Table 1

List of amphi‐oceanic boreal–arctic taxa included in the study, and the distributional and phylogeographic patterns identified. Only clades where the common ancestor is considered to post‐date the opening of the Bering Strait are included. When more than one nominal species is involved in a basin, the taxon that is ‘original’ in the given ocean basin is indicated. A refers to an arctic–boreal taxon, whose distribution is not necessarily discontinuous through the Arctic basin. In each taxon (genus), the different species and deep phylogeographic lineages (plausibly >150 ky old) are identified by the numerals 1, 2, …, and their presence in five biogeographical regions is indicated in the middle panel. Brackets mark assumed human‐mediated invasion. Lineages of putative late‐ and post‐glacial age (<130 ky) are referred to with the same numeral. Additionally, an inferred trans‐arctic contact of such lineages is indicated with a + (e.g. 1+ indicates a post‐glacial reinvasion of a late‐glacial sister lineage to the region indicated). Intra‐oceanic late‐glacial phenomena are not considered. Data references for previously published data are listed in Table S2

Amphi‐boreal and boreal–arctic taxaDistribution of lineagesPatternData references
Pacific OceanAtlantic OceanNWPNEPBering/ Chukchi–BeaufortNWANEA
Molluscs: Bivalves
Hiatella sp. LAHiatella sp. KA1111+2b c dLaakkonen, Strelkov, and Väinölä (2015)
Limecola balthica/ L. petalumL. balthica/ L. petalum11b(4)12 41 3b c dNikula et al. (2007)
Macoma calcarea  AM. calcarea  A1111b1dThis study, Layton et al. (2014)
Modiolus modiolusM. modiolus1 2233aHalanych et al., 2013, this study
Mytilus trossulusM. edulis111 21 2b c dMarko et al. (2010), this study
Turtonia minutaT. minuta12?3aThis study
Molluscs: Gastropods
Acanthodoris atrogriseata/pilosaA. pilosa?1 222b c dBOLD
Aeolidia papillosaA. papillosa11 211 3b cCarmona et al., (2013), this study
Alderia modestaA. modesta11?2aEllingson and Krug (2006)
Ariadnaria insignisA. borealis?1?2aBOLD
Dendronotus dalliD. niveus1122aEkimova et al. (2015), Leray, Paulay & Mulcahy (2019), Valdés et al. (2017)
Dendronotus kamchaticusD. lacteus11?2aEkimova et al. (2015), Ekimova et al. (2016), Korshunova et al. (2016), Pola & Gosliner (2010), Korshunova et al. (2017)
Dendronotus venustusD. frondosus1 22??dDewaard (2017), Korshunova et al. (2019), Pola & Gosliner (2010), Ekimova et al. (2016)
Doris montereyensisD. pseudoargus?1?2aBOLD
Flabellina trophina/fuscaF. gracilis11?2aShields (2009), Korshunova et al. (2017), BOLD
Lacuna vinctaL. vincta122ba1Leray & Paulay GenBank MH242817, Layton, Martel & Hebert (2014), Castelin et al. (2016), BOLD
Limacina helicinaL. helicina11?11dAbyzova et al. (2018), Chichvarkhin (2016), Shimizu et al. (2017), Hunt et al. (2010), Layton, Martel & Hebert (2014), Jennings et al. (2010)
Littorina aleuticaL. obtusata12aReid, (1990), Wares and Cunningham (2001), Reid et al. (2012)
Littorina squalidaL. littorea11122aReid, (1990), Giribet et al. (2006), Reid et al. (2012), Layton et al. (2014)
Nucella heysianaN. lapillus122aCollins et al. (1996), Wares and Cunningham (2001), Marko et al. (2014)
Onchidoris muricataO. muricata1122a1Hallas & Gosliner (2015), Martynov & Korshunova (2017), Chichvarkhin et al. (2018), BOLD
Velutina velutinaV. velutina?111dBOLD
Crustaceans
Balanus crenatusB. crenatus1 21 222+2b c dThis study
Calliopius laeviusculusC. laeviusculus1?23aThis study, Radulovici et al. (2009)
Centropages abdominalisC. abdominalis111dYoung (2016), Sun, Wang & Liu (2008), BOLD
Eusergestes similisE. articus?122aBOLD
Evadne nordmanniE. nordmanni211aDurbin et al. (2008)
Gammarus setosus  AG. setosus  A1 21?2+2+b c dThis study
Neomysis awatschensisN. americana/integer1123aThis study, Radulovici et al. (2009), Remerie et al. (2009)
Pandalus borealisP. borealis111+11eThis study, Steinke: BOLD, Radulovici et al (2009)
Pleopsis polyphaemoidesP. polyphaemoides2 (1)1a ?Durbin et al. (2008)
Podon leuckartiiP. leuckartii2 (1)?11a1Durbin et al. (2008)
Pseudocalanus moultoniP. moultoni122a1Aarbakke et al. (2011), Laakmann et al. (2013), Young (2016), BOLD a
Semibalanus balanoidesS. balanoides1122aThis study, Wares and Cunningham (2001), Wares et al. (2009)
Tortanus discaudatusT. discaudatus12aYoung (2016), BOLD a
Echinodems
Asterias amurensisA. rubens, A. forbesi12 (3)3aWares (2001), Matsubara et al. (2005)
Crossaster papposusC. papposus12a1Corstorphine (2010)
Echinarachnius parmaE. parma111+dThis study
Henricia cf. oculataH. cf. oculata11dCorstorphine (2010)
Henricia sp. EAC06Henricia sp. EAC0611dCorstorphine (2010)
Henricia sp. AAB9183Henricia sp. AAD3482?11 2b c dLayton, Corstorphine & Hebert (2016)
Pteraster militarisP. militaris12a1Corstorphine (2010)
Solaster endecaS. endeca12a1Corstorphine (2010)
Strongylocentrotus droebachiensisS. droebachiensis11+1+1b c dAddison and Hart (2005), this study
Strongylocentrotus pallidusS. pallidus11111dCorstophine (2010), Hardy et al. (2011)
Polychaetes
Alitta virensA. virens12?aCarr, Hardy, Brown, Macdonald, and Hebert (2011)
Glycera capitata  AG. capitata  A1 Ber11dCarr et al. (2011)
Harmothoe imbricata groupAH. imbricata groupA71 41 51 2 3 52 3b c dCarr et al. (2011), Hardy et al. (2011)
Harmothoe rarispina groupAH. rarispina group A1 211?b c dCarr et al. (2011), BOLD
