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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 inPaleontology: Past, Present, and Future. Paleontological Journal, 51(6), 576-599. https://doi.org/10.1134/S0031030117060028
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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).
12
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
13
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
14
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
15
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
16
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
17
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
18
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
19
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.
20
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.
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
22
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
23
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
24
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
25
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
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
27
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
28
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
29
(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
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
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
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,
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Paris,andStPetersburgin1845.
Fig.2.Ediacarafossils.(a)Dickinsoniasp.,ZimniiGory,WhiteSea,Russia(photoJ.
Vinther,seeFedonkinetal.2007,fig.222).(b)Charniamasoni,CharnwoodForest,
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.
52
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)
53
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.
54
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