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LifeinthePaleozolic

Introduction

ThePalaeozoicErawasaveryimportanttimeinthehistoryoflife.Using
evidencefromfossils,westartbylookingattheCambrianexplosion,whenmany
formsofanimallifefirstappearedabout545millionyearsago.Thenwemove
ontostudycreatureslivingintheOrdovicianseas,includingtheextinct
trilobites.Next,we'llinvestigatetheinvasionoflandbyplantsandinvertebrates
thatoccurredintheSilurianPeriod,andlookatlifeinSilurianseas.You'llalso
learnabouttheDevonianPeriod,whenvertebratesfirstmovedontoland.The
unitfinisheswithabriefoutlineofvertebrateevolution.

ThisunitismodifiedfromtheOpenUniversityLevel1courseS193;Fossilsand
theHistoryofLife.Forthispartofthatcourse,studentsarealsogiventwo
replicasofPalaeozoicfossilstodeveloptheskillofmakingandrecording
observationsonthreedimensionalspecimens.Referencetostudyingthese
replicasisomittedfromhere.

LearningOutcomes

Afterstudyingthisunityoushouldbeableto:

describesomekeyeventsintheevolutionoflifeduringthePalaeozoic
Era,suchasthefirstappearanceofmajorgroupsofinvertebratesand
vertebrates,andtheinvasionoftheland;

identifysomecommontypesoffossilorganismsthatwerelivingin
Palaeozoicseas,andcommentontheirlikelyenvironmentandgeological
age;

makeinferencesfromfossilsaboutthebiologyandmodeoflifeofsome
Palaeozoicorganisms.

1TheCambrianexplosion

1.1Aburstofevolution
Oneofthemostimportanteventsinthehistoryoflifebeganabout545million
years(Ma)ago,i.e.somefourbillionyearsaftertheoriginoftheEarth,andover
3.3billionyearsaftertheoriginoflife.ThetermCambrianexplosionreflects
asuddenburstofevolution,whenawidevarietyoforganisms,especiallythose
withhard,mineralisedparts,firstappearinthefossilrecord.Thusbeganthe
PhanerozoicEonthetimeofvisiblelifeandthePalaeozoicEra.

AspartofthisunityouwilloftenbereferredtoapdffiletakenfromDouglas
Palmer'sAtlasofthePrehistoricWorld.

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AtthestartoftheCambrianPeriod,bodyfossilswithhardshellypartsbecame
abundantforthefirsttime.Actually,theveryfirstevidenceofpreservablehard
partscomesfromsometinyfossils,includingtubesmadeofcalciumcarbonate
about0.2mmacross,foundinlatePrecambrianstrataabout550Maold.The
unknownorganismsthatproducedthemwerelivingatthesametimeasthe
Ediacaranfauna.

What,ingeneral,wasdifferentaboutthesizeoffossilswithhardshellypartsin
theearliestCambrian,comparedwithlatePrecambrianlifesuchastheEdiacaran
organisms?

Nowreadtheanswer

Awidevarietyofsmall(12mm)shellyfossilsappearedintheearliestpartof
theCambrianPeriodassortedshapessuchastubesandcones,aswellasspines,
scales,andknobs(Atlas,pp.5859).It'softendifficulttotellwhetherafossilis
thecompleteskeletonofasingleorganismoranisolatedpartofsomelarger
creature.Thesofttissuesassociatedwiththesehardpartsarealmostentirely
unknown,andthereconstructionsofsoftpartsshownintheAtlas(pp.6061)are
conjectural.

Whatseemslikelytohavebeenthemainstimulusfortheacquisitionofhard
partsbyorganismsintheearliestCambrian?

Nowreadtheanswer

Protectionfrompredationorotherdamageiscertainlyconsistentwiththefirst
appearanceofdefensivestructuressuchasspinesandscales,andtubesand
conicalshellsthatcouldprotectvulnerablesoftpartsinside.Thereare,however,
anumberofotherpossiblereasonswhyhardpartsfirstevolved,asdiscussedin
theAtlas,p.61,someorallofwhichmayhavebeeninvolved.

Agreatlyincreasedvarietyoftypesoftracefossils,especiallyburrowsofsoft
bodiedanimals,arefoundaroundthestartoftheCambrianPeriod.Whatisthe
significanceofthisfinding?

Nowreadtheanswer

Althoughenvironmentalchangeswereoccurring(suchasariseinglobalsea
level(Atlas,p.58),thereis,todate,littleevidenceofspecial,widespread
environmentalchangesthatcouldhavedirectlytriggeredtheCambrian
explosion.Whateverthecauses,oncetriggered,awiderangeofecological
opportunitiespresumablybecameavailableforexploitation,promotingtherapid
evolutionofnew,quitedifferenttypesofanimals.Byabout530Ma,mostofthe
animalphylathatareinexistencetodayhadappeared.Notsurprisingly,afew
entirelysoftbodiedphylalivingtodayhavenoknownfossilrecord,sowedon't
knowwhentheyevolved.Evidencefromgeneticssuggeststhatsomeanimal
phylahaddivergedfromeachothermuchearlierthanthestartoftheCambrian
explosion,butthetimingremainsuncertain.

ManyofthenewlyevolvedCambrianphylashoworganisationofthebodyinto
specialisedareasespeciallyaheadendwithfoodtrappingandsensoryorgans,
atubulargutandlimbs.ThereisnodoubtthatmanyCambriananimalswere
equippedwithadaptationsforpreyingonotheranimals,andwereabletopursue
foodmuchmoreactivelythancouldtheEdiacaranfaunasuchasbyscuttling
overtheseafloor,swimmingactively,andburrowing.Notethatallformsoflife
(exceptpossiblyafewalgae)wereasyetconfinedtothesea.

Althoughmanytypesofthesmallshellyfossilsdisappearfromthefossilrecord
soonafterthestartoftheCambrian,somearethoughttohavebeenmolluscsat
anearlyevolutionarystage,inwhichcasetheydidleavedescendants.

1.2TheBurgessShale

HighintheCanadianRockiesisexposedadepositofmiddleCambrianage,
about530Maold,calledtheBurgessShale.Itcontainsthefossilsofanimalsthat
livedonamuddyseafloor,andwhichweresuddenlytransportedintodeeper,
oxygenpoorwaterbysubmarinelandslides.Theircatastrophicburialhasgiven
usanexceptionalviewofCambrianlife.Notonlyhaveanimalswithhardshelly
partsbeenpreservedbutentirelysoftbodiedformsarealsopreservedasthin
filmsonthesedimentsurface.

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SomeofthemostcommonCambrianfossils,whichappearimmediatelyafterthe
firstshellyfossils,aretrilobites.Thesewereagroupofexclusivelymarine
arthropods,membersoftheenormouslydiversephylumofanimalswithjointed,
externalskeletonsthattodayincludeformssuchascrabs,lobsters,insectsand
spiders.ThetrilobitefossilsoftheBurgessShalearelikemanytrilobitesfound
elsewherebutexceptionalinthatnotonlyisthemainpartoftheirouterskeleton
(orexoskeleton)preserved,butsotooaretheirappendagessuchasantennaeand
legs(see,forexample,thoseofOlenoides,Atlas,p.63).Elsewhere,trilobite
appendagesareextremelyrareastheywerepoorlymineralised.Wewillstudy
trilobitesinmoredetailshortly.Othertypesofarthropods,especiallyones
lackingwellmineralisedexoskeletons(suchasMarrella,Atlas,p.64),are
particularlyabundantintheBurgessShale.

Onlyabout15percentofthe120generapresentintheBurgessShaleareshelly
organismssuchastrilobitesandbrachiopodsthatdominatetypicalCambrian
fossilassemblages(fossilsthatoccurtogether)elsewhere.Theshellycomponent
wasthereforeinaminority,andorganismswithhardpartsprobablyformedless
than5percentofindividualsinthelivingcommunity.