Hediste atokaH. diversicolor12 3aAudzijonyte et al. (2008), Tosuji & Sato (2010)
Lepidonotus squamatus CMC02L. squamatus CMC0112aCarr et al. (2011), Hardy et al. (2011)
Myxicola infundibulum CMC02M. infundibulum CMC0112aCarr et al. (2011)
Nereis pelagica CMC01, CMC03AN. pelagica CMC02, CMC03A132 32b c dCarr et al. (2011), Hardy et al. (2011), BOLD
Nothria conchylega CMC01AN. conchylega CMC0212aCarr et al. (2011)
Pectinaria granulata CMC01P. granulata CMC02 A12aCarr et al. (2011)
Pholoe baltica  AP. baltica  A1 Ber1?dCarr et al. (2011)
Pholoe minuta  AP. minuta  A1 Ber1dCarr et al. (2011)
Phyllodoce sp. CMC01AP. sp. CMC01A1 Ber1dCarr et al. (2011)
Priapulida
Priapulus cf. caudatusP. caudatus sp. 112332 3 4b dThis study
Fishes
Anisarchus medius  AA. medius  A??11?eMecklenburg et al. (2011)
Aspidophoroides monopterygiusA. olrikii1b 2b1b 2b1 21 22bb c dMecklenburg et al. (2011)
Clupea pallasiiC. harengus11121 2b c dLaakkonen et al. (2013)
Eleginus nawagaE. gracilis11b12aThis study
Eumesogrammus praecisusAE. praecisusA1a1a1+1+eMecklenburg et al. (2011)
Gadus chalcogrammus (finnmarchicus)G. morhua11121 2b c dUrsvik et al. (2007), Gorbachev et al. (2014)
Gadus macrocephalus (ogac)G. morhua1111 22b c dCoulson et al. (2006)
Gasterosteus aculeatusG. aculeatus11 21 23+3+a1Ortí et al. (1994), Mäkinen & Merilä (2008), Lescak et al. (2015), this study
Glyptocephalus stelleriG. cynoglossus123aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016), Noren et al. (2009)
Gymnocanthus spp.G. tricuspis123451a 2a1234564a4ab c dYamazaki et al. (2013)
Hippoglossoides dubius / robustusH. platessoides122aVinnikov et al. (2018), Noren et al. (2009)
Hippoglossoides robustusH. platessoides11122+aBOLD
Hippoglossus stenolepisH. hippoglossus111b22aSanjuan & Comesaña (2002), Espineira et al. (2008), Mjelle et al. (2008), Mecklenburg et al. (2011)
Leptagonus decagonus  AL. decagonus  A1a1+11eMcCusker et al. (2012), BOLD
Leptoclinus maculatusAL. maculatusA11a1+11eMecklenburg et al. (2011)
Limanda asperaL. limanda1112aThis study
Liopsetta pinnifasciata  AL. glacialis  A1 + 2?1 2b c dThis study, Kartavtsev et al. (2014)
Lumpenus fabriciiL. lampretaeformis??1 Ber?/22 1b cThis study, Mecklenburg et al. (2011)
Mallotus villosus  AM. villosus  g  A1211 34b c dDodson et al. (2007)
Megalocottus platycephalusMyoxocephalus quadricornis11a1 2a22aKontula & Väinölä (2003), Podlesnykh & Moreva (2014)
Myoxocephalus stelleri + othersM. scorpius  A1?1 2a1 22+2+b c dThis study, Mecklenburg et al. (2011)
Osmerus dentexO. dentex11a11dSkurikhina et al. (2013)
Pholis laetaP. gunnellus11122aSteinke et al. (2009)
Platichthys stellatusP. flesus1112aThis study
Pleuronectes quadrituberculatusP. platessa??12aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016)
Pungitius pungitiusP. pungitius1222+2a1Shikano et al. (2010), Mecklenburg et al. (2011)
Reinhardtius hippoglossoidesAR.. hippoglossoides  A1b1b1 Ber‐Chu11dVis et al. (1997), Mecklenburg et al. (2011), Kartavtsev et al. (2014)
Zoarces fedoroviZ. viviparus12a1This study
Mammals (seals)
Pusa hispida ochotensis  AP. h. hispida  A/ladogensis1111dThis study
Phoca vitulinaP. largha121a212a2b cNakagawa et al. (2010), Stanley et. al (1996)
Amphi‐boreal and boreal–arctic taxaDistribution of lineagesPatternData references
Pacific OceanAtlantic OceanNWPNEPBering/ Chukchi–BeaufortNWANEA
Molluscs: Bivalves
Hiatella sp. LAHiatella sp. KA1111+2b c dLaakkonen, Strelkov, and Väinölä (2015)
Limecola balthica/ L. petalumL. balthica/ L. petalum11b(4)12 41 3b c dNikula et al. (2007)
Macoma calcarea  AM. calcarea  A1111b1dThis study, Layton et al. (2014)
Modiolus modiolusM. modiolus1 2233aHalanych et al., 2013, this study
Mytilus trossulusM. edulis111 21 2b c dMarko et al. (2010), this study
Turtonia minutaT. minuta12?3aThis study
Molluscs: Gastropods
Acanthodoris atrogriseata/pilosaA. pilosa?1 222b c dBOLD
Aeolidia papillosaA. papillosa11 211 3b cCarmona et al., (2013), this study
Alderia modestaA. modesta11?2aEllingson and Krug (2006)
Ariadnaria insignisA. borealis?1?2aBOLD
Dendronotus dalliD. niveus1122aEkimova et al. (2015), Leray, Paulay & Mulcahy (2019), Valdés et al. (2017)
Dendronotus kamchaticusD. lacteus11?2aEkimova et al. (2015), Ekimova et al. (2016), Korshunova et al. (2016), Pola & Gosliner (2010), Korshunova et al. (2017)
Dendronotus venustusD. frondosus1 22??dDewaard (2017), Korshunova et al. (2019), Pola & Gosliner (2010), Ekimova et al. (2016)
Doris montereyensisD. pseudoargus?1?2aBOLD
Flabellina trophina/fuscaF. gracilis11?2aShields (2009), Korshunova et al. (2017), BOLD
Lacuna vinctaL. vincta122ba1Leray & Paulay GenBank MH242817, Layton, Martel & Hebert (2014), Castelin et al. (2016), BOLD
Limacina helicinaL. helicina11?11dAbyzova et al. (2018), Chichvarkhin (2016), Shimizu et al. (2017), Hunt et al. (2010), Layton, Martel & Hebert (2014), Jennings et al. (2010)
Littorina aleuticaL. obtusata12aReid, (1990), Wares and Cunningham (2001), Reid et al. (2012)
Littorina squalidaL. littorea11122aReid, (1990), Giribet et al. (2006), Reid et al. (2012), Layton et al. (2014)
Nucella heysianaN. lapillus122aCollins et al. (1996), Wares and Cunningham (2001), Marko et al. (2014)
Onchidoris muricataO. muricata1122a1Hallas & Gosliner (2015), Martynov & Korshunova (2017), Chichvarkhin et al. (2018), BOLD
