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Benton, M. J., Briggs, D. E. G., Clack, J. A., Edwards, D., Galway-Witham, J., Stringer, C. B., & Turvey, S. T. (2017). Russia–UK Collaboration in Paleontology: Past, Present, and Future. Paleontological Journal, 51(6), 576- 599. https://doi.org/10.1134/S0031030117060028 Peer reviewed version Link to published version (if available): 10.1134/S0031030117060028 Link to publication record in Explore Bristol Research PDF-document This is the accepted author manuscript (AAM). The final published version (version of record) is available online via Springer at https://doi.org/10.1134/S0031030117060028 . Please refer to any applicable terms of use of the publisher. University of Bristol - Explore Bristol Research General rights This document is made available in accordance with publisher policies. Please cite only the published version using the reference above. Full terms of use are available: http://www.bristol.ac.uk/pure/user- guides/explore-bristol-research/ebr-terms/

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Page 1: Paleontology: Past, Present, and Future. Paleontological ... · monograph on geology and palaeontology of European Russia in 1845. Since then, and continuing today, there have been

Benton, M. J., Briggs, D. E. G., Clack, J. A., Edwards, D., Galway-Witham,J., Stringer, C. B., & Turvey, S. T. (2017). Russia–UK Collaboration inPaleontology: Past, Present, and Future. Paleontological Journal, 51(6), 576-599. https://doi.org/10.1134/S0031030117060028

Peer reviewed version

Link to published version (if available):10.1134/S0031030117060028

Link to publication record in Explore Bristol ResearchPDF-document

This is the accepted author manuscript (AAM). The final published version (version of record) is available onlinevia Springer at https://doi.org/10.1134/S0031030117060028 . Please refer to any applicable terms of use of thepublisher.

University of Bristol - Explore Bristol ResearchGeneral rights

This document is made available in accordance with publisher policies. Please cite only the publishedversion using the reference above. Full terms of use are available: http://www.bristol.ac.uk/pure/user-guides/explore-bristol-research/ebr-terms/

Page 2: Paleontology: Past, Present, and Future. Paleontological ... · monograph on geology and palaeontology of European Russia in 1845. Since then, and continuing today, there have been

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Russia-UKcollaborationinpalaeontology:past,present,andfutureM.J.Bentona,D.E.G.Briggsb,J.A.Clackc,D.Edwardsd,J.Galway-Withame,C.B.

Stringere,andS.T.Turveyf

aSchoolofEarthSciences,UniversityofBristol,Bristol,BS81RJ,U.K.

bDepartmentofGeologyandGeophysics,YaleUniversity,NewHaven,Connecticut06520,

U.S.A.

cUniversityMuseumofZoology,Cambridge,DowningStreet,CambridgeCB23EJ,U.K.

dSchoolofEarthandOceanSciences,CardiffUniversity,MainBuilding,ParkPlace,Cardiff,

CF103AT,U.K.

eDepartmentofEarthSciences,NaturalHistoryMuseum,CromwellRoad,London,SW7

5BD,U.K.

fInstituteofZoology,ZoologicalSocietyofLondon,Regent'sPark,London,NW14RY,U.K.

Abstract–ThereisalonghistoryofcollaborationbetweenRussiaandtheUnited

Kingdominpalaeontology.Thisbegan,arguably,in1821,withtheseminalworkby

WilliamFox-Strangways,whoproducedageologicalmapoftheareaaroundSt

Petersburg.Mostfamously,RoderickMurchisoncarriedoutextensivesurveyingand

observationsthroughoutEuropeanRussiain1840and1841,andpublishedamajor

monographongeologyandpalaeontologyofEuropeanRussiain1845.Sincethen,and

continuingtoday,therehavebeenmanyfruitfulcollaborationsonPrecambrianlife,

Palaeozoicmarineorganisms,terrestrialisationofplantsandvertebrates,thePermian-

Triassicmassextinction,fossilmammals,humanevolution,andconservation

palaeobiology.

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INTRODUCTION

TheRoyalSociety(RS)andRussianAcademyofSciences(RAN)agreedtohold

meetingstodiscussfuturecollaborativeresearchdirectionsindifferentareasofmutual

scientificinterest.Thefirstthemetobeselectedwaspalaeontology,andaRoyalSociety

delegationofsixpalaeontologiststravelledtoRussiainOctober2016toexplorepossible

collaborationswiththeirRussiancounterparts.Co-chairedbyProfessorMikeBenton

FRS(Bristol)andProfessorSergeiRozhnov,DirectorofthePalaeontologicalInstitute

(PIN)andcolleagues,itfeaturedadiversityofpresentationsencompassingavastrange

ofgeologicaltime,fromthePrecambriantotheemergenceofHomosapiens.

Inthisaccount,weprovideaperspectivefromtheBritishsideofthemeeting,

outliningthemesthatwerepresentedanddiscussedduringadayofsymposium

presentationsattheBorissiakPaleontologicalInstitute,Moscow(PIN),akeydivisionof

RAN.Sectionsofthispaperwerewrittenasfollows:Introduction(M.J.B.),Theearly

yearsofcollaboration(M.J.B.),Exceptionalpreservationrevealstheearlyevolutionof

lifeintheoceans(DEGB),Terrestrialisation,earlyplantsandchangingearthsystems

(D.E.),Documentingmajorevolutionarytransitions–thefirsttetrapods(J.C.),The

impactofmassextinctionsonthehistoryoflife(M.J.B.),Theearlyhumanoccupationof

Britain(C.B.S.andJ.G.-W.),andConservationpalaeobiology(S.T.T.)

THEEARLYYEARSOFCOLLABORATION

CollaborationsbetweenRussianandBritishearthscientistsandpalaeontologists

beganattheveryrootsofgeologyasascience.Anunusualearlymanifestationwasan

articleentitled‘GeologicalsketchoftheenvironsofPetersburg’publishedinthe

TransactionsoftheGeologicalSocietyofLondonin1821.TheauthorwasWilliamFox-

Strangways,4thEarlofIlchester(1795–1865),andhewroteitwhenhewasattachéat

theBritishEmbassyinStPetersburg.Fox-Strangwaysventuredoutaroundtheglittering

imperialcapitalcityforsome40versts(c.30miles)inalldirections,exploringthe

alluviumandunderlyingrocksaroundthedeltaoftheRiverNeva.Amongsthisdiverse

observationsontheroads,rivers,villages,soils,androcks,Fox-Strangwaysmentioned

fossilorthoceratites,‘sometimesayardinlength’andtwospeciesoftrilobiteinthe

‘Pleta’LimestoneatKrasnoeSelo,identifiedastheLowerSilurianOrthoceratite

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Limestone,knownoverpartsofFinlandandSwedenalso,andsubsequentlywidelyused

asadecorativestoneinbuilding.Fox-Strangways(1821)providedacolouredgeological

mapoftheStPetersburgregion,distinguishingfourrocktypes,aswellassomecross-

sectionsandscenicviewsalongtheriverbanks.

Fox-StrangwayswaswritingatatimebeforethesynopticworksofLyell(1830–

1833)andMurchison(1839),whodidsomuchtoestablishtheprinciplesofmethodsin

geologicalobservationandtheinternationalstratigraphicsystem.Infact,Roderick

ImpeyMurchison(1792–1871;Fig.1)isregardedbymanyasthefounderofgeological

researchinRussia.MurchisonfirstvisitedRussiain1840,andsawthegeologyofthe

westandtheMoscowBasin.Hisintentionwastocheckthevalidityofthestratigraphic

systemofthePalaeozoicthatheandAdamSedgwickhadestablishedintheUnited

Kingdom.HetravelledthroughGermanyandtheBalticcoast,observingexamplesofthe

Silurian,Devonian,andCarboniferoussystemsashewent,andconfirmingtheir

occurrence,includingtheOrthoceratiteLimestoneatStPetersburg.Buthismainquest

wastofillthefinalgapintheuniversalstratigraphicschemehehaddrawnup,namely

identifyingwhathappenedbetweenthealreadynamedCarboniferousandtheTriassic.

Afterlengthynegotiations,MurchisonhadachievedthebackingofTsarNicholas

I,andherevelledintheroyalpartiesanddiscussions.HewascommissionedbytheTsar

tocarryoutamuchmoreextensivesurveyofthegeologyofEuropeanRussia,andhe

readextensivelyinplanningforthetrip.Inparticular,hebenefitedfromthedetailed

accountswrittenbyAlexandervonHumboldt(1769–1859)whohadtravelledwidely

aroundRussiain1829exploringtheUralMountainsfortheireffectsontopographyand

climate,butalsotheminesforvaluablemetals(Humboldt1831).Murchisonwas

enthralledbyHumboldt’sadventures,especiallysincetheRussianofficialsandprinces

themselvesknewsolittleoftheremoterpartsoftheirownland.Murchison’sexpedition

wascommissionedinasimilarwaytoHumboldt’s:hewasfundedandinstructedinhis

aimsbyTsarNicholasIandRussianForeignMinisterCountGeorgvonCancrin.

MurchisonbeganhissecondRussianexpeditionin1841,settingoutfromSt

Petersburgwithhisscientificcompanions,theFrenchpalaeontologistEdouardde

Verneuil(1805–1873),theGerman-RussianminingexpertAlexandervonKeyserling

(1815–1891),andayoungRussianmineralogist,NikolaiKoksharov(1818–1893).Their

routetookthemtoMoscow,andtheneastontheroadtoNizhnyNovgorod.They

proceededtogethertotheoldmonasterycityofVladimir,andthenVerneuiland

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KeyserlingtookasoutherlyroutetoKazan’,andMurchisonandKoksharovcontinuedon

themainroadeast.DetailsofthetripandMurchison’sobservationsarerecordedin

detailinaseriesof13fieldnotebooksaswellasanautobiographicaljournal,innine

volumes,allpreservedinthearchivesoftheGeologicalSocietyofLondon(thelatter

transcribedandpublished:CollieandDiemer2004).

AtVyazniki,atown300kmeastofMoscow,Murchisonnotedhowatlastthe

partyhadpassedfromthegrey-colouredCarboniferousrocksoftheMoscowBasininto

younger,redbedrocksthathelateridentifiedasequivalentinagetothemarinePermian

hesawintheUrals.ThiswashisfirstsightingofthePermian(Bentonetal.2010).

MurchisoncontinuedtoKazan’,afterpassingdowntheVolga,andthepartyproceeded

toexplorethewesternflanksoftheUralMountains,reachingPerm’amonthlater.

Murchisonsawgypsum-bearingwhitelimestones,whichheassignedtothelowerpart

oftheNewRedSandstone,and,onlaterdays,marinebedsoftheLowerPermian.The

partycriss-crossedtheUralMountains,exploringthePermianandTriassicredbeds

aroundOrenburginthesouth,andthenswingingfarwesttoexaminetheDonetzcoal

field.TheyfinallyreturnedthroughMoscowtoStPetersburg,afterajourneylastingfor

some5months.BeforereturningtotheUK,Murchison(1841)wroteashortnotein

whichheestablishedthePermianSystem,foundedprimarilyonthemarinerocks

aroundKazan’andPermitself,andOrenburginthesouth:limestoneswithshells,

gypsum,salt,andcopper-bearingsandstones,lithologiesthathehadseeninthe

ZechsteinofGermanyandMagnesianLimestoneofEngland.Heusedhisinformation

fromRussiainnumerouspublications,andespeciallyinhismagisterialaccountofthe

wholeexpedition(Murchison1845),publishedinlavishstyleattheexpenseoftheTsar

(Fig.1).