IfthesoftbodiedfossilsoftheBurgessShalearetakenaway,allthatremainsis
atypicalCambrianassemblageofhardbodiedorganisms.Whyisthisimportant
tobearinmindwhentryingtointerpretotherCambrianfossilassemblages?

Nowreadtheanswer

AnotherimportantrevelationoftheBurgessShaleliesinthewidediversityof
animaltypesthatwerearoundinmiddleCambriantime,about530Maago.
Therearerepresentativesofaboutadozenofthephylathatpersisttothepresent
day.OneformcloselyrelatedtoearlyarthropodswasAnomalocaris,thelargest
knownCambriananimal,someindividualsofwhichmayhavereachedtwo
metresinlength.Itsextraordinaryjaw(Atlas,p.65)consistedofspinyplates
encirclingthemouth,whichprobablyconstricteddownonpreyinmuchthesame
waythattheplatesofanirisdiaphragmcutdownthelightinacamera.This
fearsomemouthisseeninplaceinthereconstructionofthecloselyrelated
Laggania(Atlas,pp.6465).Notethatthecoloursoforganismsshowninthis
andothersuchreconstructionsareconjectural.

AboutadozentypesofBurgessShalefossilshavebeensaidtobesounlike
anythinglivingtodayandsodifferentfromeachotherthat,hadtheybeenliving
now,eachwouldhavebeenplacedinaseparatephylum.Withfurtherstudy,
however,therelationshipsofthesepuzzlinganimals(suchasHallucigenia,
Atlas,p.63)arebecomingclearer.ItseemsthatsomeBurgessShaleformsare
hardtoclassifysimplybecausetheboundariesbetweenmajorcategoriesof
animallifewerestillblurredshortlyaftertheCambrianexplosion.Inother
words,bymidCambriantime,therestillhadnotbeenenoughtimeforsome
groupstohavedivergedsufficientlyfromtheirrecentcommonancestorstobe
distinctlydifferent.

BurgessShaletypefaunashavebeenfoundinabout30sitesrangingfromNorth
AmericaandGreenland,toChinaandAustralia.Thewiderangeofanimalsthey
containseemstoreflectanunprunedbushofdiversityresultingfromthe
Cambrianexplosion.Notlongafter,though,extinctionloppedoffsomeofthe
branches,leavingphylawiththerelativelydistinctfeaturesthathaveremainedto
thisday.

1.3Anoverviewofanimalphyla

Wehavealreadymetquiteafewdifferentanimalphyla,andit'susefultogetan
overviewofalltheonescommonlyfoundinthefossilrecordandtheirmodeof
lifebeforestudyingsomeinmoredetail.Exceptforafewsoftbodiedphylawith
verypoorfossilrecords,itisclearthatalltheanimalphylahadappearedbythe
OrdovicianPeriod.

Modesoflife

Thefollowingtermsareveryusefulfordescribingthemodeoflifeandthe
environmentalsettingoforganisms.Thetermsareexplainedhereastheyare
appliedtomarineorganisms,buttheyarealsosometimesappliedtoorganisms
livinginlakes.

benthicanimalsandplantsthatliveontheseafloor;collectivenoun:benthos.

pelagicanimalsandplantsthatliveabovetheseafloor.Theymaybeeither
nektonic(animalsonly)orplanktonic(animalsandplants).

nektonicanimalsthatswimactively(e.g.fish,squid);collectivenoun:nekton.

planktonicanimalsandplantsthatdriftpassivelyorswimfeebly,mainlyin
thesurfacewatersofseas;collectivenoun:plankton.Thetermincludes
phytoplankton(photosyntheticorganisms,mostlyalgae)andzooplankton
(mostlymicroscopicanimals,includinglarvaeoflargerones,butalsosome
macroscopicanimalsthatarereadilyvisiblewiththenakedeye,e.g.jellyfish).

epifaunalanimalsthatliveontheseafloor,eitheronsoftsediment,orattached
torocks,seaweed,etc.(sessile),orthatmoveovertheseafloor(vagrant);
collectivenoun:epifauna.
infaunalanimalsthatlivewithinsediment,ofteninburrowsorboringsinto
hardermaterial;collectivenouninfauna.

Likemostclassificationsinvolvinglivingorganisms,someinvertebratesdonot
fitneatlyintothesecategories,e.g.thecommonprawnburiesitselfinsediment
duringtheday(i.e.isinfaunal),butatnightemergestojointheepifaunaasit
feeds.Otherepifaunalanimalsburythemselvesduringlowtide.

Thefollowingtextgivessomekeypointsabouttheimportantanimalphylamost
commonlyfoundinthefossilrecord.Theagerangeandmodeoflifeofsome
commongroupsaregiven,andphylathataremicroscopicthroughoutlifeare
excluded.Figure1showstypicalfossilrepresentativesofsomeofthephyla.

Porifera.Sponges.CambriantoRecent.Mainlymarine;somefreshwater.
Sessile.Thesimplestmulticellularanimals,spongeslackdefinitetissuesand
organs,e.g.theyhavenonervoussystem.Theyhaveaskeletonofcalcium
carbonate,silica,or,asinsomebathsponges,hornyorganicmaterial.Water
passesinthroughthesponge'smanysurfacepores,oftentothecentralcavityofa
sacklikebody,andoutthroughalargeholeatthetop.Somehaveastalk(Figure
1a),othersareencrustingandirregularinshape.Spongesfeedbyfilteringoff
minuteorganicparticlesfromthewater.Spongesarelocallyabundantfossils,
especiallyinCretaceousrocks,wheretheyareverycommonlyenclosedinflint
nodulesintheChalk.

Cnidaria(pronouncedwithasilentC:nighdareeea).LatePrecambrian
(Ediacaran)toRecent.Almostentirelymarine.Cnidarianshaveacentralmouth
aroundwhicharestingingtentaclesforcatchingprey.Byfarthemostimportant
fossilgrouparetheentirelymarine,generallybenthic,corals,whichsecretea
skeletonofcalciumcarbonatebelowthesoft,anemonelikepartsatthetop.
Coralsmaybeeithercolonial(withmanygeneticallyidentical,linkedindividuals
sharingaskeleton)orsolitaryindividuals.Therearethreemaingroupsofcorals:
rugosecorals(solitary,Figure1b,orcolonial),OrdoviciantoPermian;tabulate
corals(alwayscolonial),OrdoviciantoPermian;andscleractiniancorals,
sometimesalsocalledhexacorals(solitaryorcolonial),TriassictoRecent.Sea
anemonesandjellyfisharealsocnidarians,butbeingsoftbodiedthesegroups
aremuchlesscommoninthefossilrecord.

Bryozoa.Bryozoans.OrdoviciantoRecent.Sometimescalledmossanimals.
Normallymarine,rarelyfreshwater.Sessile,tinyanimalswhichliveincolonies,
withaskeletonusuallyofcalciumcarbonate.Thecoloniesvaryinshape;some
areencrustingsheets(seamats),othersmaybedelicatenetlikefronds(Figure
1c)orbranchingtwigs.Eachcolonyconsistsofafewtothousandsof
interconnectedindividuals.Whenfeeding,tentaclesfiltermicroorganismsfrom
thewater.Bryozoansoftenoccurasfossilsamongthediversefaunaofreefs.
Somebryozoanslookratherlikesmallcorals;afewspeciescanlookalittlelike
graptolites(seebelow).