Velutina velutinaV. velutina?111dBOLD
Crustaceans
Balanus crenatusB. crenatus1 21 222+2b c dThis study
Calliopius laeviusculusC. laeviusculus1?23aThis study, Radulovici et al. (2009)
Centropages abdominalisC. abdominalis111dYoung (2016), Sun, Wang & Liu (2008), BOLD
Eusergestes similisE. articus?122aBOLD
Evadne nordmanniE. nordmanni211aDurbin et al. (2008)
Gammarus setosus  AG. setosus  A1 21?2+2+b c dThis study
Neomysis awatschensisN. americana/integer1123aThis study, Radulovici et al. (2009), Remerie et al. (2009)
Pandalus borealisP. borealis111+11eThis study, Steinke: BOLD, Radulovici et al (2009)
Pleopsis polyphaemoidesP. polyphaemoides2 (1)1a ?Durbin et al. (2008)
Podon leuckartiiP. leuckartii2 (1)?11a1Durbin et al. (2008)
Pseudocalanus moultoniP. moultoni122a1Aarbakke et al. (2011), Laakmann et al. (2013), Young (2016), BOLD a
Semibalanus balanoidesS. balanoides1122aThis study, Wares and Cunningham (2001), Wares et al. (2009)
Tortanus discaudatusT. discaudatus12aYoung (2016), BOLD a
Echinodems
Asterias amurensisA. rubens, A. forbesi12 (3)3aWares (2001), Matsubara et al. (2005)
Crossaster papposusC. papposus12a1Corstorphine (2010)
Echinarachnius parmaE. parma111+dThis study
Henricia cf. oculataH. cf. oculata11dCorstorphine (2010)
Henricia sp. EAC06Henricia sp. EAC0611dCorstorphine (2010)
Henricia sp. AAB9183Henricia sp. AAD3482?11 2b c dLayton, Corstorphine & Hebert (2016)
Pteraster militarisP. militaris12a1Corstorphine (2010)
Solaster endecaS. endeca12a1Corstorphine (2010)
Strongylocentrotus droebachiensisS. droebachiensis11+1+1b c dAddison and Hart (2005), this study
Strongylocentrotus pallidusS. pallidus11111dCorstophine (2010), Hardy et al. (2011)
Polychaetes
Alitta virensA. virens12?aCarr, Hardy, Brown, Macdonald, and Hebert (2011)
Glycera capitata  AG. capitata  A1 Ber11dCarr et al. (2011)
Harmothoe imbricata groupAH. imbricata groupA71 41 51 2 3 52 3b c dCarr et al. (2011), Hardy et al. (2011)
Harmothoe rarispina groupAH. rarispina group A1 211?b c dCarr et al. (2011), BOLD
Hediste atokaH. diversicolor12 3aAudzijonyte et al. (2008), Tosuji & Sato (2010)
Lepidonotus squamatus CMC02L. squamatus CMC0112aCarr et al. (2011), Hardy et al. (2011)
Myxicola infundibulum CMC02M. infundibulum CMC0112aCarr et al. (2011)
Nereis pelagica CMC01, CMC03AN. pelagica CMC02, CMC03A132 32b c dCarr et al. (2011), Hardy et al. (2011), BOLD
Nothria conchylega CMC01AN. conchylega CMC0212aCarr et al. (2011)
Pectinaria granulata CMC01P. granulata CMC02 A12aCarr et al. (2011)
Pholoe baltica  AP. baltica  A1 Ber1?dCarr et al. (2011)
Pholoe minuta  AP. minuta  A1 Ber1dCarr et al. (2011)
Phyllodoce sp. CMC01AP. sp. CMC01A1 Ber1dCarr et al. (2011)
Priapulida
Priapulus cf. caudatusP. caudatus sp. 112332 3 4b dThis study
Fishes
Anisarchus medius  AA. medius  A??11?eMecklenburg et al. (2011)
Aspidophoroides monopterygiusA. olrikii1b 2b1b 2b1 21 22bb c dMecklenburg et al. (2011)
Clupea pallasiiC. harengus11121 2b c dLaakkonen et al. (2013)
Eleginus nawagaE. gracilis11b12aThis study
Eumesogrammus praecisusAE. praecisusA1a1a1+1+eMecklenburg et al. (2011)
Gadus chalcogrammus (finnmarchicus)G. morhua11121 2b c dUrsvik et al. (2007), Gorbachev et al. (2014)
Gadus macrocephalus (ogac)G. morhua1111 22b c dCoulson et al. (2006)
Gasterosteus aculeatusG. aculeatus11 21 23+3+a1Ortí et al. (1994), Mäkinen & Merilä (2008), Lescak et al. (2015), this study
Glyptocephalus stelleriG. cynoglossus123aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016), Noren et al. (2009)
Gymnocanthus spp.G. tricuspis123451a 2a1234564a4ab c dYamazaki et al. (2013)
Hippoglossoides dubius / robustusH. platessoides122aVinnikov et al. (2018), Noren et al. (2009)
Hippoglossoides robustusH. platessoides11122+aBOLD
Hippoglossus stenolepisH. hippoglossus111b22aSanjuan & Comesaña (2002), Espineira et al. (2008), Mjelle et al. (2008), Mecklenburg et al. (2011)
Leptagonus decagonus  AL. decagonus  A1a1+11eMcCusker et al. (2012), BOLD
Leptoclinus maculatusAL. maculatusA11a1+11eMecklenburg et al. (2011)
Limanda asperaL. limanda1112aThis study
Liopsetta pinnifasciata  AL. glacialis  A1 + 2?1 2b c dThis study, Kartavtsev et al. (2014)
Lumpenus fabriciiL. lampretaeformis??1 Ber?/22 1b cThis study, Mecklenburg et al. (2011)
Mallotus villosus  AM. villosus  g  A1211 34b c dDodson et al. (2007)
Megalocottus platycephalusMyoxocephalus quadricornis11a1 2a22aKontula & Väinölä (2003), Podlesnykh & Moreva (2014)
Myoxocephalus stelleri + othersM. scorpius  A1?1 2a1 22+2+b c dThis study, Mecklenburg et al. (2011)
Osmerus dentexO. dentex11a11dSkurikhina et al. (2013)
Pholis laetaP. gunnellus11122aSteinke et al. (2009)
Platichthys stellatusP. flesus1112aThis study
Pleuronectes quadrituberculatusP. platessa??12aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016)
Pungitius pungitiusP. pungitius1222+2a1Shikano et al. (2010), Mecklenburg et al. (2011)
Reinhardtius hippoglossoidesAR.. hippoglossoides  A1b1b1 Ber‐Chu11dVis et al. (1997), Mecklenburg et al. (2011), Kartavtsev et al. (2014)
Zoarces fedoroviZ. viviparus12a1This study
Mammals (seals)
Pusa hispida ochotensis  AP. h. hispida  A/ladogensis1111dThis study
Phoca vitulinaP. largha121a212a2b cNakagawa et al. (2010), Stanley et. al (1996)

