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Left in the Wake of the Ice: Using metagenomic sedimentary ancient DNA to reconstruct ecosystems before and after the Last Glacial Maximum in Sweden
Stockholm University, Faculty of Science, Department of Geological Sciences.ORCID iD: 0009-0004-0745-2437
2026 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

As the Fennoscandian ice-sheet retreated, plants and animals began to recolonise the Scandinavian peninsula. Current knowledge about the recolonisation patterns in Sweden during the Holocene is based mostly on fossil plant remains and preserved osteological remains, of which the latter record is sparse. Even less is known about the ecosystem composition before the Last Glacial Maximum (LGM), since the last ice-sheet eroded and reworked most deposits. Previous limitations due to gaps in the fossil record can now be addressed, as sedimentary ancient DNA (sedaDNA) is established as a powerful tool for reconstructing past ecosystems.

This thesis investigates how ancient environmental genomics based on shotgun sequencing of lake sediments can be used to reveal which species were present at certain points in time during the Holocene in central and northern Sweden. It also includes a glimpse into the Pleistocene pre-LGM ecosystem, as a comparison to the Holocene environment. 

In Paper I, we present an ancient environmental genome from a male brown bear extracted from lake sediment from northern Sweden dated to 9.6 cal. ka BP. With a mitochondrial coverage of 231x and a nuclear genome coverage of 0.05x, we could investigate the genome using population genomic methods. The paper highlights the potential of sedaDNA and non-destructive methods for generating high-quality genomic data. 

For metagenomic processing of sedaDNA data, an open-access ready-to-use bioinformatic pipeline was compiled from existing tools. The pipeline, called metaJAM, is presented in Paper II. It includes a pre-processing step of adapter trimming and removal of low complexity regions and microbial filtering, followed by competitive mapping against reference databases. Lastly, a module of filtering and authentication finalises the output.

From a sediment sequence from lake Orsasjön in central Sweden (Paper III), we reconstructed the local plant composition between ca. 8.1–0.1 cal. ka BP using sedaDNA. The results show a constant presence of woody taxa throughout the sediment sequence, with little turnover in the sedaDNA record. We demonstrate the expansion of spruce (Picea) in the region, with modelled ages overlapping with previous studies. The arrival of maple (Acer) to central Sweden, previously unreported, is also shown. Additionally, we present authenticated sedaDNA from vertebrates, including humans, fish, beaver and water vole.

There are some pre-LGM sediment deposits in Sweden left intact. In Paper IV, we investigated samples from Vålbacken outside Östersund, northern Sweden. The sediments are dated to 55–35 ka BP, i.e. within marine isotope stage 3 (MIS 3). We present indications of a mammoth steppe-like plant composition, and the presence of woolly mammoth (Mammuthus primigenius) in the sedaDNA. Our findings add to the sparse knowledge of pre-LGM ecosystems in Sweden.

A literature review is also included (Paper V). It outlines and summarises the setting for the project, and introduces many relevant concepts for readers not familiar with the field.

In conclusion, this thesis expands the knowledge of how the landscape in Scandinavia was developed before and after the LGM, based on shotgun sequenced sedaDNA. It provides an example of how lake sediments can yield genomic data from a single individual of comparable quality to osteological aDNA, reveals the arrival of maple to central Sweden, and confirms the presence of a mammoth steppe ecosystem in Sweden during MIS 3.

Place, publisher, year, edition, pages
Stockholm: Department of Geological Sciences, Stockholm University , 2026. , p. 51
Series
Meddelanden från Stockholms universitets institution för geologiska vetenskaper ; 399
Keywords [en]
Sedimentary ancient DNA, sedaDNA, eDNA, lake sediments, ancient environmental genomics, palaeogenetics, metagenomics, bioinformatics, palaeoecology, ecosystem reconstruction, Holocene, MIS 3, postglacial recolonisation
National Category
Multidisciplinary Geosciences Palaeontology and Palaeoecology
Research subject
Marine Geology
Identifiers
URN: urn:nbn:se:su:diva-253801ISBN: 978-91-8107-578-6 (print)ISBN: 978-91-8107-579-3 (electronic)OAI: oai:DiVA.org:su-253801DiVA, id: diva2:2049650
Public defence
2026-05-22, William-Olssonsalen, Geovetenskapens hus, Svante Arrhenius väg 14, and online via Zoom, public link is available at the department website, Stockholm, 13:00 (English)
Opponent
Supervisors
Available from: 2026-04-27 Created: 2026-03-30 Last updated: 2026-04-21Bibliographically approved
List of papers
1. Ancient environmental genome reveals a migratory brown bear individual in Early Holocene Scandinavia
Open this publication in new window or tab >>Ancient environmental genome reveals a migratory brown bear individual in Early Holocene Scandinavia
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2026 (English)In: Proceedings of the National Academy of Sciences of the United States of America, ISSN 0027-8424, E-ISSN 1091-6490, Vol. 123, no 16, article id e2527944123Article in journal (Refereed) Published
Abstract [en]