Brachiopoda.Brachiopods(pronouncedbrackyopods).CambriantoRecent.
Entirelymarine,benthicanimals.Sometimescalledlampshells(aftertheir
resemblancetoRomanoillamps).Brachiopodsaretypically25cmlong,but
theyrangeinsizefromafewmmtoasmuchas30cm.Theshell,whichencloses
thesofttissues,hastwoparts,calledvalves.Onevalveisalmostalwayslarger
thantheother.Manybrachiopodsareattachedtotheseafloorbyastalkofhorn
likeorfleshymaterial,thepedicle;infossilbrachiopodsthepresenceofthis
pedicleisindicatedbyaholepassingthroughthelargerofthetwovalves(Figure
1d).Somearefreelyingontheseafloor,andafewarecementedorattachedby
spines.Inmostbrachiopodstheshelliscomposedofcalciumcarbonate,though
somearephosphatic.Theyfeedbydrawingwaterintotheshellandfilteringoff
foodparticleswithacomplexfeedingdevicecalledalophophore.Brachiopods
aremuchlessabundantanddiversetodaythanduringthePalaeozoicand
Mesozoic.TheyarethecommonestfossilinmanyPalaeozoicshallowmarine
limestonesandshales.About20speciesoccurtodayofftheBritishIsles,mostly
indeeperwaters,andarerarelyseen.

Mollusca.Molluscs.CambriantoRecent.Mainlymarine.Averydiverse
phylum,perhapsnumericallythemostabundantlargeinvertebratesinthefossil
record.Thereareshelledandunshelledforms.Althoughatfirstthesixorso
livingclassesmayseemunrelated,theyrepresentevolutionaryvariationsonthe
samethemethemolluscanbodyplan.Threeclasses,eachofwhichrangefrom
CambriantoRecent,areparticularlyimportant,bothasfossilsandtoday:the
Bivalviabivalves(e.g.cockles,musselsandoysters)(Figure1e);the
Gastropodagastropods(e.g.slugsandsnails);andthenektonicCephalopoda
cephalopods(e.g.squid,cuttlefish,octopusandnautilus).Mostbivalvesare
marine(vagranttosessilebenthos),thoughsomearefreshwater.Themajorityof
gastropodsareaquatic,andmostoftheseliveinshallowseas,buttheyarealso
widespreadinfreshwaterandondryland.Cephalopodsare(andhavebeen)
entirelymarine,andarethemosthighlyevolvedmolluscs.Themostimportant
fossilcephalopodgroupsareallformswithchamberedshellsthenautiloids
(CambriantoRecent),thefamiliarspiralshelledammonites(Triassicto
Cretaceous),andthebulletshapedbelemnites(JurassictoCretaceous).

Echinodermata.Echinoderms.CambriantoRecent.Entirelymarine.Mostare
benthic.Manyarevagrant,somearesessile,andafewarefreeswimming
(nektonic).Mostechinodermskeletonsaremadeofmanyporousplatesofcalcite
(calciumcarbonate)whichareverythinlycoveredwithsofttissue.Multipurpose,
extendibletentaclescalledtubefeetemergetotheoutside,andareused
especiallyinfeeding,respirationandlocomotion.Inmanyformsthereisa
distinctivefiverayedarrangementofplatesandtubefeet.Echinodermmeans
spinyskin,referringtothefactthatsomegroupshavespinesorhard,warty
bumpsprojectingfromthesurface.Themostcommonfossilgroupsaresea
urchins(echinoids),OrdoviciantoRecent,andsealilies(crinoids),Cambrian
toRecent.Twootherwellknownlivinggroups,bothOrdoviciantoRecent,are
starfish(asteroids)andbrittlestars(ophiuroids).Therearealsoseveralextinct
groups.

Arthropoda.Arthropods.CambriantoRecent.Thelargestphylumofanimals,
withagreatdiversityofmorphologyandmodeoflife;livingformsarefoundin
mostpossiblehabitatsinwaterandonland.Apartiallistofgroupsincludes
crustaceans(crabs,lobsters,barnaclesandshrimps),insects,millipedes,
centipedes,spiders,kingcrabs,scorpions,mitesandseveralextinctgroups,of
whichtheentirelymarine,extincttrilobites(pronouncedtrylobites)arethe
mostimportant.Themostcharacteristicfeaturesofthephylumarethehardouter
coating(exoskeleton)whichisdividedintosegments,andthepaired,jointed
appendageswhichvaryinnumberandfunction.Theexoskeleton,usuallyof
chitin(astrong,lightweightorganicmaterial),maybefurtherstrengthenedby
calciumcarbonateorcalciumphosphate,increasingthepreservationpotential.
Growthoccursduringperiodicmoultingwhentheexoskeletonisshedandanew,
largeroneisformed.
Figure1:Representativefossilsofsomeofthephylamentionedabove.(a)A
sponge,Siphoniatulipa,Cretaceous(5.5cm).(b)Acnidarian,thesolitary
rugosecoralClisiophyllum,Carboniferous(3.5cm),(c)Abryozoan,
Fenestellaplebeia,Carboniferous(fieldofview3cm).(d)Abrachiopod,
Terebratulagigantea,Cretaceous(4cm).(e)Amollusc,thebivalve
Pleuromyacostata,Jurassic(4.5cm).(f)Ahemichordate,thegraptolite
Hustedograptusteretiusculus,Ordovician(2.5cm)

Hemichordata*.Hemichordates.CambriantoRecent.Byfarthemostimportant
fossilgrouparethegraptolitesextinct,entirelymarinecoloniesconfinedtothe
PalaeozoicEra,particularlyabundantinOrdovicianandSilurianrockswhere
theyareveryusefulzonefossils.Manylooklikesawbladesafewcentimetres
longontherock,withteethononeorbothsidesofthesaw(Figure1f).The
teethwereactuallytinycupsthathousedindividualswhichmadeupthecolony
andpossessedfilterfeedingtentacles.Somegraptoliteswerebenthicandsessile,
butmostwerepelagic,eitherdriftingorpossiblyswimmingfeeblyaspartofthe
zooplankton.

*SomebiologistsplacethehemichordatesasasubphylumwithinthePhylum
Chordata.

Chordata.Chordates.CambriantoRecent.Allchordatespossessanotochord(a
flexiblerodrunningalongthelengthofthebody).Minorlivingchordategroups,
lackingavertebralcolumn(backbone),includeseasquirtsandthelancelet.The
majorgrouparethevertebrates,whichhavebonyorcartilaginousskeletonsand
ahead.Today,vertebratesincludefiveclasses:fish(CambriantoRecent),
amphibians(DevoniantoRecent),reptiles(CarboniferoustoRecent),
mammals(TriassictoRecent)andbirds(JurassictoRecent).Importantextinct
reptilegroupsincludethelanddwellingdinosaurs(TriassictoCretaceous),the
marineichthyosaurs(TriassictoCretaceous)andplesiosaurs(Jurassicto
Cretaceous),andtheflyingpterosaurs(TriassictoCretaceous).Vertebrate
fossilsarerelativelyrarecomparedwithinvertebrates,andareusuallyonly
fragments.

1.4Theoriginofthevertebrates

Vertebratessuchasourselvesarebydefinitionanimalswithabackbone(or
vertebralcolumn,pairedlimbs,askullandvariousotherstructures.Until
recentlyvertebrateswerethoughttoextendbackonlyintolateOrdoviciantimes,
some450millionyearsago.Atthistimefossilsofstrangelookingfishwith
bonyheadshields,suchasSacabambaspis(Atlas,pp.7071),appearinthefossil
record.Thesejawlessfish(calledagnathans)areonlyverydistantlyrelatedto
thesolelivingagnathans,thelampreysandhagfish.However,theseOrdovician
creaturesarealreadyhighlyevolvedandclearlyhadyetmoreancientancestors.

Tounderstandwhatsuchancestorsmighthavebeenlikeweneedtoconsiderthe
backbonethefundamentalvertebratefeatureinabitmoredetail.The
precursortothebackboneisastiffeningrod,calledthenotochord,detailsof
whichweknowfromthestudyofdevelopingvertebrateembryosandthefew
survivinganimalswhichhaveretainedanotochord,suchasthelancelet
Branchiostoma.

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Whatisthemainfunctionofthenotochord?