NEP, Northeast Pacific; NWP, Northwest Pacific; NEA, Northeast Atlantic; NWA, Northwest Atlantic.

Samples from the Bering Strait region ‘Ber’ are included in the Chukchi column:

Classification of inferred biogeographical patterns (Figure 2):

a) long vicariant histories (descending from the Pliocene–Early Pleistocene)

a1) more recent vicariant histories (descending from the Middle Pleistocene, 1–0.2 Myr).

b) minimum of two distinct trans‐Arctic divergence events.

c) possible secondary contact following repeated trans‐Arctic dispersal.

d) incomplete lineage sorting or close relationship, interpreted as post‐ or late‐glacial dispersal.

e) hypothesized post‐glacial Arctic contact following short vicariance (e.g. late‐glacial).

?—Presence of the taxon, but without genetic assignment.

a

Absence of genetic data (but presence of the morphologically identified species).

b

Inference based on unpublished data or on indirect evidence (not critical for the conclusions).

Table 1

List of amphi‐oceanic boreal–arctic taxa included in the study, and the distributional and phylogeographic patterns identified. Only clades where the common ancestor is considered to post‐date the opening of the Bering Strait are included. When more than one nominal species is involved in a basin, the taxon that is ‘original’ in the given ocean basin is indicated. A refers to an arctic–boreal taxon, whose distribution is not necessarily discontinuous through the Arctic basin. In each taxon (genus), the different species and deep phylogeographic lineages (plausibly >150 ky old) are identified by the numerals 1, 2, …, and their presence in five biogeographical regions is indicated in the middle panel. Brackets mark assumed human‐mediated invasion. Lineages of putative late‐ and post‐glacial age (<130 ky) are referred to with the same numeral. Additionally, an inferred trans‐arctic contact of such lineages is indicated with a + (e.g. 1+ indicates a post‐glacial reinvasion of a late‐glacial sister lineage to the region indicated). Intra‐oceanic late‐glacial phenomena are not considered. Data references for previously published data are listed in Table S2