FurthercollaborationsbetweenRussiaandtheUKfocusedonmorespecialist

topics.Forexample,boneshadbeenreportedasearlyas1770fromthePermianCopper

SandstonesofOrenburg,andMurchisonbroughtspecimensbackin1841forSirRichard

Owen(1804–1892)tostudy.Variousspecimenshadbeendescribedinthe1830sand

1840sbyRussiangeologistsS.S.Kutorga,F.WangenheimvonQualen,andG.I.Fischer

vonWaldheim(OchevandSurkov2000),atthesametimethatOwenwasnaming

materialsofsimilaragefromtheKaroobasininSouthAfrica.Eventually,Owen(1876)

providedathoroughoverviewoftheRussiansynapsidreptiles,subsequentlythesubject

ofextensiveexpeditionsintheSouthUralsandalongthebanksoftheRiverNorthDvina,

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andamajorfocusinmodernRussianpalaeontology(OchevandSurkov2000;Bentonet

al.2000).

Inthisaccount,wesummarisekeypalaeontologicalthemesofinternational

importance,emphasizingthepotentialforfurtheringcollaborationbetweenresearchers

inRussiaandtheUK.Weseetheseasareasforfuturedevelopment.Thetopicsrange

fromtheoriginandearlyevolutionoflifeinthePrecambrian,exceptionallypreserved

Palaeozoicmarinefaunas,theoriginofplantsandanimalsonland,majormass

extinctions–especiallythePermo-Triassicmassextinction252millionyearsago,

humanevolution,andconservationpalaeobiology.

EXCEPTIONALPRESERVATIONREVEALSEARLYEVOLUTIONOFLIFEINTHEOCEANS

Themarinefossilrecordisdominatedbythebiomineralizedelementsof

organisms–shells,bonesandteeth.Surveysofmodernmarinecommunitiesshowthat

some60%ofanimalslackbiomineralized‘hard’parts(Schopf1978),aproportion

exceededinsomeassemblagesofexceptionallypreservedfossilssuchasthoseofthe

MiddleCambrianBurgessShaleofCanada(ConwayMorris1986).Itfollowsthathard

partsaloneprovideasignificantlyimpoverishedsampleoftheorganismsoriginally

present.Conversely,exceptionalpreservations(Konservat-Lagerstätten)arecriticalto

ourunderstandingofthehistoryoflifeonEarth.Thesoft-bodiedfossilspreservedin

Konservat-Lagerstättenfillgapsintherecordofgroupswithanotherwisepoorfossil

record.Theyareimportantinfleshingouttheevolutionaryhistoryofthesegroupsand

theyprovidedataonmorphologiesnolongerrepresentedtoday,whicharecriticalfor

reconstructingphylogenies.Thepreservationofsoft-bodiedtaxaalsoallowsthe

calibrationofmolecularclocks,whichprovideestimatesofthetimingofeventsinclade

evolution.

BiomineralizationwasrarepriortotheCambrianexplosion.Fossilizationin

Precambrianrocksgenerallydependsonthesurvivaloforganicremains,whichrequires

exceptionalconditions.Nonethelessthereisasubstantialrecordofmicroscopicorganic

fossilsinthePrecambrianofRussiawhichistooextensivetosummarizehere:Archean

andProterozoicexamplesaretreatedinreviewsofglobaloccurrencesbyKnolland

Sergeev(1995)andSergeevetal.(2007,2010)andimportantnewdiscoveriescontinue

tobemade(e.g.Golubkovaetal.2015).Thefirstlargeorganisms,whicharelikewise

non-biomineralized,becamewidespreadduringtheEdiacaranPeriod.Someofthemost

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celebratedexamplesaretheclassicEdiacarafossilsoftheWhiteSeaarea(Fedonkinet

al.2007).Suchexceptionalpreservationsarecriticaltounderstandinghowthe

evolutionofoceanecosystemsrelatestoenvironmentalfactorssuchasthechemistryof

theatmosphereandoceans,andviceversa.

TheWhiteSeafossilsarepreservedasthree-dimensionalimpressionsresulting

fromearlycementationofthehostsediment(Fig.2a),butotherfossilsofEdiacaranage

surviveasorganicmaterial(bothmicro-andmacrofossils)andasaresultofdiagenetic

replacementinphosphate.LargeEdiacarafossilsareknownworldwide,mostnotably

fromMistakenPointinNewfoundland,theFlindersRangesinAustralia,andfrom

Namibia.Theirpreservationhasbeenexplainedbyearlyprecipitationofpyrite–the

‘DeathMask’hypothesisdevelopedbyGehling(1999)basedonspecimensfromthe

FlindersRanges.RecentevidencefromAustralianexamplesindicatesthatthepyrite

coatingformslaterintheFlindersRangesandfossilizationappearstobearesultof

elevatedconcentrationsofsilicaintheoceans(Tarhanetal.2016)priortotheonsetof

silicabiomineralizationbyspongesandradiolarians.Thisnewmodel(Tarhanetal.

2016)remainstobetestedbyinvestigatingoccurrenceselsewhereintheworldwhere

pyriteisassociatedwithEdiacarafossils,includingMistakenPoint(Liu2016)andthe

WhiteSea(Ivantsov2016).

LocalitiesyieldingEdiacarafossilsaremuchrarerintheUnitedKingdomthanin

RussiabutexamplesfromCharnwoodForestwerethefirsttobepositivelyidentifiedas

Precambrian,inthe1950s(Fig.2b).RecentresearchintheCharnwoodForestinlierhas

revealedsome14taxa(P.R.Wilby,pers.comm.)andshownthatthebiotaismostsimilar

totheAvalonAssemblageofNewfoundland(Wilbyetal.2011).Beddingplanesurfaces

withhundredsofspecimenshaveallowedpopulationstudieswiththeaidofnew

techniquesformouldingandimaging(Wilbyetal.2015).AfewexamplesofEdiacara

fossilshavealsobeenrecoveredfromtheCarmarthenareainWales.

Amajorareaofpalaeontologicalresearchisthenatureofthetransitionfromlife

intheEdiacarantothatintheCambrian(Rozhnov2010).FortunatelytheCambrian,like

theEdiacaran,ischaracterizedbyanabundanceofsitesofexceptionalpreservation:

therearemanymoreKonservat-LagerstättenofCambrianagethaninyoungerrocks.

InvestigationofthesmallshellyfossilsthatpredatemorefamiliarCambrian

macrofossilswaspioneeredbyA.Yu.RozanovandothersinRussia(Rozanovand

Missarzhevsky1966;MatthewsandMissarzhevsky1975;seeBengtson2005).These

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phosphatizedshellsweredissolvedoutoflimestones,revealingaremarkable

abundanceanddisparityofearlyCambrianformsotherwiselargelyunknown.

Diageneticphosphatizationcanalsopromotethepreservationofembryosandlarval

stages:so-calledOrsten-typedepositsareknownfromtheCambrianofSiberia(Müller

etal.1995;BengtsonandZhao1997).TheLowerCambrianofSiberiahasalsoyielded

someoftheearliestspongereefs,dominatedbyarchaeocyaths(e.g.Ridingand

Zhuravlev1995).

OurunderstandingoftheCambrianExplosionisintimatelyassociatedwith

Walcott’sdiscoveryoftheBurgessShaleintheearly20thcenturyandthereinvestigation

ofthebiotainitiatedinthe1960sbytheGeologicalSurveyofCanadaandcarriedoutin

CambridgeunderthedirectionofHarryWhittington(Gould1989).Itisimportantto

understandthefactorsthatinfluencepreservationsothatwecandistinguishreal

patternsinthediversificationoflifefromthosethatreflectpreservationalbiases.

BurgessShale-typepreservations,likethoseofEdiacaranage,areglobalindistribution

anditislikelythatthecontrolsontheiroccurrencewerealsoglobal.Anumberof

factorshavebeeninvoked,includingtheobservationthatburialmayhaveplaced

potentialfossilsbeyondthereachofburrowinganimals,mostofwhichcouldonly

penetratetoshallowdepthsduringtheCambrian;deeperburrowingformsevolvedlater

(butseeGainesetal.2012b).Thereismountingevidence,however,thatocean

chemistryplayedacentralroleinfossilpreservation.Theremayhavebeenlowerlevels

ofsulphateintheCambrianoceans,inhibitingdecaybysulphatereducers.Inaddition

marinewaterswerecharacterizedbyahigherconcentrationofalkalis,whichpromoted

precipitationofcarbonatecements,reducingdiffusionandslowingdegradation(Gaines

etal.2012a).Recentstudiessuggestthatthecompositionofclaymineralsintheearly

oceansmayalsohaveservedtoinhibitbacterialactivity(WilsonandButterfield2014;

McMahonetal.2016;Naimarketal.2016).Suchconsiderationsmayexplainwhy

Sedgwick’sclassicCambriansequencesintheUKhaveyieldednothingtomatchBurgess

Shale-typepreservationelsewhereintheworld.

OurknowledgeofthediversificationofmetazoansintheearlyPalaeozoicis

increasingapace.Explorationofareaswhereexceptionalpreservationsoccur,inrocks

ofvariousages,havedemonstratedthatKonservat-Lagerstättenarerarelyunique:

exceptionalpreservationoccureswheresimilarconditionsprevail.Examplesincludethe

multipleoccurrencesofBurgessShalefossilsinthevicinityofWalcott’sfamousquarry

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intheMiddleCambrianofBritishColumbia(Briggs2014).SpectacularnewCambrian

assemblageshavealsobeendiscoveredinChina,notablytheolderChengjiangand

GuanshanbiotasandthemiddleCambrianKailibiota.Furtherexcavationshave

augmentedourknowledgeofotherearlyCambrianbiotas,includingthatfromSirius

PassetinGreenland,andtheEmuBayShaleinAustralia.Stratigraphicoccurrencesof

someiconicCambriangroupssuchasanomalocarididshavealsobeensignificantly

extendedbydiscoveriesintheOrdovicianofMorocco(VanRoyetal.2010,2015).The

EarlyCambrianSinskbiotaontheSiberianPlatformnearYakutskisanimportant

sourceofdataonBurgessShale-typefossils,astheycomenotfromshalesbutfroma

carbonateunitknownastheAlgalLens.Thebiotaisdiverse,includingtrilobites,

bradoriids,lobopods,otherarthropodsincludingPhytophilaspis,paleoscolecids,

brachiopods,sponges,chancelloriids,eldoniidsandprobablepterobranchs,arangeof

taxasimilartothatofWalcott’sBurgessShale(Ivantsov1999;Ivantsovetal.2005).The

MiddleCambrianZelenotsvetnayaFormationontheUkukitRiverontheSiberian

Platformhasyieldedrhabdopleuridswithpreservedzooids(Sennikov2016).Burgess

Shale-typepreservationsarerareintheintervalbetweentheMiddleCambrianandthe

LowerOrdovician,presumablyreflectingalackofsuitablesedimentaryfacies(i.e.

extensivedepositsofmudstonesintroducebyepisodicevents).Recentresearchisfilling

thisgapwithdatafromsmallcarbonaceousfossils(e.g.Smithetal.2015;Slateretal.

2017).

FewmarineKonservat-Lagerstättenhavebeenreportedfrompost-Cambrian

stratainRussia,presumablyreflectingthechallengesinvolvedinexploringsuchavast

country.ThePalaeontologicalInstituteinMoscow,however,holdssignificantmaterial

fromanumberofimportantterrestrialsiteswhichhaveyieldedprolificfossilinsectsof

Permian,TriassicandMesozoicage,togetherwithsimilarcollectionsfromadjacent

territories(GrimaldiandEngel2005).RussianKonservat-Lagerstätten,bothnowandin

thefuture,representafundamentalresourcefortheinvestigationofthehistoryofthe

planet.