Nowreadtheanswer

Thefundamentalchordatecharacteristicofanotochord(Section1.3)hasnow
beenidentifiedinvariousCambrianfossils,suchasPikaiafromtheBurgess
Shale(Section1.2).

WhyisPikaiaclassifiedasachordatebutnotavertebrate?

Nowreadtheanswer

Theextinctconodonts(lateCambriantoendTriassicinage)havebeenshownto
possessnotonlyanotochordbutalsotoothstructuresandpairedeyes,which
seemtosuggestthattheyweremoreadvancedthanchordatessuchasPikaiaand
closetotheearliestvertebrates.(Youwillreadaboutconodontsshortlyonpp.70
oftheAtlas.)SomeearlyCambrianfossilsrecentlyfoundinChinaarethoughtto
preserveallthebasicchordatefeaturesplussomemoreadvancedvertebrate
ones,namelygillsaswellaspairedeyes.Ifso,thenitisclearthatsomeanimals
closetothevertebratebodyplanhadalreadyappearedbyearlyCambriantimes,
andchordateancestryprobablyreachesbackintothelatePrecambrian.

Humansbelongtothevertebrategroupknownasmammals.UsingSection1.3,
placemammalsandtheotherfourmainvertebrategroupsinorderoftheir
evolutionaryappearance.

2TheOrdovicianseas

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CollectingseashellsonanOrdovicianbeachwouldhavebeenarathercurious
experience.Whilstmostshellsweremadeofsimilarmaterialstothosefoundon
amodernbeach,thedetailedformofmanywouldhavebeenquiteunfamiliar,
andallthespecieshavelongbeenextinct.

Havealookatthepanoramicillustrationonpp.7071oftheAtlas.Fromthis
andtheAtlas,pp.6871,thinkaboutwhichorganismsappearmostunfamiliarto
you.

Therearemanyorganismsthatprobablyseemunfamiliar,andwhenreading
aboutthemyoumaywanttorefertoSection1.3.Brachiopods,e.g.Strophomena,
aresuperficiallyclamlikeanimals,andalthoughnotextinct,aremuchrarernow
thaninthePalaeozoic;mostpeoplehaveneverseenalivingone.Graptolites,e.g.
Orthograptus,areextinctcolonialanimalsthatmostlydriftedintheocean
currents(Figure2).Trilobites,e.g.Triarthrus,areextinctmarinearthropods.
Conodonts,e.g.Promissum,areextinct,andjawless(agnathan)fish,e.g.
Sacabambaspis,areextinctexceptforhagfishandlampreys(seeSection1.4).
Thestraightshellednautiloidcephalopods,e.g.Endoceras,areextinctandonly
distantlyrelatedtotoday'sNautilus,whichhasaspiralshell.Thegroupsstill
verymuchwithusincludehorseshoecrabs,snails(gastropods),e.g.Cyclonema,
andcorals(althoughPalaeozoiccoralsweresignificantlydifferentfrommodern
corals).Bivalves(notshown)weremuchrarerintheOrdovicianseasthanthey
aretoday.

Allthisfaunawasmarine.Verylittle,ifany,animallifehadmadeitoutofthe
seaontodrylandbytheendoftheOrdovician.Tracefossilsofanunknown,
possiblymillipedelikeanimal(Atlas,p.71)arerareevidencethatinvertebrates
wereexploringtheedgeofthelandbyafreshwaterroute.Thereisfossil
evidencethatbryophytelikeplantsandfungihadbeguntocoloniseland
environmentsbackinOrdoviciantimes.Apartfromasmalladvanceguard,
however,themaininvasionoffreshwaterandlandenvironmentsbyplantsand
animalsdidnotreallygetgoinguntiltheSilurianPeriod.
Figure2:SpecimensofthegraptoliteDiplograptusinOrdovicianshales
(fieldofview3.5cm).Graptolitesoftenlookliketinysawbladeslyingonthe
rocksurface

3TheSilurianPeriodandtheinvasionoftheland

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WhatglobaleventhadreducedglobalsealevelattheendofOrdoviciantimes,
drasticallyaffectingshallowmarineorganisms,andleavingthediversityofearly
Silurianlifeseverelycurtailed?

Nowreadtheanswer

RecentresearchshowsthatminorglaciationscontinuedintotheearlySilurian,
butafterawhiletheoceansbecamewarmeragainandsealevelsrose.Ittook
severalmillionyearsformarinelifetorecover.Atfirst,stromatoliteswere
relativelycommon,apparentlybecausetheorganismsthatnormallysuppressed
themhadsufferedseverely,butbymidSiluriantimes,vastcoralreefswere
establishedintropicalwaters,promotingarichdiversityoforganisms.The
growthofcoralswiththeirtough,mineralisedskeletonsturnsflatseabedsinto
complex3Dtopographies,bafflingoceancurrents,andprovidingsheltered
surfacesandnewecologicalniches.Wewillsoonstudysomeoftheorganisms
thatthrivedinsuchenvironments.

Meanwhile,theanimalsandplantsthatwereinvadingthelandfacedallsortsof
environmentalchallengestowhichtheyhadtoadapt.Forexample,ifamarine
plantcellisdirectlysurroundedbyfreshwater,thewatertendstoflowintoitby
theprocessofosmosis,causingittoburst.Alternatively,ifthecellisdirectly
surroundedbyair,itlosesallitswater,justasseaweedsbecomehardandcrisp
whenstrandedabovehightideandexposedtothewindandthesun.So,to
surviveinair,plantshadtoacquireaneffectiveoutercoattokeeptheright
amountofwaterin.Theyalsohadtoevolvesmall,controllableporestoenable
gasestobeexchangedthroughthiscoat.Expressedthisway,itisalltooeasyto
givetheimpression,quitewrongly,thatsuchinnovationscouldbeachieved
intentionally,almostasifbysomedirectedeffort.Onthecontrary,asinall
evolutionaryexplanations,naturalselectionwouldhavefavouredthose
organismsthatwere,bychance,betteradaptedtothesenewenvironmental
challenges.
Withoutthebuoyancyprovidedbyimmersioninwater,adaptationsinboth
plantsandanimalswereneededtosupportabodyonlandagainstthepullof
gravity.Therelativelyhighdensityofwaterprovidesmuchgreatersupportfor
thebodiesofmarineorganismsthandoeslessdenseairhencetheexpression
likeafishoutofwater.Togrowupoffthelandsurface,plantshadtodevelop
groupsofspecialplumbingcellstoconductwater,nutrientsandtheproductsof
photosynthesisaroundtheirbodies.Onlyabout4cmtall,Cooksonia(seeFigure
3aandillustration,Atlas,p.75)lackedleaves,andsentshort,forkingshoots
upwardtocapturesunlightandreleasespores(reproductivecells)intothewind.
Thesefirsttruevascularlandplantswithspecialisedcellsforcarryingfluids
aroundanuprightplantwerestilldependentonwaterforreproduction,andlived
inswampsandonriverbanksandfloodplains.Theearliestmovementontoland,
forbothplantsandanimals,seemstohavebeenthroughthemediumoffresh,as
opposedtosaline,water.

Air,beingagas,hasalowcapacitytostoreheatcomparedwithseawaterand,to
survive,lifeonlandmustbeabletowithstandrelativelylargetemperature
fluctuationsaswellashigheramountsofultravioletradiationfromtheSun.
Exchangeofgametesduringreproductionisalsomucheasierinwaterwhere
malegametescanswimtofertilisethefemaleones.Theanimalsthat,bychance,
werebestadaptedtolifeonlandwerethearthropods.Theywereverystrongfor
theirsize,andalreadyhadanalmostwaterproofouterskeletonthatwasresistant
todamagebyultravioletlight.Numerouspairedandjointedlimbswithinternal
muscleshelpedovercomegravityandallowmovementoveruneventerrestrial
surfaces.However,thearthropodscouldnothaveinvadedthelandiftherehad
notalreadybeensomesupplyoffoodthere.