Amphi‐boreal and boreal–arctic taxaDistribution of lineagesPatternData references
Pacific OceanAtlantic OceanNWPNEPBering/ Chukchi–BeaufortNWANEA
Molluscs: Bivalves
Hiatella sp. LAHiatella sp. KA1111+2b c dLaakkonen, Strelkov, and Väinölä (2015)
Limecola balthica/ L. petalumL. balthica/ L. petalum11b(4)12 41 3b c dNikula et al. (2007)
Macoma calcarea  AM. calcarea  A1111b1dThis study, Layton et al. (2014)
Modiolus modiolusM. modiolus1 2233aHalanych et al., 2013, this study
Mytilus trossulusM. edulis111 21 2b c dMarko et al. (2010), this study
Turtonia minutaT. minuta12?3aThis study
Molluscs: Gastropods
Acanthodoris atrogriseata/pilosaA. pilosa?1 222b c dBOLD
Aeolidia papillosaA. papillosa11 211 3b cCarmona et al., (2013), this study
Alderia modestaA. modesta11?2aEllingson and Krug (2006)
Ariadnaria insignisA. borealis?1?2aBOLD
Dendronotus dalliD. niveus1122aEkimova et al. (2015), Leray, Paulay & Mulcahy (2019), Valdés et al. (2017)
Dendronotus kamchaticusD. lacteus11?2aEkimova et al. (2015), Ekimova et al. (2016), Korshunova et al. (2016), Pola & Gosliner (2010), Korshunova et al. (2017)
Dendronotus venustusD. frondosus1 22??dDewaard (2017), Korshunova et al. (2019), Pola & Gosliner (2010), Ekimova et al. (2016)
Doris montereyensisD. pseudoargus?1?2aBOLD
Flabellina trophina/fuscaF. gracilis11?2aShields (2009), Korshunova et al. (2017), BOLD
Lacuna vinctaL. vincta122ba1Leray & Paulay GenBank MH242817, Layton, Martel & Hebert (2014), Castelin et al. (2016), BOLD
Limacina helicinaL. helicina11?11dAbyzova et al. (2018), Chichvarkhin (2016), Shimizu et al. (2017), Hunt et al. (2010), Layton, Martel & Hebert (2014), Jennings et al. (2010)
Littorina aleuticaL. obtusata12aReid, (1990), Wares and Cunningham (2001), Reid et al. (2012)
Littorina squalidaL. littorea11122aReid, (1990), Giribet et al. (2006), Reid et al. (2012), Layton et al. (2014)
Nucella heysianaN. lapillus122aCollins et al. (1996), Wares and Cunningham (2001), Marko et al. (2014)
Onchidoris muricataO. muricata1122a1Hallas & Gosliner (2015), Martynov & Korshunova (2017), Chichvarkhin et al. (2018), BOLD
Velutina velutinaV. velutina?111dBOLD
Crustaceans
Balanus crenatusB. crenatus1 21 222+2b c dThis study
Calliopius laeviusculusC. laeviusculus1?23aThis study, Radulovici et al. (2009)
Centropages abdominalisC. abdominalis111dYoung (2016), Sun, Wang & Liu (2008), BOLD
Eusergestes similisE. articus?122aBOLD
Evadne nordmanniE. nordmanni211aDurbin et al. (2008)
Gammarus setosus  AG. setosus  A1 21?2+2+b c dThis study
Neomysis awatschensisN. americana/integer1123aThis study, Radulovici et al. (2009), Remerie et al. (2009)
Pandalus borealisP. borealis111+11eThis study, Steinke: BOLD, Radulovici et al (2009)
Pleopsis polyphaemoidesP. polyphaemoides2 (1)1a ?Durbin et al. (2008)
Podon leuckartiiP. leuckartii2 (1)?11a1Durbin et al. (2008)
Pseudocalanus moultoniP. moultoni122a1Aarbakke et al. (2011), Laakmann et al. (2013), Young (2016), BOLD a
Semibalanus balanoidesS. balanoides1122aThis study, Wares and Cunningham (2001), Wares et al. (2009)
Tortanus discaudatusT. discaudatus12aYoung (2016), BOLD a
Echinodems
Asterias amurensisA. rubens, A. forbesi12 (3)3aWares (2001), Matsubara et al. (2005)
Crossaster papposusC. papposus12a1Corstorphine (2010)
Echinarachnius parmaE. parma111+dThis study
Henricia cf. oculataH. cf. oculata11dCorstorphine (2010)
Henricia sp. EAC06Henricia sp. EAC0611dCorstorphine (2010)
Henricia sp. AAB9183Henricia sp. AAD3482?11 2b c dLayton, Corstorphine & Hebert (2016)
Pteraster militarisP. militaris12a1Corstorphine (2010)
Solaster endecaS. endeca12a1Corstorphine (2010)
Strongylocentrotus droebachiensisS. droebachiensis11+1+1b c dAddison and Hart (2005), this study
Strongylocentrotus pallidusS. pallidus11111dCorstophine (2010), Hardy et al. (2011)
Polychaetes
Alitta virensA. virens12?aCarr, Hardy, Brown, Macdonald, and Hebert (2011)
Glycera capitata  AG. capitata  A1 Ber11dCarr et al. (2011)
