 ()36479848 (PubMedID)2-s2.0-85145725694 (Scopus ID)
Funder
Swedish Research Council, 2017- 04548
Available from: 2023-01-19 Created: 2023-01-19 Last updated: 2024-09-16Bibliographically approved
Organisations
Identifiers
ORCID iD: ORCID iD iconorcid.org/0000-0001-8814-0013

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