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Divergent evolutionary processes associated with colonization of offshore islands

Identifieur interne : 001657 ( Istex/Corpus ); précédent : 001656; suivant : 001658

Divergent evolutionary processes associated with colonization of offshore islands

Auteurs : Natália Martínková ; Ross Barnett ; Thomas Cucchi ; Rahel Struchen ; Marine Pascal ; Michel Pascal ; Martin C. Fischer ; Thomas Higham ; Selina Brace ; Simon Y. W. Ho ; Jean-Pierre Quéré ; Paul O'Higgins ; Laurent Excoffier ; Gerald Heckel ; A. Rus Hoelzel ; Keith M. Dobney ; Jeremy B. Searle

Source :

RBID : ISTEX:48CC05C77BA9B2A631CBFCFFA3B9D019915362C5

Abstract

Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.

Url:
DOI: 10.1111/mec.12462

Links to Exploration step

ISTEX:48CC05C77BA9B2A631CBFCFFA3B9D019915362C5

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<div type="abstract">Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.</div>
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<note>Data S1 Material and Methods. Table S1 List of modern M. arvalis samples used for morphometrics, including country and site of origin. Table S2 List of all modern M. arvalis used for cytb analysis and collected for this study, including country and site of origin; arranged according to cytb haplotype. Table S3 List of all ancient specimens of M. arvalis that successfully provided a cytb sequence with details of location collected, calibrated age range (where obtained) and GenBank Accession Number for the sequence. Table S4 List of primers used for the amplification of cytochrome b from M. arvalis. Table S5 Prior distributions of the ABC model parameters (as illustrated in Fig. S3). Table S6 Pairwise FST values between population samples analysed with microsatellites (see Table  and Fig. S1). Fig. S1 Map showing distribution of population samples of modern M. arvalis used for microsatellite typing, labelled for mtDNA lineage. Population names as listed in Table : 1 – Heerenveen, 2 – Dinteloord, 3 – Stalhille, 4 – Veurne, 5 – Pihen lès Guînes, 6 – Fressenneville, 7 – Daubeuf, 8 – Thaon, 9 – Ste Marie du Mont, 10 – St Jean du Thomas, 11 – Baie d'Aiguillon, 12 – Aiffres, 13 – Avallon, 14 – Clérmont‐Ferrand, 15 – Alflen, 16 – Schiltach, 17 – Loch of Swartmill, 18 – Ness, 19 – Whitemill Bay, 20 – Settiscarth, 21 – Harray Stenness, 22 – St Ola, 23 – Grimness, 24 – Wind Wick. Fig. S2 Map showing Orkney localities where ancient specimens of M. arvalis successfully provided either radiocarbon dates and/or cytb sequences. Localities as listed in Tables  and S3: 1 ‐ Holm of Papa Westray, Westray, 2 ‐ Point of Cott, Westray, 3 ‐ Pierowall Quarry, Westray, 4 ‐ Quanterness, Mainland, 5 ‐ Earl's Bu, Mainland, 6 ‐ Howe, Mainland, 7 ‐ Skara Brae, Mainland, 8 ‐ Green Hill, South Walls, Hoy. Fig. S3 Diagram illustrating the ABC model parameters.</note>
<note>Swiss National Science Foundation - No. 31003A‐127377; No. 3100A0‐112072; No. 3100‐126074;</note>
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<note>Arts and Humanities Research Council - No. 119396;</note>
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<street>F‐75231 Paris Cedex 05</street>
<city>Paris</city>
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<city>Bern</city>
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<street>South Parks Road</street>
<city>Oxford</city>
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<country>UK</country>
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<postCode>TW20 0EX</postCode>
<country>UK</country>
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<orgName>University of Sydney</orgName>
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<postCode>2006</postCode>
<country>Australia</country>
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<orgDiv>UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro)</orgDiv>
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<address>
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<street>CS 30016</street>
<city>F‐34988 Montferrier‐sur‐Lez cedex</city>
<country>France</country>
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<orgDiv>Centre for Anatomical and Human Sciences</orgDiv>
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<address>
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<city>York</city>
<postCode>YO10 5DD</postCode>
<country>UK</country>
</address>
</affiliation>
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<orgDiv>Department of Ecology and Evolutionary Biology</orgDiv>
<orgName>Corson Hall</orgName>
<orgName>Cornell University</orgName>
<address>
<city>Ithaca</city>
<countryPart>NY</countryPart>
<postCode>14853‐2701</postCode>
<country>USA</country>