Harmothoe imbricata groupAH. imbricata groupA71 41 51 2 3 52 3b c dCarr et al. (2011), Hardy et al. (2011)
Harmothoe rarispina groupAH. rarispina group A1 211?b c dCarr et al. (2011), BOLD
Hediste atokaH. diversicolor12 3aAudzijonyte et al. (2008), Tosuji & Sato (2010)
Lepidonotus squamatus CMC02L. squamatus CMC0112aCarr et al. (2011), Hardy et al. (2011)
Myxicola infundibulum CMC02M. infundibulum CMC0112aCarr et al. (2011)
Nereis pelagica CMC01, CMC03AN. pelagica CMC02, CMC03A132 32b c dCarr et al. (2011), Hardy et al. (2011), BOLD
Nothria conchylega CMC01AN. conchylega CMC0212aCarr et al. (2011)
Pectinaria granulata CMC01P. granulata CMC02 A12aCarr et al. (2011)
Pholoe baltica  AP. baltica  A1 Ber1?dCarr et al. (2011)
Pholoe minuta  AP. minuta  A1 Ber1dCarr et al. (2011)
Phyllodoce sp. CMC01AP. sp. CMC01A1 Ber1dCarr et al. (2011)
Priapulida
Priapulus cf. caudatusP. caudatus sp. 112332 3 4b dThis study
Fishes
Anisarchus medius  AA. medius  A??11?eMecklenburg et al. (2011)
Aspidophoroides monopterygiusA. olrikii1b 2b1b 2b1 21 22bb c dMecklenburg et al. (2011)
Clupea pallasiiC. harengus11121 2b c dLaakkonen et al. (2013)
Eleginus nawagaE. gracilis11b12aThis study
Eumesogrammus praecisusAE. praecisusA1a1a1+1+eMecklenburg et al. (2011)
Gadus chalcogrammus (finnmarchicus)G. morhua11121 2b c dUrsvik et al. (2007), Gorbachev et al. (2014)
Gadus macrocephalus (ogac)G. morhua1111 22b c dCoulson et al. (2006)
Gasterosteus aculeatusG. aculeatus11 21 23+3+a1Ortí et al. (1994), Mäkinen & Merilä (2008), Lescak et al. (2015), this study
Glyptocephalus stelleriG. cynoglossus123aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016), Noren et al. (2009)
Gymnocanthus spp.G. tricuspis123451a 2a1234564a4ab c dYamazaki et al. (2013)
Hippoglossoides dubius / robustusH. platessoides122aVinnikov et al. (2018), Noren et al. (2009)
Hippoglossoides robustusH. platessoides11122+aBOLD
Hippoglossus stenolepisH. hippoglossus111b22aSanjuan & Comesaña (2002), Espineira et al. (2008), Mjelle et al. (2008), Mecklenburg et al. (2011)
Leptagonus decagonus  AL. decagonus  A1a1+11eMcCusker et al. (2012), BOLD
Leptoclinus maculatusAL. maculatusA11a1+11eMecklenburg et al. (2011)
Limanda asperaL. limanda1112aThis study
Liopsetta pinnifasciata  AL. glacialis  A1 + 2?1 2b c dThis study, Kartavtsev et al. (2014)
Lumpenus fabriciiL. lampretaeformis??1 Ber?/22 1b cThis study, Mecklenburg et al. (2011)
Mallotus villosus  AM. villosus  g  A1211 34b c dDodson et al. (2007)
Megalocottus platycephalusMyoxocephalus quadricornis11a1 2a22aKontula & Väinölä (2003), Podlesnykh & Moreva (2014)
Myoxocephalus stelleri + othersM. scorpius  A1?1 2a1 22+2+b c dThis study, Mecklenburg et al. (2011)
Osmerus dentexO. dentex11a11dSkurikhina et al. (2013)
Pholis laetaP. gunnellus11122aSteinke et al. (2009)
Platichthys stellatusP. flesus1112aThis study
Pleuronectes quadrituberculatusP. platessa??12aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016)
Pungitius pungitiusP. pungitius1222+2a1Shikano et al. (2010), Mecklenburg et al. (2011)
Reinhardtius hippoglossoidesAR.. hippoglossoides  A1b1b1 Ber‐Chu11dVis et al. (1997), Mecklenburg et al. (2011), Kartavtsev et al. (2014)
Zoarces fedoroviZ. viviparus12a1This study
Mammals (seals)
Pusa hispida ochotensis  AP. h. hispida  A/ladogensis1111dThis study
Phoca vitulinaP. largha121a212a2b cNakagawa et al. (2010), Stanley et. al (1996)
Amphi‐boreal and boreal–arctic taxaDistribution of lineagesPatternData references
Pacific OceanAtlantic OceanNWPNEPBering/ Chukchi–BeaufortNWANEA
Molluscs: Bivalves
Hiatella sp. LAHiatella sp. KA1111+2b c dLaakkonen, Strelkov, and Väinölä (2015)
Limecola balthica/ L. petalumL. balthica/ L. petalum11b(4)12 41 3b c dNikula et al. (2007)
Macoma calcarea  AM. calcarea  A1111b1dThis study, Layton et al. (2014)
Modiolus modiolusM. modiolus1 2233aHalanych et al., 2013, this study
Mytilus trossulusM. edulis111 21 2b c dMarko et al. (2010), this study