AccordingtotheAtlas(p.74),whatarethefirstarthropodsbelievedtohave
eaten?

Nowreadtheanswer

Oncethegreeningofthelandhadbegun,smallmillipedesandwinglessinsects
wereapparentlytemptedontoittoeattherottingplantdebris,andtheyandtheir
remainswereeateninturnbypredatoryorscavengingcarnivorousarthropods
suchascentipedes,scorpionsandsmallspiderlikecreatures(Figure3b).
Figure3:(a)Earlylifeonland.Steganotheca,averyearlylandplant(aclose
relativeofCooksonia)fromSilurianrocksinWales,about4cmtall.Notice
thesporebearingstructuresattheendofeachsimplebranchingstem.(b)
Oneoftheearliestknownlandanimalsaspiderlikecreature(4mmlong
includinglegs)fromSilurianrocksinShropshire,England

4LifeintheSiluriansea

4.1Trilobites

Aswe'veseen,theCambrianexplosionlefttheseasteemingwithahugevariety
ofanimals.Inthefollowingactivityyouwillstudysomeofthemarinelifeatone
particulartimeinthePalaeozoicErathemiddlepartoftheSilurianPeriod,
430Maago.You'lllookindetailatsomefossilswhichcomefromadepositin
theUKcalledtheWenlockLimestone,famousforitsmanybeautifulfossils.The
WenlockLimestonecropsoutmainlyaroundBirminghamandthebordersof
Wales.
Figure4showsatrilobitefromtheWenlockLimestone,calledCalymene(kal
iminny).

Figure4:Calymene,atrilobitefromtheWenlockLimestone

Trilobiteswereamajorgroupofentirelymarinearthropodsthatthrivedinthe
seasofthePalaeozoicEraandeventuallybecameextinctinthePermianPeriod.
Thearthopodsareanimmenselydiversephylum(seeSection1.3).Theirexternal
jointedskeleton(exoskeleton)formsarobustarmourplatingforthebody,
thoughitisflexibleatthejointstoallowmovement.

Beingarthropods,howdidtrilobitesgetaroundtheproblemofytheirgrowth
beingconstrainedbyanexternalskeleton?

Nowreadtheanswer

Trilobitesaresonamedbecausetheyhavethreelobesrunningupanddowntheir
lengthacentralaxis,andtwolobes:oneoneitherside.Theyarealsodivided
crosswaysintoaheadshield,atrunk,andatailpiece.Haveapreliminiarylook
atthefossilinFigure4toseeifyoucanidentifythesedivisions.Thetrunkhasa
numberofseparatesegments,andthetailpieceismadeofasingleplate.The
moreformalnamesforthethreemaindivisionsofthetrilobitebody,oftenused
elsewhere,areasfollows:

headshield=cephalon;trunk=thorax;tailpiece=pygidium.

Trilobitesalsohadappendagessuchasantennaeandlegs,buttheseareextremely
rarelyfound.Anexamplewherelegsarepreservedisshownonp.63ofthe
Atlas.

Whatdoestherarityoftrilobiteappendagessuggestabouttheirstructure?

Nowreadtheanswer

Becausetrilobitescastofftheirshellduringmoulting,mosttrilobitefossilsare
actuallyshedshells,ratherthancarcasses(deadanimals).Thetrilobite'shard
outershellneededplacesofweaknessalongwhichitcouldbreakapartduring
moulting,abitlikehavingweakareasbetweenthepiecesofabarofchocolate.
Therewerelinesofweaknessbetweentheheadshieldandthefrontofthetrunk;
betweenanytwotrunksegments;andbetweentheendofthetrunkandthefront
ofthetailpiece.Trilobitesalsohadoneormorelinesofweaknesswithinthe
headshield.Figure4showstheselinesofweaknessforCalymene;theyallowed
twosidepiecestodetachfromthelarger,centralpartoftheheadshield.

Manytrilobiteshadeyes.InCalymene(Figure4),theyareatthecresentshaped
areasonthedetachablesidepieceseithersideofthecentralpartofthe
headshield.Theeyeswereratherlikethecompoundeyesofamodernfly
(anotherarthropod)inhavingmanylenses.Eachtrilobitelenswasmadeof
calcite,anditspreservationpotentialwasasgoodastherestoftheskeleton(also
mademostlyofcalcite).Trilobiteshadtheearliestrecordedeyesintheanimal
world.

Thepresenceofwelldevelopedeyessuggeststhatwhereverthetrilobiteswere
livingtherewasenoughlighttoseeby,and,aslightfadeswithdepth,thewater
isunlikelytohavebeenverydeep.

ItislikelythatCalymenespentmoretimescuttlingaroundontheseafloorthan
swimminghighupinthewater.Itprobablyrestedontheseafloor,stretchedout
inthesamewayasinFigure4.Thesetrilobitesprobablyatesmallorganismsand
organicdebrisonorneartheseafloor.

Sometrilobitescouldrollup,tuckingthetailsnuglyunderneaththehead.Figure
5showsaspecimenofCalymenethathasrolleditselfupintoaball,likeapill
bug(atypeofwoodlouse,anotherarthropod)cando.Theflat,outeredgesofthe
trunksegmentshadtoslideovereachothertoenablethetrilobitetorollup.
Whymighttheabilitytorollupbeuseful?

Nowreadtheanswer

Bystudyingfossilsonecanobservevariousfeaturesofthemorphologyofan
extinctorganism,andinferaspectsofitsmodeoflifeandenvironment.Wecan
dothisbycomparingobservedfeatureswithwhatisknownoflivingrelatives
andsimilarfossils.Inpractice,whenstudyingfossilsonewouldalsotakeother
informationintoaccount,suchasevidencefromthesedimentenclosingthe
fossil,andsoon.Theresultisalwaysaninterpretationthatreflectsthebalance
ofprobabilitiesfromtheavailableinformation,ratherthanthecertaintruth.

Figure5:AspecimenofCalymeneblumenbachifromtheWenlock
Limestonethatrolleditselfupintoaball(2.5cm)

4.1.1Moreontrilobites

Manythousandsoftrilobitespeciesareknown,mostlyfromCambrianto
Silurianrocks,andallwereconfinedtothePalaeozoicEra.Bythetimetrilobites
becameextinctinthelatePermian,theirdiversityhaddwindledtoasmall
numberofspecies,andthegroupwaslongpastitspeak.Thevariationintrilobite
formisenormous,butthebasicthreelobeddivisionoftheexoskeletonisalways
present.Thenumberoftrunksegmentsvariesfrom2to40.Notallhaveeyes.
Mostareabout210cminlengthbutsomeare12mmandafewspeciesgrew
tonearly1mlong.Themajorityoftrilobiteslivedonornearthefloorofshallow
seas,butsometrilobitesswaminthesurfacewatersoftheopenocean,andsome
wereadaptedtolowconcentrationsofoxygeninwaterhundredsofmetresdeep.
Howtheyreproducedisnotclear.Figure6showsarangeoftrilobitesfromthe
OrdovicianPeriod.
Figure6:AselectionofOrdoviciantrilobitesfromcentralWales.(a)
Oygiocarella(7.5cm).(b)Cnemidopyge(3cm).(c)Telaeomarrolithus(2cm).
Trilobiteslikethiswithapittedfringearoundtheheadarecalled
trinucleids.Themainfunctionofthefringe,whichhastinyholesatthe
centreofeachlittlepit,mayhavebeentoallowwatertopassoutfroma
feedingchamberunderneaththeheadaftertheanimalhadstrainedoff
smallfoodparticles.Thisspecimenisenrolledthetailpiece,alongwhich
thespecimenhaspartlysplit,istuckedupunderneaththeheadshield.(d)
Segmentagnostus.Thistrilobite,about1cmlong,belongstoagroupofsmall
trilobitescalledagnostids,whichhaveonlytwoorthreetrunksegmentsand
aheadshieldandtailpieceofsimilarsizeandshape