</address>
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</affiliationGroup>
<keywordGroup type="author">
<keyword xml:id="mec12462-kwd-0001">demographic analysis</keyword>
<keyword xml:id="mec12462-kwd-0002">genetic replacement</keyword>
<keyword xml:id="mec12462-kwd-0003">island colonization</keyword>
<keyword xml:id="mec12462-kwd-0004">
<i>
<fc>M</fc>
icrotus arvalis</i>
</keyword>
<keyword xml:id="mec12462-kwd-0005">phylogeography</keyword>
</keywordGroup>
<fundingInfo>
<fundingAgency>Swiss National Science Foundation</fundingAgency>
<fundingNumber>31003A‐127377</fundingNumber>
<fundingNumber>3100A0‐112072</fundingNumber>
<fundingNumber>3100‐126074</fundingNumber>
</fundingInfo>
<fundingInfo>
<fundingAgency>Synthesis of Systematic Resources</fundingAgency>
<fundingNumber>226506‐CP‐CSA‐Infra</fundingNumber>
</fundingInfo>
<fundingInfo>
<fundingAgency>Wellcome Trust University</fundingAgency>
<fundingNumber>GR071037</fundingNumber>
</fundingInfo>
<fundingInfo>
<fundingAgency>Arts and Humanities Research Council</fundingAgency>
<fundingNumber>119396</fundingNumber>
</fundingInfo>
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<caption>
<p>
<b>Data S1</b>
Material and Methods.</p>
<p>
<b>Table S1</b>
List of modern
<i>M. arvalis</i>
samples used for morphometrics, including country and site of origin.</p>
<p>
<b>Table S2</b>
List of all modern
<i>M. arvalis</i>
used for
<i>cytb</i>
analysis and collected for this study, including country and site of origin; arranged according to
<i>cytb</i>
haplotype.</p>
<p>
<b>Table S3</b>
List of all ancient specimens of
<i>M. arvalis</i>
that successfully provided a
<i>cytb</i>
sequence with details of location collected, calibrated age range (where obtained) and GenBank Accession Number for the sequence.</p>
<p>
<b>Table S4</b>
List of primers used for the amplification of cytochrome
<i>b</i>
from
<i>M. arvalis</i>
.</p>
<p>
<b>Table S5</b>
Prior distributions of the ABC model parameters (as illustrated in Fig. S3).</p>
<p>
<b>Table S6</b>
Pairwise
<i>F</i>
<sub>ST</sub>
values between population samples analysed with microsatellites (see Table 
<link href="#mec12462-tbl-0001"></link>
and Fig. S1).</p>
<p>
<b>Fig. S1</b>
Map showing distribution of population samples of modern
<i>M. arvalis</i>
used for microsatellite typing, labelled for mtDNA lineage. Population names as listed in Table 
<link href="#mec12462-tbl-0001"></link>
: 1 – Heerenveen, 2 – Dinteloord, 3 – Stalhille, 4 – Veurne, 5 – Pihen lès Guînes, 6 – Fressenneville, 7 – Daubeuf, 8 – Thaon, 9 – Ste Marie du Mont, 10 – St Jean du Thomas, 11 – Baie d'Aiguillon, 12 – Aiffres, 13 – Avallon, 14 – Clérmont‐Ferrand, 15 – Alflen, 16 – Schiltach, 17 – Loch of Swartmill, 18 – Ness, 19 – Whitemill Bay, 20 – Settiscarth, 21 – Harray Stenness, 22 – St Ola, 23 – Grimness, 24 – Wind Wick.</p>
<p>
<b>Fig. S2</b>
Map showing Orkney localities where ancient specimens of
<i>M. arvalis</i>
successfully provided either radiocarbon dates and/or
<i>cytb</i>
sequences. Localities as listed in Tables 
<link href="#mec12462-tbl-0002"></link>
and S3: 1 ‐ Holm of Papa Westray, Westray, 2 ‐ Point of Cott, Westray, 3 ‐ Pierowall Quarry, Westray, 4 ‐ Quanterness, Mainland, 5 ‐ Earl's Bu, Mainland, 6 ‐ Howe, Mainland, 7 ‐ Skara Brae, Mainland, 8 ‐ Green Hill, South Walls, Hoy.</p>
<p>
<b>Fig. S3</b>
Diagram illustrating the ABC model parameters.</p>
</caption>
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<abstract type="main" xml:id="mec12462-abs-0001">
<title type="main">Abstract</title>
<p>Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome
<i>b</i>
(
<i>cytb</i>
) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the
<fc>O</fc>
rkney archipelago by the common vole (
<i>
<fc>M</fc>
icrotus arvalis</i>
), a species found in continental
<fc>E</fc>
urope but not in
<fc>B</fc>
ritain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 
<sc>bp</sc>
(confirmed by radiocarbon dating). We used approximate
<fc>B</fc>
ayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread
<i>cytb</i>
replacement event in the continental source area purged
<i>cytb</i>
variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental
<i>
<fc>M</fc>
. arvalis</i>
was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.</p>
</abstract>
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<note xml:id="mec12462-note-0001">These authors contributed equally. </note>
<note xml:id="mec12462-note-0002">These authors contributed equally. </note>
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<title>Divergent evolutionary processes associated with colonization of offshore islands</title>
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<title>Divergent evolutionary processes associated with colonization of offshore islands</title>