TERRESTRIALISATION,EARLYPLANTSANDCHANGINGEARTHSYSTEMS

Thecolonisationofthelandbyplantswasnotonlyamajoreventinthehistoryof

lifeonEarth,butonethathadfarwiderimpactsonthelithosphereandatmosphere.

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Initially,intensivestudiesofterrestrialisation,whichbeganinthe1960s,concentrated

ontheoriginsandradiationsofvascularplants(tracheophytes)intheLateSilurianand

EarlyDevonian,but21stcenturyworkhasyieldedincreasingevidence,beginninginthe

mid-Ordovician,forearlierlandplantsbasedonpalynomorphsratherthanmegafossils.

Configurationsandultrastructureofthesespores,plusnewphylogenetictrees,indicate

affinitieswithbryophytes.Moreconjecturalarehypothesesrelatingtotheexistenceof

furthervegetationinwhichbasalembryophytes(bryophytes+tracheophytes)were

accompaniedbyassociationsoffungi,includinglichens,algae,cyanobacteriaand

bacteria,collectivelytermedcryptogamiccovers.Theseorganismstodaycolonise

habitatsinhospitabletovascularplants.Researchonearlyterrestrialanimalshasbeen

lessproductive,butprovidesinformationonbasalmembersofmanylineagesandthe

evolutionofrudimentaryfoodwebs.Interactionsbetweenplantsandfungi,particularly

involvingmycorrhizae,arebeingincreasinglystudied,asaretheimpactsofplantson

sedimentologicalandenvironmentalprocesses.Theseincludetheirroleinthechemical

weatheringofrocks,withimplicationsfortheevolutionofatmosphericcompositionand

thesequestrationofcarbondioxide.

Returningtotheevolutionoftracheophytes,workonfossilsfromNorthAmerica

andEuropehasshownaninitialproliferationofplantsoftheCooksoniatypeinthelatest

SilurianandearliestDevonian.Thiswasfollowedbythediversificationofplantswith

lateralsporangiaandthebeginningsofthelycophytelineageinthebasalDevonian(the

LochkovianZosterophyllumFlora)andthentheemergenceandsubsequentradiationof

theeuphyllophytes(thePragian-EmsianPsilophytonFlora).Thisapparentlystepwise

chronologicalprogressionofeverincreasingplantcomplexitywasshatteredbythe

discoveryofthelateSilurianlycophyte,BaragwanathiainAustraliaandzosterophyllsin

ArcticCanada,aswellasthedisparityandcomplexitydisplayedinPragian/Emsian

plantsfromYunnan,China,manyofwhichdonotfitcomfortablyintoexisting

classifications.

ButwhatofpalaeobotanicalknowledgeofRussiansequencesofthisage?Ina

landmarkpaperin1967,NinaPetrosyanproducedlistsofLowerDevonianspecies

collectedfromthreegeographicalareas–theTunguskaregionofwesternSiberia,

centralKazakhstan,andtheVolyno-PodolianmarginoftheRussiancontinental

platform,andwith,forthefirsttimeinEnglish,acommentaryontheir

palaeophytogeographicsignificance.Theseareasareconsideredhere,emphasizingthe

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roleofRussianpalaeobotanistsinthecollection,descriptionandsubsequentdeposition

offossilsinRussianInstitutions,andincludestheUkraineforcompleteness.

TheSiberianplantassemblagesareimportantnotonlybecausetheycontain

plantsfromLochkovianthroughEmsianstrata(termedtheZosterophyllumand

PsilophytonFloras),butalsobecausetheywerelocatedontheSiberianpalaeocontinent.

ThiswastheonlymajorcontinentinthenorthernhemisphereinDevoniantimes,soits

isolationpotentiallyoffersevidenceforglobalprovincialismintheearlyhistoryof

terrestrialplants.ComprehensivespecieslistswerebasedinitiallyonstudiesbyA.

Ananiev(e.g.1955,1960),whoincorporatedvaluablegeologicaldataontheplant

localities.SubsequentpublicationsbyAnanievandStepanov(1969),Stepanov(1975),

andZacharova(1981)testifytothelegacyofhisleadership.Dataaresupplementedby

collectionsofLepekhina,PetrosyanandRadchenko(1969),whichD.E.wasprivilegedto

examineinStPetersburgonherfirstvisittoRussiaintheearly1970s.In2002,she

studiedsomeoftheAnanievschool’scollectionsinthemuseumofTomskUniversity,

courtesyofDrV.M.Podobina.

Interrogationofspecieslistsandsomepersonalobservationsindicatethatthere

areanumberofSiberianendemicgenera,pluscosmopolitanrepresentatives,that

requirereinvestigation.Thus,forexampleStepanov(1975)describedCooksoniapertoni

andC.hemisphaericaintheKuznetskBasin.TheseareimportanttaxaintheUpper

SilurianandbasalDevonianofEurope,buttheRussianspecimensshowmuchgreater

architecturalcomplexityandshouldbeassignedtoanewgenus.Thedifficultiesof

namingsterilespinyaxeshavebeenexemplifiedinEuropeanmembersofPsilophyton

whereithasbeenshownthatP.goldschmidtiiisayoungersynonymofP.burnotense.

SpinyaxesfromSiberiawerenamedP.goldschmidtiibyAnanievandinitiallyby

Zacharova.TheidentityandaffinitiesoftheRussianmaterialweresubsequently

resolvedbythedemonstrationofitslateralsporangia,whichnecessitateditsremoval

fromthegenusPsilophyton(withterminalsporangia)tothenewgenusMargophyton

(Zacharova1981).Inafinaltwist,thedemonstrationofterminalsporangiainEuropean

P.burnotensehasconfirmeditstrimerophytestatusandhencecannotbethesametaxon

astheSiberianexampleswhichbelongtothelycophytelineageandnowrequireanew

speciesname(Schweitzer1989).Sucharethepitfallsofrecognisingvegetative

convergenceintheabsenceofanatomyincompressionandimpressionfossilsandthis

exampleprovidesanindicationoftheextentofresearchactivityrequiredtoconfirmthe

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presenceofcosmopolitantaxa!Well-illustrated,intriguingendemics(e.g.Christophyton

kuznetzkianum)needsimilarattention.Afurthercharacteristicisthelackoflycopsidsin

theSiberianassemblages.Drepanophycusspinaeformisisanexception,butisitselfin

needofrevision.

TheKaragandaandJunggarbasinsinKazakhstanpresentdifferentkindsof

phytogeographicpuzzles,becausetheareaisanamalgamofanumberofsmallplates,

reflectingaverycomplexgeologicalhistory.Theplantsofthecentralarea(Karaganda)

thusflourishedonaplatedistinctfromthatoftheneighbouringJunggarBasin.Fossils

fromKaraganda,centralKazakstan,weredescribedbyYurina(1969)whosecollections

D.E.wasabletoperuseinMoscow,andbySenkevich(1978,1980).Marineincursions

allowsomeindependentdatinginvolvingbrachiopodsandgraptolites.Again,

representativesofcosmopolitantaxasuchasCooksonia(C.crassiparietilisYurina,1964,

1969),PsilophytonandZosterophyllumneednewassignationasdothenumerous

lycopsidswhich,incontrasttoSiberia,dominatetheassemblages.Theselycopsids

includeEuropean/NorthAmericantaxasuchasDrepanophycusspinaeformis,D.

gaspianusandProtolepidendronwahnbachenseandendemicLidasymophytonakermensis

(Senkevich1980)—allinneedofrevision.Suchuncertaintiespreventadequate

assessmentofpalaeogeographicrelationshipsoftheflora,althoughsuperficialscrutiny

suggestsgreatsimilaritieswithLaurussiaratherthanSiberia.Equallyinterestingarethe

olderassemblagesfromtheJunggarbasininthesouth.FossilsintheTokrauFormation

occurinterrestrialrocksinterbeddedwithmarinefaciesthatyieldgraptoliteswhich

allowconfidentdating(Senkevich1975,1986)asuppermostSilurian(Přídolí).The

plantsincludemembersoftherhyniophytecomplex,butwithorganisationsmore

complexthanincoevalrockselsewhere,plusspinyand‘leafy’axes.Asimilar

compositionisfoundinadjacentXinjiang(China)whereJunggariaisprobably

congenericwithSenkevich’sCooksonella,althoughtherearefurthernomenclatural

problems,whilegraptolitesconfirmtheageasPřídolí(Caietal.1993).Intensive

fieldworknearHoboksar,north-westXinjiangisprovingveryproductiveand

demonstratestheimportanceoftheJunggarassemblagestounderstandingpossible

provincialismearlyinthecolonisationofthelandbytracheophytes,whilereinforcing

theneedforrevisionoftheKazakhstanfossils(Caietal.1993).

TheVolyno-PodoliamarginoftheRussianplatform,Ukrainehasyielded‘leafy’

stemsofuncertainaffinityfromtheLateSilurianSkalahorizon(Istchenko1969).

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SectionsexposedinthebanksoftheDniestrriverextendintotheLowerDevonianand

preservevascularplantsandalgaeinamarginalfacies.Duringameetingofthe

InternationalSubmissionoftheIUGConSilurianstratigraphyin1983,ledby

academicianSolokov,itwaspossibletoexaminetheextensivecollectionsofT.A.and

A.A.IstchenkoinKiev.PlantsfromtheSkalahorizon(Přídolí)includealgae

(Primochara),rhyniophytes(Cooksonia,Eorhynia(?Salopella))andpossiblytheearliest

hepatic,Prehepaticites(seebelow),aswellastheleafyforms(includingLycopodolia),

whoselycopsidcredentialsneedfurtherscrutiny.Inaddition,thesucceedingLower

Devonianassemblages(Istchenko1965)containplantsalsofoundinLaurussia,aswell

aslessfamiliartaxasuchasTirasiophytoneuropea,buttheirageisbasedontheplants

themselves

ThereisevidenceofevenolderlandplantsfromtheUpperOrdovicianofthe

TungusBasinontheSiberianPlatform,basedonpalynologicalassemblages(Raevskaya

etal.2016).Theyinclude,togetherwithacritarchs,prasinophytesandchitinozoans,

cryptosporesintheformofnakedandenvelopedmonads,dyads,tetradsandpolyads,

similartothosefromrocksofthesameage(Fig.3).Previousultrastructuralstudieshad

indicatedhepaticaffinitiesofthedyads,whilethetetradconfigurationisnotedin

certainextanthepatics.Basalembryophytes(bryophytes+tracheophytes)arepoorly

knownworldwide.InsitudyadsandtetradsfoundinsporangiaintheLowerDevonian

ofsouthernBritainledtotherecognitionofanewgroupofbasalembryophytesthat

possessbothtracheophyteandbryophytecharacters(Edwardsetal.2014).Themid

Palaeozoicmegafossilrecordofbryophytesismeagre.TheSilurianPodolianfossil

namedPrehepaticitespredatestheoldestunequivocalhepaticintheUpperDevonian,as

dotheoverlookedrosette-likethalloidfossilsnamedSciadophytopsisandRiccielliopsis

fromtheMiddleDevonianofPodolia(IstchenkoandSclyakov1979),allofwhichare

urgentlyinneedofrevaluation.