4.2Crinoids

Figure7showsthefossilisedremainsofatypeofechinodermcalledacrinoid
(crynoyed).Althoughcrinoidsoccurtoday,theywerefarmorecommoninthe
PalaeozoicandMesozoicEras.Mostcrinoidsfeedbybendingtheirumbrellalike
arrangementofflexibleappendages(calledarms)downstreamsoastocatcha
current,ratherasinanumbrellabeingcaughtinthewind.Tubefeet
(multipurposetentacles)onthearmsgatherfoodparticlessuspendedinthe
water,whicharethenwaftedbysmallhairlikethreadsingroovesalongthearms
tothecentralmouth.Mostancientcrinoidslivedgregariouslyinshallow,
currentsweptareas,freeofmuddysedimentthatwouldotherwisehavetendedto
killthembycloggingtheirfeedingmechanism.
Figure7:Marsupiocrinites,aSiluriancrinoidfromtheWenlockLimestone.
Thelowerpartofthestemisbrokenoff.Thelengthofthepartofthe
specimenshownhereis9cm

Mostancientcrinoidswereattachedtotheseafloorbyastemorstalkwitha
rootlikeholdfast.Themouthandgutweresituatedinanenclosedcupatthetop
ofthestem.Inlife,thestemwasfairlyflexible,likethearms.Themajorityof
livingcrinoidsdonothaveastem,andarecapableofcreepingaroundoreven
swimming.Thefewsurvivingstemmedformsgenerallyliveindeepwater.
Althoughtheyareanimals,crinoidswithstemslookatfirstsolikeplantsthat
theyareofteninformallycalledsealilies.Shortlyafterdeath,thetinyorganic
fibresthataloneholdthecalciteplatestogetherrotaway,oftencausingthe
crinoidtodisintegrateintoseparateplatesthatcanbereadilydispersedby
currentsorthemovementofotherorganisms.Insomeancientenvironments
wherecrinoidswereabundant,rockshaveformedthatarelargelycomposedof
isolatedplatesandstemfragments(Figure8).

Crinoidshaveanarrangementoffiverays,eachofwhichcarriesoneormore
armsthattogetherformanefficientfoodgatheringstructure.Attachedtoeach
armaremanyfinesidebranchescalledpinnules,eachmadeoftinyplates,
whicharealsopartofthefoodgatheringapparatus.
Figure8:AlimestoneofCarboniferousagecomposedofcrinoiddebris,
especiallystemplates.Thestemsareinvaryingdegreesofdisarticulation.
Fieldofview8cm

4.3Corals

CoralsareespeciallyabundantintheWenlockLimestone.

AccordingtoSection1.3,whichtwomaingroupsofcorals,eachconfinedtothe
PalaeozoicEra,mightyouexpecttofindintheSilurianPeriod?

Nowreadtheanswer

Whichofthesetwocoralgroupsonlyformscolonies?

Nowreadtheanswer

Figure9showstworugosecorals.Whenalive,theAcervulariacolonyinFigure
9awouldprobablyhavelookedlikeabunchofseaanemones,witheachofthem
sittinginoneofthebowlshapedhollowsoftheircalciteskeletons.Theskeleton
secretedbyanindividual,whetherpartofacolonyornot,iscalledacorallite.In
rugosecorals,eachcoralliteisusuallydividedbyaseriesofconspicuousradial
partitions,calledsepta(Figure9).ThecorallitesinacolonysuchasAcervularia
shareadjacentwalls.InthecentreofmanycorallitesofAcervulariaisasingle,
bowlshapedhollow.Withinsomecorallitesthereareoneormoresmaller
hollows.Theselittlehollowshavethesamebasicarrangementofseptaasinthe
largercorallites:theyrepresentnewindividualsformedbythesplittingupof
theirparent,enablingthecoralcolonytogrowupwardandoutwardbyaprocess
ofasexualreproduction.
Figure9:RugosecoralsfromtheWenlockLimestone.(a)Acervularia
ananas,arugosecoralcolony(5.5cm);(b)Kodonophyllumtruncatum
(2.5cm).Thisisasolitaryindividualfromaspeciesthatmoreoftenforms
colonies.Mostotherrugosecoralspeciesareeitheralwayssolitaryoralways
colonial

Tabulatecoralsaredistinguishedbyhavingmuchsmallercorallitesthanthoseof
rugosecorals,andseptaareeitherabsentorshortandinconspicuous.Tabulate
coralcoloniesmaytakevariousforms,includingmassive(Figure10a)orchains
(Figure10b).

SomeWenlockLimestonecoralsareshownonpp.74and75oftheAtlas;seeif
youcandistinguishthesolitarycoralsfromcolonialones.Thecoralatthe
bottomofthegrouponp.75isatabulatecoralcalledHeliolites.

Whatsortofenvironmentsdocoralstendtobeassociatedwithtoday?

Nowreadtheanswer

AccordingtoSection1.3,whatisthemaingroupofmoderncorals,andwhatis
theiragerange?

Nowreadtheanswer

Figure11showssomeexamplesofmoderncoralskeletons,bothsolitaryand
colonial.Likerugosecorals,scleractiniansusuallyhaveconspicuoussepta.
However,unlikebothrugoseandtabulatecorals,theskeletonofscleractiniansis
madeofaragonite,notcalcite.Althoughitisalsoaformofcalciumcarbonate,
aragonitetendstoaltertocalciteordissolveaway,sothatMesozoicand
Cenozoiccorals(i.e.scleractinians)areoftenlesswellpreservedthanPalaeozoic
ones.

Most,butnotall,scleractiniancoralstodaygrowinwarm,clear,shallowseas.
Thesameseemstohavebeentrueofmanyancientcorals,includingPalaeozoic
ones.Therequirementofmanyoftoday'scoralsforclear,shallowwateris
mostlybecausewithinthecoralssofttissuesaretinyalgaewithwhichtheylive
inanassociationofmutualbenefit.Thealgaerequireclear,welllitwaterfor
photosynthesis.Byanalogywiththepreferencesofmostmoderncorals,and
whatisknownofancientones,itthereforeseemsverylikelythatthewaterofthe
WenlockLimestoneseawaswarmratherthancold.Thisissupportedby
palaeomagneticevidenceshowingthatatthistimeBritainwasabout25southof
theEquator.Palaeozoicandscleractiniancoralsare,however,unrelatedandthere
isevidencethatPalaeozoiccoralslackedaspecialassociationwithalgae.

Figure10:TabulatecoralsfromtheWenlockLimestone.(a)Favosites(field
ofview8cm);(b)Halysites,thechaincoral(fieldofview3.5cm)
Figure11:Examplesofmodernscleractiniancoralskeletons.Thelargestis
9cmacross

4.4OtherWenlockLimestonefossils

AmongtheotherfossilscommonintheWenlockLimestonearebrachiopods
(Figure12aandb),gastropods(Figure12c)andbryozoans(Figure12d).You
mayneedtorereadSection1.3toremindyourselfaboutvariousaspectsofthese
groups.

Figure13(theunitimage)isareconstructionofatypicalscenefromaWenlock
Limestoneenvironment.Seeifyoucanidentifytrilobites(alive,dead,ormoulted
piecesofexoskeleton),corals(coloniesandisolatedindividuals),andcrinoids
amongthevariousformsoflifedepicted.Theconeshapedsquidlikeanimalsare
straightnautiloids(cephalopodmolluscs,Section1.3).
Figure12:SomeWenlockLimestonefossils:(a)thebrachiopodLeptaena
depressa(2.5cm);(b)thebrachiopodAtrypareticularis(2cm);(c)the
gastropodOriostoma(4cm);(d)thebryozoanHalloporaelegantula(2cm)
Figure13:ReconstructionofatypicalscenefromaWenlockLimestone
environment

5TheDevonianPeriod

Beforegoinganyfurther,clickon'Viewdocument'belowandreadpages7677
fromDouglasPalmer'sAtlasofthePrehistoricWorld.Donotworrytoomuch
aboutallthedifferentnamesoffishgroupsinthis,theAgeofFishes.