Turtonia minutaT. minuta12?3aThis study
Molluscs: Gastropods
Acanthodoris atrogriseata/pilosaA. pilosa?1 222b c dBOLD
Aeolidia papillosaA. papillosa11 211 3b cCarmona et al., (2013), this study
Alderia modestaA. modesta11?2aEllingson and Krug (2006)
Ariadnaria insignisA. borealis?1?2aBOLD
Dendronotus dalliD. niveus1122aEkimova et al. (2015), Leray, Paulay & Mulcahy (2019), Valdés et al. (2017)
Dendronotus kamchaticusD. lacteus11?2aEkimova et al. (2015), Ekimova et al. (2016), Korshunova et al. (2016), Pola & Gosliner (2010), Korshunova et al. (2017)
Dendronotus venustusD. frondosus1 22??dDewaard (2017), Korshunova et al. (2019), Pola & Gosliner (2010), Ekimova et al. (2016)
Doris montereyensisD. pseudoargus?1?2aBOLD
Flabellina trophina/fuscaF. gracilis11?2aShields (2009), Korshunova et al. (2017), BOLD
Lacuna vinctaL. vincta122ba1Leray & Paulay GenBank MH242817, Layton, Martel & Hebert (2014), Castelin et al. (2016), BOLD
Limacina helicinaL. helicina11?11dAbyzova et al. (2018), Chichvarkhin (2016), Shimizu et al. (2017), Hunt et al. (2010), Layton, Martel & Hebert (2014), Jennings et al. (2010)
Littorina aleuticaL. obtusata12aReid, (1990), Wares and Cunningham (2001), Reid et al. (2012)
Littorina squalidaL. littorea11122aReid, (1990), Giribet et al. (2006), Reid et al. (2012), Layton et al. (2014)
Nucella heysianaN. lapillus122aCollins et al. (1996), Wares and Cunningham (2001), Marko et al. (2014)
Onchidoris muricataO. muricata1122a1Hallas & Gosliner (2015), Martynov & Korshunova (2017), Chichvarkhin et al. (2018), BOLD
Velutina velutinaV. velutina?111dBOLD
Crustaceans
Balanus crenatusB. crenatus1 21 222+2b c dThis study
Calliopius laeviusculusC. laeviusculus1?23aThis study, Radulovici et al. (2009)
Centropages abdominalisC. abdominalis111dYoung (2016), Sun, Wang & Liu (2008), BOLD
Eusergestes similisE. articus?122aBOLD
Evadne nordmanniE. nordmanni211aDurbin et al. (2008)
Gammarus setosus  AG. setosus  A1 21?2+2+b c dThis study
Neomysis awatschensisN. americana/integer1123aThis study, Radulovici et al. (2009), Remerie et al. (2009)
Pandalus borealisP. borealis111+11eThis study, Steinke: BOLD, Radulovici et al (2009)
Pleopsis polyphaemoidesP. polyphaemoides2 (1)1a ?Durbin et al. (2008)
Podon leuckartiiP. leuckartii2 (1)?11a1Durbin et al. (2008)
Pseudocalanus moultoniP. moultoni122a1Aarbakke et al. (2011), Laakmann et al. (2013), Young (2016), BOLD a
Semibalanus balanoidesS. balanoides1122aThis study, Wares and Cunningham (2001), Wares et al. (2009)
Tortanus discaudatusT. discaudatus12aYoung (2016), BOLD a
Echinodems
Asterias amurensisA. rubens, A. forbesi12 (3)3aWares (2001), Matsubara et al. (2005)
Crossaster papposusC. papposus12a1Corstorphine (2010)
Echinarachnius parmaE. parma111+dThis study
Henricia cf. oculataH. cf. oculata11dCorstorphine (2010)
Henricia sp. EAC06Henricia sp. EAC0611dCorstorphine (2010)
Henricia sp. AAB9183Henricia sp. AAD3482?11 2b c dLayton, Corstorphine & Hebert (2016)
Pteraster militarisP. militaris12a1Corstorphine (2010)
Solaster endecaS. endeca12a1Corstorphine (2010)
Strongylocentrotus droebachiensisS. droebachiensis11+1+1b c dAddison and Hart (2005), this study
Strongylocentrotus pallidusS. pallidus11111dCorstophine (2010), Hardy et al. (2011)
Polychaetes
Alitta virensA. virens12?aCarr, Hardy, Brown, Macdonald, and Hebert (2011)
Glycera capitata  AG. capitata  A1 Ber11dCarr et al. (2011)
Harmothoe imbricata groupAH. imbricata groupA71 41 51 2 3 52 3b c dCarr et al. (2011), Hardy et al. (2011)
Harmothoe rarispina groupAH. rarispina group A1 211?b c dCarr et al. (2011), BOLD
Hediste atokaH. diversicolor12 3aAudzijonyte et al. (2008), Tosuji & Sato (2010)
Lepidonotus squamatus CMC02L. squamatus CMC0112aCarr et al. (2011), Hardy et al. (2011)
Myxicola infundibulum CMC02M. infundibulum CMC0112aCarr et al. (2011)
Nereis pelagica CMC01, CMC03AN. pelagica CMC02, CMC03A132 32b c dCarr et al. (2011), Hardy et al. (2011), BOLD