Finally,mentionshouldbemadeofprogressindisentanglingtheaffinitiesof

membersoftheDevonianorderSpongiophytales,betheytracheophytes,algaeoreven

lichens.Theyareunitedinbeingessentiallyaxialorganismswithverythickcuticlesand

includesuchdisparatetaxaasSpongiophyton,OrestoviaandBiteleria.Spacedoesnot

allowadetailedreviewofthecontroversiessurroundingdescriptionandidentification

ofspecimensfromRussia,butBroushkinandGordenko’s(2009)descriptionofanew

tracheophyte,Istchenkophytonfiliciforme,fromtheKuznetskBasindemonstrateshow

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rigorousexamination,especiallyemployingelectronmicroscopy,canresolvesuch

issues.

Inencouragingthesearchfornewfossils,particularlywithanatomy,and

renewedactivityonRussianassemblages,aswellasthosefromotherpartsofthe

formerSovietUnion,D.E.concurswiththelateSergeiMeyen,themostinfluentialand

insightfulpalaeobotanistofhisgeneration(Meyen1987),whenhewroteinapersonal

letterrespondingtoherrequestforinformationontheKazakhstanassemblages,‘This

enigmawillnotdisappearifwescrutinizenowandagainpublishedlistsandbadly

reproducedphotographs.Oneshouldmerelywaituntilanewgenerationof

palaeobotanistscleantheAugeanstables’.Perhapsnowisthetime!

DOCUMENTINGAMAJOREVOLUTIONARYTRANSITION

Thetransitionfromaquaticvertebrateswithfinsandscales(fish)tothosethat

hadlimbswithdigits(tetrapods),andsocouldliveonland,isoneofabidinginterestnot

onlytopalaeontologists,butalsotothegeneralpublic(Clack2012).Overthepast

twentyyears,ourknowledgehasexpandedmoreorlessexponentially,withdiscoveries

ofDevonianandearliestCarboniferoustetrapods,andthepredecessorsofthetetrapods,

thetetrapodomorphfishes.

Nearly20taxaofDevoniantetrapodsarenowrecognised,includingsomestill

awaitingdescriptionandanalysis,andtheyoccurworldwide(Clack2012).Although

originallyonlyfoundinEastGreenland,someofthemostimportantsubsequentfinds

weremadeinRussia(Fig.4),andthesameistrueforthetetrapodomorphfishesclosest

totetrapods.Thissectiondescribesthesediscoveriesandsomeofthecollaborations

betweenRussian,BritishandotherEuropeanscientiststhathavehelpedbuildapicture

ofthisimportanttransition.

In1984,OlegLebedevannouncedtheDevoniantetrapodTulerpetoncurtumfrom

theTulaRegion.Thisspecimenconsistedofanodulecontainingafewskullbonesbut

alsofore-andhindlimbs,shouldergirdleandpartsofthepelvicgirdle,somevertebral

elementsandgastraliaofasinglepartiallyarticulatedindividual(Lebedev1984,1985).

Intriguingly,itshowsseveraltraitsthataremorereminiscentoflaterCarboniferous

tetrapodsthanthetwothenknownDevonianforms,especiallyintheshouldergirdle

andhumerus.Itwasthoughtanomalous,however,inthattheforelimbcarriesa

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complementofsixdigitsratherthantheconventionalfive.Thehindlimb,althoughthe

pesisincomplete,showsbasesforsixtoes.Aseriesoffurtherpublications,

reconstructionsandanalysesfollowed,includingcomparisonwiththetrackwaysmade

byDevoniantetrapodsfromAustralia.Collaborationsonmoredetaileddescriptions

weremadewithJ.C.andMikeCoates,theninCambridge(LebedevandClack1993;

LebedevandCoates1995).

FurtherdiscoveriesofDevoniantetrapodswithcompletelimbsputTulerpetonin

context.Allthreehadmorethanfivedigits(CoatesandClack1990).Infact,Tulerpeton

wasclearlypartofapatternofmultidigitedtetrapods,initiatingnewinterestinthe

evolutionarydevelopmentalbiologyoftetrapoddigits.Itgeneratednewcollaborations

betweenpalaeontologistsanddevelopmentalbiologists,previouslyquiteseparate

researchendeavours,whichcontinuestothisday(e.g.Shubinetal.1997;Davisetal.

2007;Standenetal.2014).

ThenumberoftoeswasnottheonlysurpriseoftheTulerpetonfind.Itsgeological

contextwashighlyunusualandunexpectedrepresentingahypersalineenvironment

manykilometresfromthenearestlandmass(Alekseyevetal.1994).Itappearedto

contradictthecommonpreviousassumptionthattetrapodsoriginatedinfreshwaters.

Haditbeenwashedouttoseaasamummifiedcarcass?Itsurelycouldnothavebeen

livingthere.BasedonthebiasfollowingthelimitedevidenceprovidedbyEast

Greenland,afreshwateroriginhadbeenassumed(butseeThomson1980foranearly

alternativeview).Subsequentdiscoverieshaveincreasinglyshownthatamarginal

marineorlagoonaloriginismuchbettersupported(Lebedev2004;Lukševičsand

Zupins2004).TulerpetonwasthefirstDevoniantetrapoddiscoverytopromptquestions

aboutourassumptionsonthis.

SincethediscoveryofTulerpeton,otherDevoniantetrapodshavebeenfoundin

Russia.Jakobsonia,describedbyLebedev(2004),camefromanewlocalitywiththe

potentialforfurtherdiscoveries.InthenorthernprovinceofTiman,furthernew

materialawaitingformaldescriptionconsistsofisolatedskullandgirdlebones

preservedinalimestonematrix.Thesecanbeextractedbyaciddigestionandrevealnot

onlythatseveralofthebonescanbefittedtogetherandbelongedtoasingleindividual,

butthatthisnewgenusshowssomeremarkablyprimitivefeatures.Workisinprogress

withPavelBesnosovandcolleagueswithJ.C.andPerAhlbergfromUppsala(Ahlberget

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al.2011).UnexploredLateDevoniansequencesinSiberiaalsohavegreatpotentialfor

newdiscoveriesinthisimportantfield(P.Ahlbergpers.comm.).

TheseearlytetrapodfindsrevealtherichpotentialinRussiaforilluminatingthis

crucialtransitioninthehistoryoflifeonearth,withsignificantimpactinproducingnew

ideasandoverturningestablishedhypotheses(Fig.4).

ThetetrapodomorphfishesrepresentanotherareainwhichRussian

palaeontologistshavemadeandcancontinuetomakekeycontributions.Chiefamong

theresearchersinthisareawasthelateEmilaVorobyeva.ShedescribedseveralLate

DevoniantetrapodomorphsincludingPanderichthysandObruchevichthysaswellasa

rangeoflesswellknownearlierforms.

Panderichthyswasregardedasaproblematictaxonfromsomeyears,butwith

thediscoveriesofElpistostegeintheFrasnianofCanada,itssignificancebecame

recognised.VorobyevaandSchultze(1991)describedtheskullofPanderichthysin

detail,drawingattentiontofeaturesofitsskullthatforeshadowedthoseinlimbed

tetrapods.In1992withAlexandrKuznetzov,Vorobyevamadeanassessmentofitslikely

locomotorymodebasedonitsshouldergirdleandforelimb.Althoughmostofits

postcraniumhadnotbeenfullydescribedatthattime,severalarticulatedspecimens

existandnewtechniqueshaveincreasedourknowledge(Boisvert2005;Boisvertetal.

2008).FeaturesofPanderichthysandElpistostege,furtherilluminatedbythefindsof

TiktaalikinCanada(Daeschleretal.2006),revealthatthesethreetaxaaremembersofa

seriesofnear-tetrapodtaxaeachofwhichshowstetrapod-likefeaturesoftheskulland

postcranium.TheRussiancollectionofPanderichthysspecimensremainstobemore

fullyexploited.

Vorobyeva(1977)firstdescribedObruchevichthysbeforethemorerecentstudies

ofDevoniantetrapodshadgotunderway.Thespecimenscomprisetwofragmentsof

lowerjaw.WiththedescriptionofthelowerjawofElginerpetonfromtheFrasnianof

Scotland,Obruchevichthyswasrecognizedasatetrapod(Ahlberg1991).Thephylogeny

oftheTetrapodomorphaiscurrentlyinneedofrevisionwiththeadditionofmembersof

thecladefromRussia.Theseneedurgentrestudyinthelightofmorerecentfindsandof

currentsystematicpractice(Fig.4).

TheendoftheDevonianwasmarkedbyanextinctioneventthatterminatedthe

reignofmanyfishtaxacommonintheso-called‘AgeofFishes’.Itmarkedthedemiseof

boththefish-liketetrapodsandthenear-tetrapodfishes,aswellasothermorearchaic

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groups,atleastasfarasthefossilrecordshows(SallanandCoates2010).Followingthat

extinction,adearthoffindsofcontinentalvertebratessuggestedthatlifeonlandtooka

verylongtimetorecover,inanintervalcolloquiallyknownas‘Romer’sGap’(Coatesand

Clack1995).Bytheendofthisinterval,tetrapodshadradiatedintomorefamiliarforms

andgeneratedfullyterrestriallycapableanimals.RecentdiscoveriesintheTournaisian

ofScotlandhaveinsertedseveralnewtetrapodsinthisgap(Clacketal.2016).Someof

theseindicatecross-overtaxabetweentheDevonianandCarboniferous–ofwhich

Tulerpetonisanearlierindication–suggestingthattheextinctioneventmightnothave

beensodrasticfortetrapods(Andersonetal.2015).Manynewlungfish(Smithsonetal.

2015)andchondrichthyantaxa(Richardsetal.2015)havealsobeenfound.Few

indisputablyTournaisianlungfishwerepreviouslyknown:onefromRussia,

Parasagenodus,fromthe‘LowerCarboniferous’waspoorlydatedandwaslikelynot

Tournaisian(Vorobyeva1972;Lebedevpers.comm.).StudiesinScotlandshowthat

tetrapods,lungfishesandchondrichthyansalldiversifiedrapidlyfollowingthe

extinctioneventandthatcontinentalTournaisianstratacouldproviderich

palaeontologicalrewards.AlthoughRussianTournaisianstrataaregenerallyconsidered

tobemainlymarine,therearepresumablysequencesthataremarginalmarine(e.g.

Lebedev1996)orevencontinental.StudiesofFamennian–Tournaisiansequencesmight

helpresolvesomeofthecurrentdisputesregardingthecauseoftheend-Devonian

extinction,anditsextent(McGhee2013).Continentalvertebratesarealsopoorlyknown

inViseanstrata,andthesetoocouldyieldrichrewardsforthosedeterminedenoughto

seekthemout.

THEIMPACTOFMASSEXTINCTIONSONTHEHISTORYOFLIFE

Therehavebeenmanyextinctioneventsinthehistoryoflife,timeswhenlarge

numbersofplantandanimalspecieshavediedoutinasingleevent,oftentriggeredby

someshockenvironmentalchange.Largestoftheseeventsarethemassextinctions,

when50%ormoreofspecieshavedisappeared,andthemostfamousofthesehappened

66Myrago,whenthedinosaurs,marinereptiles,andammonitesdisappearedatatime

ofshockenvironmentalchangetriggeredbytheimpactonEarthofagiantasteroid.This

eventisnotwelldocumentedineitherRussiaortheUK,andattentionhasfocusedon

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olderevents,mostnotablythelargestmassextinctionofalltime,thePermian-Triassic

massextinction(PTME),whichhappened252Myrago.