Viewdocument

Environmentalchangeisknowntohaveasignificantimpactontheevolutionof
life.Forexample,wideningoceansgeneratebarriersbetweenpopulationsand
promoteincreasinggeneticdivergencebetweenthemovertime.Narrowing
oceans,risingmountainbeltsandtheformationoflargecontinentalareascan
alsohavemajoreffects,causingbothevolutionandextinction.Duringthe
Silurian,theIapetusOceanhadbeennarrowing(Atlas,p.72),allowingthe
progressivemixingofmarineorganismsthathadpreviouslybeenseparatedbya
wideoceanicbarrier.BytheDevonian,thecontinentalareasofLaurentia,
AvaloniaandBalticahadamalgamatedtoformalargelandmass,withhuge
riversandlakesthatwerecolonisedfirstlybyjawlessfishandthenbypredatory
jawedfish.(JawedfishhadfirstevolvedintheOrdovician(Atlas,p.71),but
onlybecameabundantinthelateSilurianandDevonian,evolvingintomany
differentmarineandfreshwatergroups.)Therisingmountainchainsseparated
riversystemsandpromotedtheevolutionofdifferentspeciesoffishindifferent
inlandwaterways.Likewise,landplantswerespreadingglobally,evolving
distinctflorasinparticularregionswithdifferentclimates.

Wenowlookatoneofthemostsignificantdevelopmentsintheevolutionofour
vertebrateancestorsthefirsttetrapodsinthelateDevonian.

6Vertebratesmoveontoland

6.1Adifficultevolutionarytransition

Beforegoinganyfurther,clickon'Viewdocument'belowandreadpages7879
and8283fromDouglasPalmer'sAtlasofthePrehistoricWorld.

Viewdocument

AswesawinSection3,themoveoutofwaterontolandwasaparticularly
difficultevolutionarytransition,requiringmanyadaptations.

Whatspecificadaptationsdidvertebratesevolveforlivingonland,andwhy
weretheyneeded?

Nowreadtheanswer

Theearliestfourlimbedvertebrates(i.e.tetrapods)areknownfromlate
DevonianriverandlakedepositsinGreenland.AcanthostegaandIchthyostega
haveasomewhatpuzzlingmixtureofcharacteristics.Theyhavefourjointed,
muscularlimbsthatwereoriginallyinterpretedaslegsforlivingonland,but
theyalsohavelong,sidewaysflattened,fishtypetailsforswimming,asensory
systemfordetectionofvibrationsinwaterand,mostimportant,fishtypegills.
Thecurrentinterpretationthereforeseesthelimbsasadaptationsforlifeina
particularaquaticenvironmentratherthanforlifeonland.However,theselimbs
alsohadveryappropriatepreadaptationsforthetetrapoddescendantswhodid
leavethewater.BytheearlyCarboniferous,limbswithfivedigitshadbecome
thenorm,whichhasremainedsoeversince.

Lookatthepanoramaonpp.7879oftheAtlas.Howmanydigits(fingersor
toes)didAcanthostegaandIchthyostegahaveoneachlimb?

Nowreadtheanswer

WhatweretheearliesttetrapodlimbssupposedlyusedforinthislateDevonian
aquaticenvironment?
Nowreadtheanswer

ByearlyCarboniferoustimes,fossilsfromlocalitiessuchasEastKirktonin
Scotlandrevealtheexistenceofseveraldifferenttetrapodgroups(Atlas,pp.82
83).Someofthesewerestillfullyaquatic,butothers,suchasthesuperficially
lizardlikeWestlothianaandthesalamanderlikeBalanerpeton,hadprobably
madethetransitiontoasemiaquaticlife.Lifeinthewaterwashazardous,with
intensecompetitionforfood,andpartoftheincentiveforearlytetrapodstoleave
thewatermayhavebeentoexploitabundantfoodresourcesonland.However,
lifeonlandcouldalsobedangerous,aswitnessed,forexample,bythescorpion,
Pulmonoscorpius,whichgrewupto70cmlong.

Youmightwonderwhythewordtetrapodisusedhereinsteadofamphibianor
reptile.Certainly,moderntetrapodsareclearlyseparatedintoamphibians,
reptiles,birdsandmammals,largelyonthebasisoftheirdistinctivereproductive
systems.Amphibiansbreedinwater,wheretheylayunprotectedeggswhichare
externallyfertilised.Bycontrast,reptiles,birdsandmammalsuseinternal
fertilisation.Inreptilesandbirds,theembryodevelopsinfluidsurroundedbya
protectiveshellaneggthatcanbelaidonland.Innearlyallmammals,the
developingembryoisalsosurroundedbyfluidandaprotectivemembrane,butis
retainedinthemother'sbodyforsometimebeforebirth.Suchprotective
membranesarenotconfinedtothemammalsbutarealsofoundinreptilesand
birds,allofwhicharegroupedasamniotes.Theamniotesarethereforemore
independentofwaterthanamphibianswhoseembryosarenotprotectedbysuch
membranesandthereforehavetodevelopinwater.Theproblemisthattheearly
tetrapodfossilsdonotpreservedirectevidenceofreproductivehabits.Also,
duringearlytetrapodevolutiontherewouldnothavebeensoclearaseparation
betweenamphibiansandreptilesaswefindtoday.Onlylater,whenclearly
identifiableamphibianorreptiliancharacteristicsemergeinthefossils,is
classificationeasier.

ByCarboniferoustimesterrestrialfoodchainswerealreadywelldevelopedwith
agreatvarietyofplantssuchastreesizedclubmosses,seedplantsandtreeferns
andsmaller,groundcoveringferns.Theseplantssupportedawidevarietyof
invertebratelife,includingmanyarthropods,whichprovidedabundantfoodfor
thecarnivorouslandlivingtetrapods.

6.2Anoutlineofvertebrateevolution

Let'snowplacetheearlyevolutionoftetrapodsinperspectivebytakingan
overviewofthewholeofvertebrateevolution.
Figure14:Geologicalrangesofvertebratesshowingwhenvariousgroups
evolvedfromeachotherduringthePhanerozoic.Thewidthofeachgroup
indicatesitsapproximatediversity(asnumberoffamilies);notethescale
bar.Thewidehorizontalseparationofsomenewlyevolvedgroupsfrom
theirancestors(e.g.mammalsfromreptiles)doesnotindicatehuge,abrupt
changebutissimplyaconsequenceofthewaythediagramisdrawn

Clicktoviewprintableversionofimage

Viewdocument

Giventheubiquityofourownprimatespecies,andfamiliaritywithdomesticated
animalssuchascats,dogs,andrabbits,ourviewoftherelativediversityand
abundanceofthemainvertebrategroupstendstobebiasedtowardsmammals.
However,atallyoflivingvertebratespeciesremindsusthatmammalsarenotas
dominantassomemightimagine.Inroundterms,thetotalnumberofliving
vertebratespeciesisover46000(whichcompares,forexample,withsome110
000speciesofmolluscsandmorethan80000speciesofroundworms
(nematodes)).Livingmammalsandamphibianseachnumberaround4300
species,whereastherearesome6500reptilespecies,9100birdspeciesandover
21000speciesofbonyfish,andabout800speciesofsharksandrays,the
cartilaginousfish.

Whichoftoday'sfivevertebrateclasses(Section1.3)wereinexistencebythe
endoftheTriassicPeriod(seeFigure14)?

Nowreadtheanswer

Intermsoffamilies,whichlivingvertebrategroupsshowninFigure14areat
theirhighestdiversitytoday?Whichlivinggroupsweremorediverseinthepast,
andwhenwastheirpeakofdiversity?