After the last ice age, species migrated into a newly deglaciated Scandinavia. Brown bear recolonization is thought to have occurred from two directions—from the south and the northeast—resulting in a nonoverlapping distribution of two distinct mitochondrial clades. A contact zone in central Sweden separates populations with mitochondrial clade 1a in the south from those with clade 3a in the north. However, a paucity of brown bear subfossils in Scandinavia has limited testing of this prevailing model using ancient DNA. Here, we present a high-coverage brown bear mitogenome (231×) and nuclear genome-wide data (0.05×) extracted from lake sediment dated to 9.6 cal. ka BP from northern Sweden, representing the oldest known record of brown bear in the region. At this point in the Early Holocene, the Fennoscandian Ice Sheet was in its final stages of recession. Surprisingly, our analyses suggest that this environmental genome represents one male individual carrying clade 1a and with southern brown bear nuclear ancestry, despite being found far north of the contact zone. This suggests the individual was a migratory bear and had dispersed northward from its birthplace. Our finding adds to the scarce genomic record of Early Holocene brown bears and highlights the use of sedimentary ancient DNA as a powerful source of genomic information.

Keywords
sedimentary ancient DNA, Ursus arctos, phylogeography, postglacial recolonization
National Category
Evolutionary Biology
Identifiers
urn:nbn:se:su:diva-253792 (URN)10.1073/pnas.2527944123 (DOI)41973920 (PubMedID)2-s2.0-105035679479 (Scopus ID)
Available from: 2026-03-30 Created: 2026-03-30 Last updated: 2026-04-22Bibliographically approved
2. metaJAM: a Nextflow integrated metagenomic workflow for sedimentary ancient DNA
Open this publication in new window or tab >>metaJAM: a Nextflow integrated metagenomic workflow for sedimentary ancient DNA
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(English)Manuscript (preprint) (Other academic)
Keywords
bioinformatics, sedaDNA, metagenomics
National Category
Bioinformatics and Computational Biology
Identifiers
urn:nbn:se:su:diva-253763 (URN)
Available from: 2026-03-30 Created: 2026-03-30 Last updated: 2026-04-09Bibliographically approved
3. Metagenomic analysis of lake sediment from central Sweden reveals the Holocene expansion of spruce (Picea) and maple (Acer).
Open this publication in new window or tab >>Metagenomic analysis of lake sediment from central Sweden reveals the Holocene expansion of spruce (Picea) and maple (Acer).
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(English)Manuscript (preprint) (Other academic)
Keywords
bioinformatics, sedaDNA, metagenomics, palaeoecology, Holocene
National Category
Palaeontology and Palaeoecology
Identifiers
urn:nbn:se:su:diva-253766 (URN)
Available from: 2026-03-30 Created: 2026-03-30 Last updated: 2026-04-09Bibliographically approved
4. Mammoth steppe ecosystem revealed by sedimentary ancient DNA from MIS 3 sediment deposit in Sweden
Open this publication in new window or tab >>Mammoth steppe ecosystem revealed by sedimentary ancient DNA from MIS 3 sediment deposit in Sweden
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(English)Manuscript (preprint) (Other academic)
Keywords
MIS 3, woolly mammoth, sedaDNA, sedimentary ancient DNA
National Category
Palaeontology and Palaeoecology
Identifiers
urn:nbn:se:su:diva-253799 (URN)
Available from: 2026-03-30 Created: 2026-03-30 Last updated: 2026-04-09Bibliographically approved
5. The potential of lacustrine sedimentary ancient DNA for revealing human postglacial recolonization patterns in northern Sweden – a review
Open this publication in new window or tab >>The potential of lacustrine sedimentary ancient DNA for revealing human postglacial recolonization patterns in northern Sweden – a review
2024 (English)In: Boreas, ISSN 0300-9483, E-ISSN 1502-3885, Vol. 53, no 3, p. 347-359Article, review/survey (Refereed) Published
Abstract [en]

The postglacial recolonization of Fennoscandian flora and fauna was initiated when the land became accessible as the last ice sheet retreated. In northern Sweden, plants are represented in pollen and macrofossil records, but there is no genetic evidence from the first plants, animals or humans in the region, mainly owing to an absence of osteological finds. The questions of who the first postglacial peoples, or pioneers, were and where they came from therefore remain unanswered. Previous palaeogenomic analyses from remains from adjacent regions have suggested that two main routes into Sweden could have been taken by the pioneers, one from the SW through modern-day Denmark and Norway, and one from the east via Finland. However, no direct genetic evidence from the pioneers of northern Sweden exists. Modern technology has provided the ancient DNA field with an updated toolbox that could allow for novel approaches for revealing the origin and genetic profiles of the first Scandinavians, of which sedimentary ancient DNA (sedaDNA) is well placed. Lake sediments are now a routine source of sedaDNA that have been used to record environmental changes and detect species that lived in the surrounding lake catchment. This review will provide context and background, a summary of the ground-breaking studies within the field of lacustrine sedaDNA, and relevant methodology to address the scientific questions at hand. We conclude that the field is mature enough to provide insight into the origins and arrival times of the first postglacial humans that migrated into northern Sweden.

National Category
Archaeology
Identifiers
urn:nbn:se:su:diva-231186 (URN)10.1111/bor.12660 (DOI)001230949300001 ()2-s2.0-85194465644 (Scopus ID)
Available from: 2024-06-24 Created: 2024-06-24 Last updated: 2026-03-30Bibliographically approved

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