ThereareextensivebasinsoflargelyterrestrialPermianandTriassicrocks

betweenMoscowandtheUralMountains,andextendingforsome2000kmfromthe

ArcticcoasttothebordersofKazakhstan.ThePermo-Triassicboundary(PTB)hasbeen

identifiedatmanysitesthroughoutthesebasins,andevidenceforthePTMEfound

amongtheterrestrialtetrapodsinparticular(Bentonetal.2004;Benton2015).The

scaleofthePTMEwashuge.Globalcompilationsofdatashowthatmorethan50%of

familiesofanimalsintheseaandonlandwentextinct,andregionallybasedstudies

showthatthisequatesto90%ofspeciesloss.ThecauseofthePTMEhaslongbeen

debated,butitwasobviouslycatastrophic,andthebalanceofcurrentopinionindicates

thattheSiberianTrapslargeigneousprovinceprovidesthesmokinggun.

TheSiberianTrapscomprise3millioncubickilometersofbasaltlavathatcover5

millionsquarekilometresofeasternRussiatoadepthof400–3000m.Itiswidely

acceptednowthatthesemassiveeruptions,confinedtoatimespanoflessthan2myrin

all,wereasignificantfactorinthePTcrisis(Reichowetal.2009).Eruptionsofthese

basalticlavaswereaccompaniedbytheescapeofhugevolumesofgasessuchas

methane,carbondioxide,sulphurdioxide,andothers.Mixingwithwaterinthe

atmosphere,thesegasesproducedacidrain,whichkilledthelandplants,andthis

releasedthesoilsthatwerealsostrippedofftheland.Withnofood,landanimalsdied.

Warmingisoftenassociatedwithlossofoxygen,andseabedsbecameanoxic,sokilling

lifeinthesea.TheCO2fromtheeruptionsalsocausedoceanacidification,which

doubtlessledtofurtherdistressamongshelledorganisms.

Muchoftheevidenceforthis‘standard’killingmodel(Wignall2015)comesfrom

studiesofmarinesedimentsacrossthePTBinNorthItaly,westernCanada,Spitsbergen,

Greenland,andespeciallySouthChina.Inmarinesections,theend-Permiansediments

areoftenbioclasticlimestonesorintenselybioturbatedmudstonesandsandstones,

indicatingrichly-oxygenatedbottomconditionsforlife.Incontrast,sedimentsdeposited

immediatelyaftertheextinctionevent,intheearliestTriassic,aredark-coloured,often

blackandfullofpyrite.Theylargelylackburrowsandindeedanyfossils.Thesediments

andisotopegeochemistryconfirmaremarkableshifttoanoxicconditionsjustatthe

timeofthePTME.Further,adramaticshiftinoxygenisotopevaluesindicatesaglobal

temperatureriseofaround16°C.Climatemodellershaveshownhowglobalwarming

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canreduceoceancirculation,andtheamountofdissolvedoxygen,tocreateanoxiaon

theseabed.Theoxygenspikeismatchedbyanequallydramaticcarbonexcursion,

indicatingadramaticincreaseinthelightcarbonisotope(12C),comingfromdead

organisms,volcanicsources,andevenmassivereleasesofgashydratesfromfrozen

reservesontheoceanfloor.Thewarmingbecameself-reinforcing,aso-called“runaway

greenhouse”effect,andtheocean-atmospheresystemapparentlyspiraledoutof

control,leadingtothebiggestcrashinthehistoryoflife.

EvidenceforconditionsonlandcomesprimarilyfromRussia,SouthAfrica,and

northernChina.Indeed,onekeycomponentinprovidinglinkagesfromlandtoseacame

fromstudiesintheRussianPermo-Triassicredbedsinthe1990s.Geomingeologist

ValentinTverdokhlebov(Fig.5a)hadplottedgreatalluvialfansthatspreadwestwards

fromtheUralMountainsattheverybeginningoftheTriassic.Thealluvialfansspread

for100–150kmoverthelow-lyingPermianlakesandmeanderingriversonthegreat

plain.TverdokhlebovidentifiedthebouldersinthedifferentbasalTriassicalluvialfans

andtrackedthemtotheirsourceshighintheUralMountains.Theconglomerate

bouldersincludeblocksofDevonianorCarboniferouslimestones,oftenwithfossils,and

metamorphicandigneousrocks.

Duringcollaborativefieldtripsinthe1990s,ledbyValentinTverdokhlebov

(Benton2008),weinvestigatedthePTB,andespeciallythisremarkableshiftin

sedimentaryregime(Fig.5a).TheuppermostPermiansedimentarysuccessionconsists

ofrepeatedfining-upwardcycles,eachbeginningwithacross-beddedsandstone,and

thenfiningupwardsintosiltstonesandmudstonesandendingwithapalaeosol.The

palaeosolsaresometimesassociatedwithplantremainsandarenearlyalwaysinvested

withcarbonate.Thesecyclesareinterpretedasthedepositsofcyclicallakes,with

occasionalinfluxofsediment(thecoarsersands),thenfinerlakedepositsandfinallya

palaeosolwhenthelakedriedout–allperhapstheresultofabroadlymonsoonal

climate(Newelletal.1999).TheearliestTriassicconglomerateswereunexpected,

markinganentirelynewrocktype,andevidenceofahugechangeinsedimentary

regimefrommeanderingtobraidedstreams.

Earlierworkershadinterpretedthisshiftasevidenceforrenewedupliftofthe

UralMountains.TheUralshadbeenupliftedprimarilyinthelateCarboniferousand

earlyPermianastheseparateEurasianandSiberiancontinentalplatescameinto

contact,andtheycouldwellhavebecomeactiveagain.Anotherideaattributedthe

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switchfromlow-tohigh-energydepositiontoadramaticincreaseinrainfallatthePTB.

BothsuggestionswererejectedbyNewelletal.(1999)because,firstly,thesedimentary

regimeswitchisalsoevidentinSouthAfrica,Australia,India,andSpain,andsowas

apparentlyworldwide.Secondly,independentclimaticevidencepointedtoincreasing

aridityacrossthePTB,notincreasingrainfall.Newelletal.(1999)arguedforathird

model,linkingincreasingaridityandmassstrippingofforests.Climatesworldwide

switchedfromasemi-arid/sub-humidclimateinthelatestPermiantogreateraridityin

theearliestTriassic,andthiscouldincreasesedimentyieldbyreducingvegetation

cover.Ifvegetationisstrippedfromthesurfaceoftheland,ratesoferosioncanincrease

perhapstenfold.Localdataconfirmstheshort-termlossofplantsatthePTBinRussia

andelsewhere,aswellasaglobal‘forestgap’,lastingforsome10MyroftheEarlyand

MiddleTriassic,whenforestswereabsentworldwide.Withoutextensiveforestsand

theirroleingeneratingandbindingsoils,erosiononlandmayhaverevertedtoearly

Palaeozoiclevels.Thishassubsequentlybeenconfirmedbystudiesinotherpartsofthe

world,andespeciallybytheobservationofasuddenspikeinsandwashedfromland

intoshallowmarinesedimentsaroundtheworld(AlgeoandTwitchett2010).

TheRussianresearchprovideskeyevidencethatlinkslandandseainthePTME

killingmodel,andthereisenormousscopeforfuturestudiesoftheRussianPermian-

Triassicredbeds.Recentworkcontinuestorevealnewexamplesoffossiltetrapods,and

thesearestudiedinrelationtocoevalanimalsfromaroundtheworld(Fig.5c).The

associatedplants,molluscs,insects,andfisheshavebeenstudiedextensivelybyRussian

palaeontologists,butmuchoftheworkremainsunpublishedbecauseitformedpartof

themappingprogrammesbyGeomin.Forexample,wellestablishedstratigraphic

schemesbasedonpalynomorphs,bivalves,ostracods,andfishes,aswellas

magnetostratigraphy(Fig.5b),areonlyinpart,andtheserequiresubstantialre-study

andpublication.ThepotentialtousetheRussianredbedstodocumenteventsthrough

muchofthePermianandTriassicreliesonsecurestratigraphyandextensive

correlationwithdevelopingschemesinSouthAfricaandChina.Inaddition,increasing

focusonthedetailoftheSiberianTrapsshouldallowresearcherstoidentifyhowthe

killingagencyrolledoutaroundtheworld.Suchworkwillenhanceourunderstandingof

howmassextinctionshaveaffectedlifeonlandingeneral,butalsoprovidedeeper

insightsintothedevastatingPTME.

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THEEARLYHUMANOCCUPATIONOFBRITAIN

EarlyhomininoccupationinRussiaincludestheintriguingremainsfrom

DenisovaCave(AltaiMountains,Siberia),whichhashadahistoryofexcavationby

Russianarchaeologistssincethe1970s.The‘Denisovans’,agroupofhomininscurrently

representedbyonlyafragmentarymanualdistalphalanx(Krauseetal.2010),twolarge

adultmolars(Sawyeretal.2015),andadeciduousmolar(Slonetal.2015;Stringerand

Barnes2015)fromthecave,receivedconsiderableattentionwhenthepreservationof

thefossilspermittedDNAextraction,revealingalineagegeneticallydistinctfrom

modernhumansandNeanderthals(Krauseetal.2010).Thereconstructedgenome

indicatedtheDenisovansaremorecloselyrelatedtoearlyNeanderthalsthanmodern

humans(Reichetal.2010),divergingfromtheNeanderthallineage~450kya,roughly

contemporarytotheearlyNeanderthalsatSimadelosHuesos(Atapuerca,Spain)to

whichtheywerecompared(Meyeretal.2014).TheDenisovanmaterialwasuncovered

frommultiplesequenceswithinthecaveindicatinganextendedperiodofoccupation

(Sawyeretal.2015),150-50kya(StringerandBarnes2015).Neanderthalfossilstoo

havebeendiscoveredfromLatePleistocenedeposits(Prüferetal.2014),suggestingthe

cavewasalternatelyoccupiedbyNeanderthalsandDenisovans(StringerandBarnes

2015);thoughattributionofthearchaeologydatedtotheseperiodsisconsequently

ambiguous.Subsequently,thecavewasalsooccupiedbymodernhumans,basedonthe

presenceofUpperPalaeolithictoolsfrom~40kya(Dereviankoetal.2000).

Morphologically,themolarsaredistinctfromthoseofNeanderthalsandmodern

humansintheirlargesizeandrootandcrownmorphology,withsomearchaictraits

generallyconsideredreminiscentofearlyHomo(Sawyeretal.2015).Thegenetic

evidenceofintrogressionwithanadditional,currentlyunknownhomininspecieshas

beensuggestedtoaccountforthesearchaicfeatures(Sawyeretal.2015),thoughat

presentthesmallandfragmentarynatureoftheDenisovafossilsarenotsufficientto

makeanysignificantmorphologicalcomparisonswiththemajorityofotherfossils,

whichhavenotyieldedgeneticdata.However,thegeneticdiversityofthefourseparate

individualswhosegenomeshavebeensequencedindicatethattheDenisovanshada

populationsizegreaterthanthatofNeanderthals,andtheirgeographicrangelikely

extendedbeyondtheAltairegion(Pennisi2013).Furthermore,theconcentrationof

DenisovanDNAinmodernOceanicpopulations(Reichetal.2010,2011;Meyeretal.

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21

2012),withminimalamountsinmodernmainlandAsianpopulations(Prüferetal.