</titleInfo>
<name type="personal">
<namePart type="given">Natália</namePart>
<namePart type="family">Martínková</namePart>
<affiliation>Department of Biology, University of York, Wentworth Way, YO10 5DD, York, UK</affiliation>
<affiliation>Institute of Vertebrate Biology, Academy of Sciences of the Czech Republic, Květná 8, 603 65, Brno, Czech Republic</affiliation>
<description>These authors contributed equally.</description>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Ross</namePart>
<namePart type="family">Barnett</namePart>
<affiliation>Department of Biology, University of York, Wentworth Way, YO10 5DD, York, UK</affiliation>
<affiliation>School of Biological and Biomedical Sciences, Durham University, South Road, DH1 3LE, Durham, UK</affiliation>
<affiliation>Department of Archaeology, Durham University, South Road, DH1 3LE, Durham, UK</affiliation>
<description>These authors contributed equally.</description>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Thomas</namePart>
<namePart type="family">Cucchi</namePart>
<affiliation>Department of Archaeology, Durham University, South Road, DH1 3LE, Durham, UK</affiliation>
<affiliation>Department of Archaeology, University of Aberdeen, St Mary's, Elphinstone Road, AB24 3UF, Aberdeen, UK</affiliation>
<affiliation>Muséum national d'histoire naturelle, case postale 56 (bâtiment d'anatomie comparée), 55 rue BuffonF‐75231 Paris Cedex 05, Paris, France</affiliation>
<description>These authors contributed equally.</description>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Rahel</namePart>
<namePart type="family">Struchen</namePart>
<affiliation>Computational and Molecular Population Genetics (CMPG), Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH‐3012, Bern, Switzerland</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Marine</namePart>
<namePart type="family">Pascal</namePart>
<affiliation>Équipe Écologie des Invasions Biologiques, INRA, Campus de Beaulieu 35 000, Rennes Cedex, France</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Michel</namePart>
<namePart type="family">Pascal</namePart>
<affiliation>Équipe Écologie des Invasions Biologiques, INRA, Campus de Beaulieu 35 000, Rennes Cedex, France</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Martin C.</namePart>
<namePart type="family">Fischer</namePart>
<affiliation>Computational and Molecular Population Genetics (CMPG), Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH‐3012, Bern, Switzerland</affiliation>
<affiliation>Swiss Institute of Bioinformatics, Genopode, CH‐1015, Lausanne, Switzerland</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Thomas</namePart>
<namePart type="family">Higham</namePart>
<affiliation>Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art (RLAHA), Dyson Perrins BuildingSouth Parks Road, OX1 3QY, Oxford, UK</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Selina</namePart>
<namePart type="family">Brace</namePart>
<affiliation>School of Biological Sciences, Royal Holloway, University of London, TW20 0EX, Egham, UK</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Simon Y. W.</namePart>
<namePart type="family">Ho</namePart>
<affiliation>School of Biological Sciences, University of Sydney, NSW, 2006, Sydney, Australia</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Jean‐Pierre</namePart>
<namePart type="family">Quéré</namePart>
<affiliation>UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), INRA, Campus international de BaillarguetCS 30016, F‐34988 Montferrier‐sur‐Lez cedex, France</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Paul</namePart>
<namePart type="family">O'Higgins</namePart>
<affiliation>Centre for Anatomical and Human Sciences, Hull York Medical School (HYMS), University of York, John Hughlings Jackson Building, YO10 5DD, York, UK</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Laurent</namePart>
<namePart type="family">Excoffier</namePart>
<affiliation>Computational and Molecular Population Genetics (CMPG), Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH‐3012, Bern, Switzerland</affiliation>
<affiliation>Swiss Institute of Bioinformatics, Genopode, CH‐1015, Lausanne, Switzerland</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Gerald</namePart>
<namePart type="family">Heckel</namePart>
<affiliation>Computational and Molecular Population Genetics (CMPG), Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH‐3012, Bern, Switzerland</affiliation>
<affiliation>Swiss Institute of Bioinformatics, Genopode, CH‐1015, Lausanne, Switzerland</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">A.</namePart>
<namePart type="family">Rus Hoelzel</namePart>