Nothria conchylega CMC01AN. conchylega CMC0212aCarr et al. (2011)
Pectinaria granulata CMC01P. granulata CMC02 A12aCarr et al. (2011)
Pholoe baltica  AP. baltica  A1 Ber1?dCarr et al. (2011)
Pholoe minuta  AP. minuta  A1 Ber1dCarr et al. (2011)
Phyllodoce sp. CMC01AP. sp. CMC01A1 Ber1dCarr et al. (2011)
Priapulida
Priapulus cf. caudatusP. caudatus sp. 112332 3 4b dThis study
Fishes
Anisarchus medius  AA. medius  A??11?eMecklenburg et al. (2011)
Aspidophoroides monopterygiusA. olrikii1b 2b1b 2b1 21 22bb c dMecklenburg et al. (2011)
Clupea pallasiiC. harengus11121 2b c dLaakkonen et al. (2013)
Eleginus nawagaE. gracilis11b12aThis study
Eumesogrammus praecisusAE. praecisusA1a1a1+1+eMecklenburg et al. (2011)
Gadus chalcogrammus (finnmarchicus)G. morhua11121 2b c dUrsvik et al. (2007), Gorbachev et al. (2014)
Gadus macrocephalus (ogac)G. morhua1111 22b c dCoulson et al. (2006)
Gasterosteus aculeatusG. aculeatus11 21 23+3+a1Ortí et al. (1994), Mäkinen & Merilä (2008), Lescak et al. (2015), this study
Glyptocephalus stelleriG. cynoglossus123aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016), Noren et al. (2009)
Gymnocanthus spp.G. tricuspis123451a 2a1234564a4ab c dYamazaki et al. (2013)
Hippoglossoides dubius / robustusH. platessoides122aVinnikov et al. (2018), Noren et al. (2009)
Hippoglossoides robustusH. platessoides11122+aBOLD
Hippoglossus stenolepisH. hippoglossus111b22aSanjuan & Comesaña (2002), Espineira et al. (2008), Mjelle et al. (2008), Mecklenburg et al. (2011)
Leptagonus decagonus  AL. decagonus  A1a1+11eMcCusker et al. (2012), BOLD
Leptoclinus maculatusAL. maculatusA11a1+11eMecklenburg et al. (2011)
Limanda asperaL. limanda1112aThis study
Liopsetta pinnifasciata  AL. glacialis  A1 + 2?1 2b c dThis study, Kartavtsev et al. (2014)
Lumpenus fabriciiL. lampretaeformis??1 Ber?/22 1b cThis study, Mecklenburg et al. (2011)
Mallotus villosus  AM. villosus  g  A1211 34b c dDodson et al. (2007)
Megalocottus platycephalusMyoxocephalus quadricornis11a1 2a22aKontula & Väinölä (2003), Podlesnykh & Moreva (2014)
Myoxocephalus stelleri + othersM. scorpius  A1?1 2a1 22+2+b c dThis study, Mecklenburg et al. (2011)
Osmerus dentexO. dentex11a11dSkurikhina et al. (2013)
Pholis laetaP. gunnellus11122aSteinke et al. (2009)
Platichthys stellatusP. flesus1112aThis study
Pleuronectes quadrituberculatusP. platessa??12aVinnikov, Thomson & Munroe (2018), Kartavtsev et al. (2016)
Pungitius pungitiusP. pungitius1222+2a1Shikano et al. (2010), Mecklenburg et al. (2011)
Reinhardtius hippoglossoidesAR.. hippoglossoides  A1b1b1 Ber‐Chu11dVis et al. (1997), Mecklenburg et al. (2011), Kartavtsev et al. (2014)
Zoarces fedoroviZ. viviparus12a1This study
Mammals (seals)
Pusa hispida ochotensis  AP. h. hispida  A/ladogensis1111dThis study
Phoca vitulinaP. largha121a212a2b cNakagawa et al. (2010), Stanley et. al (1996)

NEP, Northeast Pacific; NWP, Northwest Pacific; NEA, Northeast Atlantic; NWA, Northwest Atlantic.

Samples from the Bering Strait region ‘Ber’ are included in the Chukchi column:

Classification of inferred biogeographical patterns (Figure 2):

a) long vicariant histories (descending from the Pliocene–Early Pleistocene)

a1) more recent vicariant histories (descending from the Middle Pleistocene, 1–0.2 Myr).

b) minimum of two distinct trans‐Arctic divergence events.

c) possible secondary contact following repeated trans‐Arctic dispersal.

d) incomplete lineage sorting or close relationship, interpreted as post‐ or late‐glacial dispersal.

e) hypothesized post‐glacial Arctic contact following short vicariance (e.g. late‐glacial).

?—Presence of the taxon, but without genetic assignment.

a

Absence of genetic data (but presence of the morphologically identified species).

b

Inference based on unpublished data or on indirect evidence (not critical for the conclusions).

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