2014),suggeststhattheDenisovanrangeextendedsignificantlybeyondSiberia,perhaps

evenhavingcrossedtheWallaceline(CooperandStringer2013).Ifthisisthecase,the

potentialrangeoftheDenisovansextendedintoareasofmainlandAsiaforwhichthere

isalreadyanextensivefossilrecord.Infact,severaltaxonomicallyambiguousspecimens

frommainlandAsiahavebeensuggestedtorepresenttheDenisovans,giventheir

relativemosaicismandcombinationsoftraitsassociatedwithNeanderthals,H.

heidelbergensisandH.erectus(CooperandStringer2013;Xingetal.2015;Lietal.

2017).

WhilecollaborationsbetweenBritishandRussianresearcherswithin

palaeoanthropologyhavebeenrarehistorically,thewealthofinformationderivedfrom

theDenisovamaterialshasledtosomecollaborativeresearch,particularlyindatingthe

site(e.g.Brownetal.2016;Doukaetal.2015).However,thereisstillroomtodevelopa

large-scaleinterdisciplinaryapproachtounderstandinghumanoccupationinnorthern

Asia.Increasingly,patternsofhumanoccupationduringtheLatePleistoceneinAsia

appeartohavebeenextremelycomplex,withmultiplespeciesoccupyingtheregionand

occasionallyinterbreeding,asexemplifiedbythedataemergingfromDenisovaand

reflectingthecomplexitythathasalsoemergedinthewesternEuropeanfossilrecord.

In2001,thefirstofthreephasesofalarge-scaleconsortiumprojectcalledthe

AncientHumanOccupationofBritain(AHOB)began,fundedbytheLeverhulmeTrust.

Overits12years,AHOBcollectedandanalyseddatafromassemblagesatnewand

existingsitesthatrelatetothecolonisationsofwesternEuropeduringthePleistocene

(Stringer2007,2011).Duringthistime(Fig.6),datahavecontinuedtoindicatethat

mainlandBritainbecameincreasinglyisolatedfromcontinentalEuropebetweenMarine

IsotopeStage(MIS)11(~400ka)andMIS7(~200ka).Fromthispoint,Britainwas

apparentlywithouthumanoccupationuntiltheendofMIS5(~71ka),afterwhichtime

Neanderthalsrecolonised,followedbymodernhumansenteringBritainforthefirst

timearound40ka.DuringtheLastGlacialmaximum,aswellasforperiodsofthe

YoungerDryas,Britainagainseemstohavebeendevoidofhumanoccupation,though

humansrecolonisedBritainrapidlyfollowingtheseperiods,asindicatedthrough

improvementsinradiocarbondating(see,e.g.JacobiandHigham2009).Theseepisodic

andfragmentedoccupationsofBritainprovideamodelthatisprobablyapplicableto

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manyregionsofEurasia,includingRussia,wheresevereclimaticandenvironmental

changesmusthavealsoregularlydisruptedthepatternsofearlyhumansettlement.

Inthe1970s,twoconflictingviewpointsemergedconcerningthenumberand

sequenceofglacialsandinterglacialsrecognisableinBritainduringtheQuaternary.The

firstfocusedonpollendatainformedbyvegetationalchanges,whichindicatedthat

therewerefourinterglacialsinBritain,withthreeinterveningglaciations(Mitchelletal.

1973).Thealternateviewpointwasthatmammalianbiostratigraphywasmoreaccurate

thanpollenalone.Thisopinionwasreinforcedbyworkonmolluscanaminostratigraphy

(Bowenetal.1986)–togethertheseindicatedthattherecordwasmorecomplexthan

thepollendataindicated.However,debatesabouttheaminostratigraphicmethods

meantthatthedebatewasnotquicklyresolved.Eventually,themammalian

biostratigraphicapproachinspiredMammalAssemblageZones(MAZ),whichhavebeen

invaluableforAHOB(CurrantandJacobi2001),andaminostratigraphywasalsoplaced

onamoresecurefooting(Penkmanetal.2013).

WorktobetterunderstandtheThamesdepositswasalsocarriedoutduringthis

time,whichwasrelevanttothedebateabouttheBritishPleistocenesequence.During

theearlyhistoryoftheThames,theriverhasbeendisplacedprogressivelysouthwards,

amovementthatmeantitaccumulatedlargeamountsofsedimentcriticaltothe

biostratigraphyoftheearlyhumanoccupationofBritain.Crucially,thesedatasuggest

thatthelargestdiversionoftheThameswascausedbyAnglianice,andthiscanbe

correlatedwithMIS12(Bridglandetal.2004).Thesignificanceofthisismarkedfor

sitessuchasSwanscombeinMIS11,withpost-diversioninterglacialdeposits.

Additionally,SwanscombeandMIS11havebeencorrelatedwiththeHoxnian

Interglacial(see,e.g.Bridglandetal.2004;Penkman,etal.2013).

InterpretationsoftheSwanscombehomininhavevariedgreatlysincethe

discoveryofitsfirstcranialbonein1935,includingbeingusedassupportforthe

Piltdownfindsandlaterasamemberofapre-sapienslineage.Sincere-evaluationsin

the1960s,however,ithasmorewidelybeenconsideredtorepresentanearly

Neanderthal(WeinerandCampbell1964;Stringer1974;Hublin1988).Itssimilaritiesin

cranialmorphologytotheSteinheimskullfromtheMiddlePleistocene,inadditionto

homininsfromthelargeassemblageatAtapuerca,Spain,intheSimadelosHuesos(SH)

(BermúdezdeCastroetal.2004;Martinón-Torres2012;Arsuagaetal.2014),provide

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supportforthearchaicaffinitiesoftheSwanscombehomininanditspotential

conspecificitywithNeanderthals.

ItmaybethathumanoccupationinBritainisevenmorecomplexthanwe

recognise,giventhecontinuingenigmaoftheClactonianindustry.TheLowerLoam,a

depositofsiltandsandbeneaththeMiddleGravelsoftheSwanscombeskull(McNabb

2007),containsanassemblageofartefactsthatarecharacterisedbyflaketoolsandthat

donotincludehandaxesattributabletotheAcheulian.Thatassemblageis

representativeoftheClactonianindustry,whichwasoriginallydescribedbasedon

materialfromClactoninEssex.Atonetime,Clactonianartefactswerethoughttobethe

oldesttechnologyinBritain,eitherbeingreplacedbyhandaxesoncetheyarrivedin

Britain,orrepresentingaprecursorindustry.However,thedatingofhandaxesitessuch

asBoxgroveasolderthandepositscontainingClactonianartefactsrefutesthisidea.This

hasraisedthequestionofwhetherthetoolsproducedbytheSwanscombeandother

peopleshaddifferentfunctions.OrdidthepeoplerepresentedbytheClactonianhavea

distinctpatternofliving,withtheabsenceofhandaxessignifyingeithertheinabilityto

producesuchtoolsoraculturaltraditionthatinhibitedtheirproduction(McNabb2007;

Stringer2007)?

ApartialskeletonofPalaeoloxodonantiquus,associatedwithClactoniantools,

wasfoundin2003,atEbbsfleetinKent.ItislikelythatthisClactonianoccupationwas

towardsthebeginningofthesameinterglacialperiodastheClactonianoccupationat

Swanscombe.Forbutchery,ithadpreviouslybeensuggestedthathandaxeswerethe

favouredtoolsduringthelaterBritishLowerPalaeolithic,andthereforethisfindwas

significant,asitassociatedClactoniantools(andanabsenceofhandaxes)withalarge

mammalskeleton.Thissupportedtheideathattheremusthavebeenseparate

populationswithdifferentlithictraditions(Wenban-Smithetal.2006).Thisviewhas

beenreinforcedbyarepetitionofthesameClactonian-AcheuliansequenceattheMIS11

siteofBarnhaminSuffolk(Ashtonetal.2016).

AnadditionalpartoftheClactonianpuzzlewasprovidedbyadiscoveryin1911

atClacton-apieceofyewwoodsharpenedtoapoint.Whileitcouldnotbeconfirmed

thatithadbeenhardenedinafire,microscopicanalysesshowedthatithadbeen

carefullyshaped(Oakleyetal.1977).Theartefactwasbroken,andat33cminlength,

hadmanyinterpretations.Itssignificancewasamplified,however,bythediscoveryof

severalwoodenspearsatSchöningeninGermany(Conardetal.2015).Itisthus

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possiblethattheClactonwoodenartefactrepresentsanimportantcomponentofLower

PalaeolithictechnologythatwasunsuspecteduntiltheSchöningendiscoveries.Neither

theClactonnorSchöningensiteshadhumanremainsassociatedwiththewooden

artefacts.However,anotherGermansite,theSteinrinnetravertinequarrynear

Bilzingsleben,didyieldsomefossilevidenceapproximatelycontemporarywithClacton

andSchöningen.Cranialremainsofthreeindividualswerefound,andfloralandfaunal

evidenceatthesiteindicatesthattheremainsderivedfromawarmMiddlePleistocene

interglacial(probablyMIS11,orperhapsMIS9).Thesehavebeenreconstructedas

eitherHomoerectus(Vlčeketal.2000)orpossiblyHomoheidelbergensis(Stringer

2012).

Atonetime,SwanscombewasconsideredastheoldestsiteintheUKtoyield

archaeologicalandfossilremainsofhumanoccupationinBritain.WhiletheMauer

mandibleofHomoheidelbergensisfromGermanyindicatedthatearlyhumanshad

reachednorthernEuropecomparativelyearlyintheMiddlePleistocene,noconvincing

evidencesuggestedthathumanshadarrivedtoBritainuntilaftertheAnglianice

advanceanditsresultantdiversionoftheThamessouthwards.Itwasarguedthat,given

theclimaticconditionsofBritainatthetime,thehumansoftheearlierMiddle

Pleistocenecouldnothaveweatheredtheconditionsbyproducingclothing,shelters,or

fire.Furthermore,theywereconsideredtobelargelyopportunistichuntersand

scavengersratherthanhuntersoflargegame,andwouldhavebeenvulnerableand

unabletocompetewiththelargecarnivoresoftheperiod.

ArchaeologicalremainsfromcavedepositsatWestbury-sub-Mendipthrewthis

intoquestion,indicatingthathumanswerepresentinBritainduringawarmstageprior

totheAnglian.Basedonbiostratigraphicfaunalanalyses,Bishop(1975)suggestedthat

theremighthavebeenhumanoccupationduringanunrecognisedinterglacialbetween

theCromerianandtheHoxnian(~600kaand~400karespectively).Further

excavationsrecoveredmoreartefactsandalsoidentifiedcut-markedbonesatWestbury

(Andrewsetal.1999).Thesefindswerecomplementedbysubsequentdiscoveriesat

Boxgrove,thatwerealsofoundtobefrominterglacialdepositsearlierthantheHoxnian,

basedontherepresentedfauna,andthereforealsopredatingtheAnglianThames

diversion(andprobablybelongingtoMIS13,~500ka;RobertsandParfitt1999).In

additiontoabout400handaxesthathavebeenexcavatedatBoxgroveandassignedto

thisperiod,bonesoflargefaunawithcutmarkshavealsobeenrecovered,andthese

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encouragedresearcherstore-evaluatetheirassumptionsaboutthehuntingcapabilities

oftheseearlyhumans.Thesubsequentdiscoveryoffossilhumanremains-apartial

tibiaandtwolowerincisors-intheearly1990sheightenedthesignificanceofthe

archaeologicalremains.ProvisionallyassignedtoH.heidelbergensis(Stringeretal.1998;

Stringer2007;Hillsonetal.2010),theseremainsrepresenttheearliestknownfossil

evidenceofhumansinBritain.Scratchesandpitsonthelabialsurfacesoftheincisors

havebeeninterpretedastheresultofdamagebytools,perhapsaspiecesoffood

grippedbetweenthefrontteethwerecutbystonetools.AidedbytheBoxgrove

specimens,a‘ShortChronology’wasdevelopedtodescribethefirsthumanoccupation

inEurope,whichwasrepresentedbysitessuchasBoxgroveandMauerfrom~500ka

(RoebroeksandvanKolfschoten1994).Priortothisdate,theevidencewasconsidered

tobesparseandunconvincing.