<affiliation>School of Biological and Biomedical Sciences, Durham University, South Road, DH1 3LE, Durham, UK</affiliation>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Keith M.</namePart>
<namePart type="family">Dobney</namePart>
<affiliation>Department of Archaeology, Durham University, South Road, DH1 3LE, Durham, UK</affiliation>
<affiliation>Department of Archaeology, University of Aberdeen, St Mary's, Elphinstone Road, AB24 3UF, Aberdeen, UK</affiliation>
<description>These authors contributed equally.</description>
<role>
<roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal">
<namePart type="given">Jeremy B.</namePart>
<namePart type="family">Searle</namePart>
<affiliation>Department of Biology, University of York, Wentworth Way, YO10 5DD, York, UK</affiliation>
<affiliation>Department of Ecology and Evolutionary Biology, Corson Hall, Cornell University, NY, 14853‐2701, Ithaca, USA</affiliation>
<description>These authors contributed equally.</description>
<description>Correspondence: Jeremy B. Searle, Fax: +1‐607‐255‐8088; E‐mail: </description>
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<abstract>Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic ‘ark’. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land‐use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.</abstract>
<note type="additional physical form">Data S1 Material and Methods. Table S1 List of modern M. arvalis samples used for morphometrics, including country and site of origin. Table S2 List of all modern M. arvalis used for cytb analysis and collected for this study, including country and site of origin; arranged according to cytb haplotype. Table S3 List of all ancient specimens of M. arvalis that successfully provided a cytb sequence with details of location collected, calibrated age range (where obtained) and GenBank Accession Number for the sequence. Table S4 List of primers used for the amplification of cytochrome b from M. arvalis. Table S5 Prior distributions of the ABC model parameters (as illustrated in Fig. S3). Table S6 Pairwise FST values between population samples analysed with microsatellites (see Table  and Fig. S1). Fig. S1 Map showing distribution of population samples of modern M. arvalis used for microsatellite typing, labelled for mtDNA lineage. Population names as listed in Table : 1 – Heerenveen, 2 – Dinteloord, 3 – Stalhille, 4 – Veurne, 5 – Pihen lès Guînes, 6 – Fressenneville, 7 – Daubeuf, 8 – Thaon, 9 – Ste Marie du Mont, 10 – St Jean du Thomas, 11 – Baie d'Aiguillon, 12 – Aiffres, 13 – Avallon, 14 – Clérmont‐Ferrand, 15 – Alflen, 16 – Schiltach, 17 – Loch of Swartmill, 18 – Ness, 19 – Whitemill Bay, 20 – Settiscarth, 21 – Harray Stenness, 22 – St Ola, 23 – Grimness, 24 – Wind Wick. Fig. S2 Map showing Orkney localities where ancient specimens of M. arvalis successfully provided either radiocarbon dates and/or cytb sequences. Localities as listed in Tables  and S3: 1 ‐ Holm of Papa Westray, Westray, 2 ‐ Point of Cott, Westray, 3 ‐ Pierowall Quarry, Westray, 4 ‐ Quanterness, Mainland, 5 ‐ Earl's Bu, Mainland, 6 ‐ Howe, Mainland, 7 ‐ Skara Brae, Mainland, 8 ‐ Green Hill, South Walls, Hoy. Fig. S3 Diagram illustrating the ABC model parameters.</note>
<note type="funding">Swiss National Science Foundation - No. 31003A‐127377; No. 3100A0‐112072; No. 3100‐126074; </note>
<note type="funding">Synthesis of Systematic Resources - No. 226506‐CP‐CSA‐Infra; </note>
<note type="funding">Wellcome Trust University - No. GR071037; </note>
<note type="funding">Arts and Humanities Research Council - No. 119396; </note>
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<genre>keywords</genre>
<topic>demographic analysis</topic>
<topic>genetic replacement</topic>
<topic>island colonization</topic>
<topic>Microtus arvalis</topic>
<topic>phylogeography</topic>
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<topic>Original Article</topic>
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<identifier type="ISSN">0962-1083</identifier>
<identifier type="eISSN">1365-294X</identifier>
<identifier type="DOI">10.1111/(ISSN)1365-294X</identifier>
<identifier type="PublisherID">MEC</identifier>
<part>
<date>2013</date>
<detail type="volume">
<caption>vol.</caption>
<number>22</number>
</detail>
<detail type="issue">
<caption>no.</caption>
<number>20</number>
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<start>5205</start>
<end>5220</end>
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<identifier type="DOI">10.1111/mec.12462</identifier>
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<accessCondition type="use and reproduction" contentType="creativeCommonsBy">This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.</accessCondition>
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