However,sincethen,earliersiteshaveemergedthatsuggestsomeearlierforays

intoEurope.TheseincludetheOrcesiteinsouthernSpain,whereafossilhominintooth

hasbeendiscovered,aswellastoolsandfauna,thatdatesto~1.4ma(Toro-Moyanoet

al.2013)andatPirroNord,whichisofasimilarage(Arzarelloetal.2012).

Furthermore,innorthernSpainatGranDolinafossilhumanremainsdatedto~850ka

havebeenexcavated(BermúdezdeCastroetal.2004;BermúdezdeCastroand

Martinón-Torres2014)andassignedtoanewtaxonHomoantecessor(Carbonelletal.

2005;Lacruzetal.2013).Similarly,anevenolderAtapuercasite,SimadelElefante,has

yieldedfragmentaryhumanremains,flakeandcoretools,andfaunawithevidenceof

butchery.SimadelElefantehasbeendatedtobetween1.1and1.2ma(Carbonelletal.

2008).Together,theevidenceatGranDolinaandSimadelElefantehasledresearchers

toarguethatthesehumansrepresentearlyEurasianlineagesthatmayhavegone

extinct.Increasingly,archaeologicalevidenceincombinationwiththeSimadelElefante

materialsuggeststhatthefirsthomininsmayhaveenteredsouthernEuropesoonafter

thehominindispersalrepresentedbytheDmanisifossilsinGeorgia,andthishas

consequentlyfalsifiedthepreviouslypopularhypothesisofaShortChronologyin

Europeanoccupation.

Forawhile,someresearcherssupportedaShortChronologynorthoftheAlps,

giventheabsenceofconvincingevidenceforhumanoccupationbefore500ka,ifMauer

andBoxgrovecouldbedatedtothattimeperiod(Roebroeks2001).However,aflint

handaxewasdiscoveredontheHappisburghforeshoreinNorfolk,in2001,whichis

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26

datedtoatleastanMIS13age(Ashtonetal.2010).ReanalysisofaHappisburgh

assemblagefroma19thcenturycollectionledtothediscoveryofabisonfootbonewith

cutmarks(Parfittetal.2005),whichrepresentsthefirststrongevidenceofhuman

occupationfromaCromerianlocalityinEastAnglia(Parfittetal.2005).Subsequently,

systematicatthesiteofHappisburgh1haveyieldedover50artefactsandseverallarge

faunalspecimenswithevidenceofimpactdamageandcutmarks.

AsiteontheshorelineatPakefield,inSuffolk,hadbeenyielding‘Cromerian’

fossilssincethe19thcentury.Then,human-modifiedflintwasalsorecovered,which

promptedformalexcavationsin2004,ultimatelyproducingover40flintflakeandcore

artefacts(Parfittetal.2005).Provisionally,Pakefieldhasbeenassignedtothewarm

peakofaninterglacialcorrelatedwithMIS17(~700ka),pushingbackhuman

occupationinBritaintotheearlyMiddlePleistocene(Fig.6).

In2006,furtherexcavationsatanewHappisburghlocality,site3,uncovered

signsofanevenearlieroccupationofBritain,lyingbetween~850and950ka,basedon

biostratigraphyandpalaeomagnetism,withevidenceoflessfavourablepalaeoclimatic

signals.Modifiedflintintheformofscrapers,notches,andflakeswasdiscovered,and

analysesofthefaunaandfloraofthesiteindicatedthatHappisburghatthistimewould

havebeenattheborderoftheborealzone(Parfittetal.2010).In2013,correlated

depositsatHappisburghrevealedevidenceofmorethan40humanfootprintsmadein

rivermudsofwhatwasthentheThamesRiversystem(Ashtonetal.2014).

Unfortunately,therearecurrentlynohumanfossilsassociatedwitheitherthePakefield

orHappisburgh3deposits,andthereforetheidentityoftheseearlynorthernEuropean

inhabitantsisstillunknown.Generally,ithasbeensupposedthatthehominins

representedatHappisburgh3andPakefieldwerederivedfromnorthernwavesof

migrationfromsouthernEurope,andthereforelikelylinkedtoH.antecessor.However,

H.antecessoriscurrentlyonlyknownfromtheGranDolinalocality,anditisunclearhow

itandH.heidelbergensisrelatephylogeneticallytothesharedancestryofNeanderthals

andH.sapiens(Stringer2016).

TheAHOBprojectshaveansweredmanyquestionsconcerningtheearliest

humanoccupationsofBritain,andsuggestthatBritainwascolonisedatleast9timesby

successivehumanpopulationsduringthePleistocene(Fig.6).Butnewquestionshave

alsoarisen,inparticularwiththediscoverythattheearliestknownoccupantsat

Happisburgh,atmorethan800ka,werealreadycopingwithclimaticconditions

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somewhatcoolerthanourpresentinterglacial.Thissuggeststhattheymusthavehad

eitherbetterculturalmeanstocopewiththecoldthancurrentlyrecognised,orthatthey

hadevolvedphysiologicaladaptationstocopewiththecold,orperhapsboth.Thisraises

thepossibilitythathumantolerancestocoldintheLowerPalaeolithicweresufficientto

havecolonisednorthernregionselsewhereinEurasia,atleastduringtheinterglacial

periods.PerhapsamultidisciplinaryconsortiumapproachlikethatofAHOBcould

revealevidenceofsuchearlyoccupationsinRussiatoo.

CONSERVATIONPALAEOBIOLOGY

Inadditiontoprovidingtheprimarysourceofdataonthehistoricalpatternof

humanevolution,fossilandarchaeologicalrecordsfromtheNeogene,andespeciallythe

Quaternary,provideuniqueinsightsintothecompositionofpastecosystemsandpast

humaninteractionswithbiodiversity.Humansarenowadominantdriverofpatternsin

globalbiodiversity;well-documentedongoinganthropogenictransformationofthe

biosphereisresponsibleforcatastrophicdeclinesacrossabroadrangeoftaxaand

disruptiontothestructureandfunctioningofecosystems,anditiswidelyacceptedthat

weareexperiencingahuman-mediatedglobalbiodiversitycrisis(McClellan2014;

McGilletal.2015).ForRussianecosystemsalone,recenthistoricalrecordsprovide

evidenceforthehuman-causedglobalextinctionofSteller’sseacow(Hydrodamalis

gigas),Japanesesealion(Zalophusjaponicus),spectacledcormorant(Phalacrocorax

perspicillatus)andcrestedshelduck(Tadornacristata)intheRussianFarEast,the

regionallossormajorrangereductionofawidevarietyofspeciesincludingtarpanor

wildhorse(Equusferus),saiga(Saigatatarica),sociablelapwing(Vanellusgregarius)

andslender-billedcurlew(Numeniustenuirostris),andextensiveanthropogenic

modificationtoecosystemsacrossthecountry(Josephsonetal.2013;IUCN2016).

However,theQuaternaryrecorddemonstratesthathumanactivitieshavesubstantially

affectedspeciesandecosystemsthroughamuchlongerperiodofrecentprehistory.Asa

result,insightsintokeypropertiesofbiodiversity—forexample,ecosystemstructure

andcompositionintheabsenceofhumanmodification,orthespecificecological

requirementsofthreatenedspeciesnowrestrictedtoremnantdistributionsin

potentiallysuboptimalhabitat—willremainincompleteandbiasedbyan“extinction

filter”ifonlyassessedusingmodern-daydata(Balmford1996).Thereistherefore

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increasingawarenessoftheneedforanewdisciplineof‘conservationpalaeobiology’

thatintegratesarangeoflong-termarchivesintoconservationresearchand

management,toidentifypastenvironmentalbaselinesandprovidenovelinsightsinto

regionalbiodiversity,extinctiondynamicsandecosystempropertiesthatare

unavailablefromshort-termstudies(Bonebrakeetal.2010;RickandLockwood2013;

Daviesetal.2014;Barnoskyetal.2017).

Thereisstrongevidencethatearlyhumanscausedtheextinctionofmanylarge

mammals.DuringtheLatePleistocene,almosttwo-thirdsofterrestrialmegafaunal

vertebratetaxa(atleast97genera),mostlymammals,disappearedwithoutecological

replacementfromtheworld’scontinentsduringaseriesof“eco-catastrophic”(Haynes

2002)events,andwithverylittlecorrespondingextinctionofsmall-bodiedspecies

(Martin1984;KochandBarnosky2006;Stuart2015).Someformofhumaninvolvement

inLatePleistocenemegafaunalextinctiondynamicsisnowwidelyacceptedbymost

palaeontologists,asthestepwisenatureoftheseextinctionsacrossdifferentcontinents

correlateswiththearrivaloftechnologicallymodernhumansineachregion.However,

themechanismbywhichhumansmayhavedriventhesemegafaunalspecieslossesis

unclear(e.g.rapiddirectoverhunting,orecosystem-levelinteractionssuchashabitat

modification).Extinctionsalsooccurredduringanintervalofmajorglobal-scaleclimatic

shiftsduringthetransitionfromglacialtointerglacialconditionsnearthePleistocene–

Holoceneboundary,sothattherelativeimportanceorpossibleinteractionsof

prehistorichumanactivityandnaturalenvironmentalchangeindrivingthisextinction

eventhavebeendebatedextensivelysincethenineteenthcentury(Grayson1984).

Thediversity,evolution,ecology,andextinctionchronologyanddynamicsofthe

LatePleistocenemegafaunaofRussiahavebeenthesubjectofextensiveresearch,

supportedbyexcellentpreservationofabundantLateQuaternarymaterialundercool

borealenvironmentalconditions(e.g.inpermafrost).Thisresearchhasbeenconducted

inpartthroughconsiderablecollaborationbetweenRussianandUKacademics(e.g.

ListerandSher2001;Stuartetal.2004,2005;Barnesetal.2007).Currentradiometric

dataindicatethatseveralrepresentativesofthenorthernEurasianmegafauna,including

woollymammoth(Mammuthusprimigenius),woollyrhinoceros(Coelodonta

antiquitatis)andcavelion(Pantheraspelaea),disappearedfrommainlandRussian

ecosystemsduringtheLateGlacialbetweenc.11,000–14,000yearsago(ya),with

severalotherlargemammalsalsobecomingregionallyextirpatedduringthisinterval

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(MacPheeetal.2002;Stuart2015).Megafaunaldisappearanceisassociatedwiththe

disappearanceofvastareasofhigh-productivityopengrass/forb/sedge-dominated

vegetation,representinganecosystemwithnoexactmodernanalogueknownas

‘mammothsteppe’or‘steppe-tundra’,anditsreplacementwithwetmossytundra,

shrubs,andconiferousanddeciduousforest(Zimovetal.2012).

Thisvegetationbiomechangeisofteninterpretedashavingoccurredinresponse

toincreasingtemperaturesattheendoftheLastGlacial,andmayhavebeenresponsible

fordrivingtheextinctionofthelargemammalcommunitythatwasadaptedfor

mammoth-steppelandscapes(e.g.Stuart2015).However,ithasalsobeenarguedthat

thisecosystemshiftmayrepresentanexampleoftop-downratherthanbottom-up

regulation,withthemammoth-steppeecosystemactuallymaintainedby

megaherbivoresanddisappearingwhenthesekeystonespeciesbecameextinctdueto

humanactivity.Heavymammaliancroppingandtramplingwouldlikelyhave

suppressedwoodygrowthandstimulatedgrowthofgrasses,leadingtohighratesof

transpirationofsoilmoistureandlikelyincreasedcarbonstorageindrysoilsand

permafrost,andrapidnutrientcyclingfrommegaherbivoredungsupportedhigh

ecosystemproductivity;conversely,megaherbivoreextinctionwouldhaveallowed

shrubsandtreestoexcludegrasses,risingwatertableswouldhavefurthersuppressed

grassesandfavouredcool,waterloggedmoss-coveredsoils,anddecreasednutrient

cyclingwouldhaveledtodecreasedsoilfertility(Zimov2005;Zimovetal.2012).This

novelhypothesissuggeststhatmammoth-steppemayrepresentastableecosystemthat

canpotentiallyberecreatedinRussiaundercurrent-dayenvironmentalconditionsif

largeherbivoresarepresent.

Investigatingthispossibilityisnowthefocusofthe‘PleistocenePark’projectled

bySergeiZimov,whichhasreintroducedextantlargemammalsformerlypresentduring

theLatePleistocene(includingferalhorses,muskoxenOvibosmoschatus,andEuropean

bisonBisonbonasus)toanaturereserveontheKolymaRiverinSakhaRepublic,totest

predictionsaboutecosystemchangeandpotentialreductioningreenhousegas

emissionsassociatedwithpredictedreductioninpermafrostthawing,andultimately

aimingtorestoreregionalecosystemprocessesthatmaybe‘natural’throughout

Quaternaryglacial–interglacialcycles(Zimov2005).Comparablebutsmaller-scale

rewildingprojects(e.g.,atOostvaardersplassenintheNetherlands;Lorimerand

Driessen2014)arealsounderwayelsewheretoassesschangesinecosystemstructure

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30

andproductivityassociatedwithreintroductionofregionallyextirpatedlarge

herbivores,andthereisconsiderablepotentialforfutureresearchinthisareatobetter

understandextinctiondynamicsandecosystemstability.

Extensiveevidenceforfurtherhumaninvolvementinprehistoricbiodiversity

lossisdocumentedacrossthesubsequentHoloceneEpoch.Thiswasanintervalof

modestorminimalclimaticvariationunderbroadly“modern”environmentaland

climaticboundaryconditions;evidenceforhumaninvolvementinnearlyallHolocene

extinctionsandecosystemchangeisnotconfoundedbymajorclimaticfactorsandis

thereforerelativelyundisputed(Turvey2009).MostHoloceneextinctionsare

documentedfromislandsystems,includingthewell-documentedextinctionofRussia’s

finalremnantmammothpopulationonWrangelIslandc.4000ya(Vartanyanetal.

1993;Stuartetal.2004).Continentalregionsexperiencedreducedlevelsofpostglacial

species-levelextinctionbeforetherecenthistoricalera,leadingtouseoftheterm

‘Holoceneunderkill’tocontrastwithhypothesized‘Pleistoceneoverkill’(Grayson2008).

However,itisincreasinglyapparentthatcontinentalecosystemsalsounderwent

considerableanthropogenicmodificationthroughouttheHolocene,associatedwith

extensiveregionalspecieslossesandpopulationdepressions(Grayson2001;Johnson

2006;CreesandTurvey2014).Reconstructingthemagnitudeandnatureofprehistoric

human-causedbiodiversitychangeunderpostglacial,‘modern’environmental

conditionsisnecessarytoidentifyregionallyextirpatedspeciesforpotential

reintroductionprojects,andthetrueevolutionaryorecologicalbasisoftheirgeographic

distributionsbeforedisruptionbyhumanactivity.Suchreconstructionwouldalso

providewidernovelinsightsintoimportantconcernsinmodernconservationbiology,

suchasthedeterminantsofvariationinpopulationvulnerabilityorresiliencethrough

timeacrossdifferentspeciesandlandscapes.

RussianecosystemsexperiencedfurtherpostglacialHolocenespecieslosses

beforetherecenthistoricalera.Theseinclude(Fig.7)globalextinctionsofthefinal

remnantpopulationsoftwoPleistocenesurvivors,thesteppebisonBisonpriscus

(persisteduntil9,800yainTaimyrand8,900yainwesternChukotka;MacPheeetal.

2002;Kirillovaetal.2013)andthegiantdeerMegalocerosgiganteus(persisteduntil

7,700yaintheUrals;Stuartetal.2004),andthemorerecentdisappearanceofaurochs

(Bosprimigenius)inthelateHolocene(Creesetal.2016).TheHolocenerecordalso

showsthatmanystill-extantspecieshadmuchbroaderpostglacialgeographical

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31

distributionsinRussia.Forexample,allwildpopulationsofEuropeanbisonwere

extirpatedbytheearly20thcentury,butthespeciesformerlyoccurredacrossalarge

areaofEuropeanRussiaasfarnorthasLakeLadoga(Sipko2009)(Fig.7);and

archaeologicalremainsdemonstratethatmultiplesturgeonspecies(Acipensersturio,A.

oxyrinchus)occurredintheNorthSeaduringtheHolocene(Ludwigetal.2002;Thieren

etal.2016).Alargenumberofmammalandbirdspeciesalonearenowknowntohave

becomeregionallyextirpatedelsewhereinEurope,e.g.Britain,duringtheHolocene

(YaldenandAlbarella2009;O'ConnorandSykes2010),anditislikelythatfuture

researchintopostglaciallong-termarchiveswillrevealgreaterlevelsofhuman-caused

biodiversitylossfrom“modern”Russianecosystems.

DatafromHolocenelong-termarchiveshavesofarbeenusedtoguidespecies

recoveryandenvironmentalrestorationprogrammes,suchasthereintroductionof

EuropeanbisontolandscapeswithinitsformerdistributioninRussiaandelsewherein

Europe(Sipko2009;IUCN2016),andtheproposedreintroductionofCaspiantiger

(Pantheratigrisvirgata)tositesinCentralAsia(Chestinetal.2017).However,wide-

scalemacroecologicalanalysisofpatternsofmammalianrangelossacrossEurope,

includingdatafromRussia,hasrevealedthatprehistorichumanimpactsonpostglacial

biodiversitybeganearlyonduringtheHolocene,andthatthedynamicpatternof

progressivefaunalmodificationofEuropeanmammalassemblagesacrosstheHolocene

challengeseasyidentificationof‘static’pastbaselinesthatcouldactasbenchmarksfor

current-dayenvironmentalmanagementandrestoration(Creesetal.2016).

Furthermore,differentterrestrialvertebrateguildshavebeendisruptedtodifferent

extentsbyhumanactivitythroughtheHolocene(e.g.large-bodiedmammaliangrazers

andbrowsersexperiencedsignificantlyearlierdeclinesthanmammaliancarnivores;

Creesetal.2016).Itisagainlikelythatreductionorremovalofdistinctfaunalguilds

duringtheHolocenewouldhavetriggeredwiderecologicalknock-oneffects,suchas

shiftsinplantcommunitycompositionfollowingbothremovaloflargeherbivores(cf.

Zimovetal.2012),andremovaloflargecarnivoresandthe‘landscapesoffear’thatthey

maintain(Kuijperetal.2013);however,thepotentialconsequencesofthisHolocene

lossoffunctionaldiversityandpotentialkeystonespecieshasnotbeenadequately

studiedinRussianecosystems.

Incorporatingdatafromlong-termarchivesisessentialforenvironmental

management,butconservationpalaeobiologyraisesimportantandchallenging

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32

questions.Ultimately,isitevenpossibletoidentifyordefinewhatconstitutesa‘natural’

Russianecosystem,whetheranysuchsystemsstilloccurtoday—andinthefaceof

conflictingpressuresofhumanresourceuseanddependencyonhigh-productivity

landscapes,could,orshould,anyRussianecosystemsberestoredtoincorporatenow-

lostbiodiversityorecologicalprocesses?

Acknowledgements:DEGBisgratefulforcommentsandinformationfromM.J.Benton,

N.J.Butterfield,M.E.McNamara,J.Vinther,andP.R.Wilby.S.Buttsassistedwiththe

figure.

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Fig.1.SirRoderickMurchison(1792–1871),aboutthetimehevisitedRussiain1840,

andthetitlepageofhismajormonographonRussiangeology,publishedinLondon,

Paris,andStPetersburgin1845.

Fig.2.Ediacarafossils.(a)Dickinsoniasp.,ZimniiGory,WhiteSea,Russia(photoJ.

Vinther,seeFedonkinetal.2007,fig.222).(b)Charniamasoni,CharnwoodForest,

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51

Leicestershire,UK(theholotype,copyrightBritishGeologicalSurvey,Natural

EnvironmentResearchCouncil).

Fig.3.CryptosporesfromtheUpperOrdovicianofSiberia,evidenceforsomeofthe

earliestvascularplants.(a,b)Permanenttetradenclosedinalaevigateenvelope,

?VelatitetraslaevigataBurgess,1991.(c)Pseudodyadenclosedinalaevigateenvelope,

SegestresporalaevigataBurgess,1991.

Fig. 4. The cladogram shows the relationships as a present understood between some of the

Devonian tetrapods. The stratigraphical column at the right shows the Russian taxa in red in

the context of a selection of other taxa and their countries of origin. Russian taxa contribute

key parts of this assemblage both phylogenetically and stratigraphically.

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Figure 5. The Permo-Triassic boundary in Russia. (a) The Kulchomovskaya Svita (latest

Permian) below the ledge, and the Kopanskaya Svita (basalmost Triassic) above, in the

Korolki Ravine, near Sol-Iletsk, on the south-western margin of the Urals, Asiatic Russia.

Mikhail Surkov, M.J.B. and Valentin Tverdokhlebov inspect the sandstone lying right at the

boundary. (b) Graeme Taylor drills rock plugs for magnetostratigraphic analysis in the

Boyevaya Gora PTB section, South Urals, assisted by M.J.B. (c) Overview of the fates of

tetrapods from the Middle Permian to Middle Triassic, showing that only prcocolophonids

and dicynodonts survived the Permo-Triassic mass extinction.

(a) (b)

(c)

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Figure 6. A timeline of key sites and events in Britain’s human past. The climate curve is a

combination of deep-sea records, and corresponds to global ice volume. This gives an

indication of changing climatic conditions through time. Copyright The Trustees of the

Natural History Museum, London.

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Figure7.ReconstructedHolocenerangesacrosseightarchaeologicalperiodsfortwo

largemammalspeciesnowextirpatedfromRussia(top:EuropeanbisonBisonbonasus;

bottom:aurochsBosprimigenius),showingminimumconvexpolygonsofavailable

localityrecordsfromarchaeologicalsites.Rangeextentisartefactuallybiasedby

variationinnumberandspatialdistributionofarchaeologicalsitesacrossdifferent

periods(i.e.,samplingbias),butavailabledatademonstratethepersistenceofboth

speciesinRussiaintotheLateMedieval.ModifiedfromCreesetal.(2016).

Bison bonasus

Bos primigenius

All zooarchaeological

records

Mesolithic Neolithic Bronze Age Age

Iron Age Roman Age