Nowreadtheanswer

Indiscussionssuchasthis,itisimportanttobeawarewhichtaxonomiccategory
(e.g.order,familyorgenus)isbeingconsideredbecausethepicturecanchange
ifothermeasuresofdiversity(e.g.species)areused.

Activity1

FromFigure14,estimateroughly(tothenearestfive)thenumberofliving
familiesofreptilesandofmammals.Then,usingthedataonspeciesnumbers
givenabove,workoutwhichofthesetwogroupshas,onaverage,morespecies
perfamily.

Nowreadtheanswer

Aswehavealreadyseen,thevertebrateswereinitiallyslowtodiversify.From
theirCambrianstart,itwasnotuntillateOrdoviciantimesthattheearlyjawed
fishevolvedintodifferentgroups,thoughsomeweresubsequentlytobecome
extinct(theplacoderms,seeAtlas,pp.7677,andacanthodians,seeAtlas,p.83).
FromFigure14itbecomesclearwhytheDevonianisoftencalledtheAgeof
Fishes:notonlywasthereagreaterdiversityoffishgroupsthanatanyother
timebutmostoftheothervertebrateclasseshadyettoevolve.

Figure14revealsavarietyofpatternsinfishevolution,includingsomemajor
setbacks.TheplacodermsbecameextinctattheendoftheDevonian,coincident
withadecreaseinearlytetrapoddiversity.Sharks,raysandbonyfishessuffered
amajorextinctionattheendofthePermian,coincidentwithasimilarsetbackfor
amphibiansandreptiles,partofamassextinctionevent.Today,thetotalnumber
offishfamiliesisatitshighestever,mainlyduetothebonyfishes.
Whendidthetotalnumberofbonyfishfamiliesbegintoincreasesteeplyfrom
some95familiestotheirpresenttotalofabout260?

Nowreadtheanswer

Let'snowlookatthehistoryoftetrapods.AfteremerginginlateDevoniantimes,
tetrapodcommunitieswereinitiallydominatedbyamphibians,especiallyin
CarboniferousandPermiantimes,whenawidevarietyofamphibiangroups
evolvedanddiedout.AmphibiansremainedlargeduringthePalaeozoicEra,
often12mlonghugebythestandardsoftoday'sfrogs,toadsandnewts,
whichhavearelativelyrecent,Mesozoicorigin.Thefirstreptilesappearedinthe
earlyCarboniferous.

Whatcrucialadaptationdidreptilesevolvethatfreedthemfromtheamphibian
dependenceonbeingnearwatertolayeggsanddeveloptadpoles?

Nowreadtheanswer

AtfirsttheearliestCarboniferousreptileswerequitesmall,superficiallylizard
likepredators.Theylivedmostlyonarthropods,suchasdragonfliesand
cockroaches,inthedenseequatorialCarboniferousforestsandswamps.The
energyofsunlighttrappedduringphotosynthesisbecamestoredinvastamounts
ofplantlitterthataccumulatedontheflooroftheforestsandinswamps.
Eventually,thisdebriswasburied,compressedandconvertedbyheatand
pressureintocoal;300MalatertheenergyfromCarboniferoussunlightfuelled
theIndustrialRevolution.

NotuntilPermiantimesdidreptilesbegintodiversifywidelyandincreasein
size;someevenreturnedtolivinginthesea.Othersevolvedcertainmammallike
features,suchasthebeginningsoftoothdifferentiation.Thisallowedforseparate
functionssuchaslargedaggershapedteethfordefenceandkillingprey,and
smallercheekteethforefficientprocessingoffoodbeforedigestion.Oneofthese
groupsthatsurvivedtheendPermianmassextinctioneventuallyevolvedintothe
firstmammalsinthelateTriassic.

Activity2

UsingFigure14,measuretheapproximatenumberoffamilies(tothenearest
five)presentinthefourmainvertebrategroups(sharksandrays,bonyfishes,
amphibiansandreptiles),firstattheendofthePermianandthenintheearly
Triassic.Fromthesetotals,calculatethepercentagechangeinfamilydiversity.

Nowreadtheanswer
AftertheendPermianextinctionevent,thereptilesrecoveredandgradually
becamethedominantgroupbytheendofTriassictimes,thoughbothreptilesand
amphibiansweresetbackbyanother,lateTriassic,extinctionevent.

Whichgroupdoyouthinkwasmainlyresponsibleforthedominanceofreptiles
intheMesozoic?

Nowreadtheanswer

Dinosaurswereaspecialgroupofreptilesspecialnotleastbecause,aftertheir
origininthelateTriassic,theybecamethedominantlandanimalsforover
150Ma.Theyfilledalmosteverynichepossibleforlargelandvertebrates.They
includedcarnivores,herbivoresandafewomnivores(mixeddiet),andrangedin
sizefromthatofachickentovastplanteaters,suchastheBrachiosaurus,which
grewupto25mlongandweighedupto40tonnes.Dinosaursnever,however,
tooktotheairortotheoceans:otherlargereptilesthepterosaursdominated
theskies,andmarinereptilessuchasichthyosaurs,plesiosaursandmosasaurs
flourishedinthesea.Themajorsurvivinggroupsofreptiles,thecrocodiles,
turtlesandtortoises,lizardsandsnakesalsoallappearedinMesozoictimes.
MuchevidencesuggeststhatbirdsevolvedfromdinosaursinlateJurassictimes.

ThegreatsuccessoftheMesozoicreptilescametoadramatichaltattheendof
theCretaceous.Reptilediversitywascutbackfromabout60familiesto30and
neverfullyrecovered(Figure14).Thebonyfishesrapidlyincreasedindiversity
intheearlyTertiary.

ForalongtimeaftertheirlateTriassicorigin,mammalsremainedsmall,perhaps
nocturnal,shrewlikecreatureslivinginthenooksandcranniesofthedinosaur
world.ButoncethelargeMesozoicreptileshadbecomeextinct,mammalswere
abletodiversifyintothewiderangeofnichesthatbecamevacant.Plants,which
alsosufferedsomeextinctionsattheendoftheCretaceous,continuedtohavea
roleintheevolutionofbothvertebratesandinvertebrates.Theriseofflowering
plants(angiosperms)andpollinatinginsects,togetherwithclimatechangeinthe
earlyTertiary,promotedtheevolutionofgrasses,whichinturnsupportedthe
spectacularriseofgrazingmammals.Itwasperhapstheclimatedrivenchangein
vegetationthatpromotedourownevolutionfromtreedwellingprimatesinlate
Cenozoictimes.GlobalcoolingaspartoftheinitialdescentintotheQuaternary
IceAgecausedretreatandfragmentationoftheextensiveforestsofcentral
Africaandtheirreplacementbymoreopengrasslands.Thisgaveanadaptive
advantagetoagroupofmoreuprightwalkingprimatesthehominids.Ourown
species,Homosapiens,appearedonly100000to150000yearsago(depending
onhowtheoriginofthespeciesisdefined).
Dothis

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Acknowledgements

ThecontentacknowledgedbelowisProprietary(seetermsandconditions)andis
usedunderlicence.

TheauthorofthisunitisPeterSheldon.

Gratefulacknowledgementismadetothefollowingsourcesforpermissionto
reproducematerialinthisunit:

Figures

Figures1and2:PeterSheldon;

Figure3a:NationalMuseumofWales;

Figure3b:CourtesyofJasonDunlop;

Figures512:PeterSheldon;

Figure13:JohnWatson;

Figure14:AdaptedfromBenton,M.J.(2000)VertebratePalaeontology,second
edition,BlackwellScience.

UnitImage

JohnWatson
ExtractsfromTheAtlasofthePrehistoricWorld

Thecontentacknowledgedbelowappearsinpages5883ofTheAtlasofthe
PrehistoricWorldbyDouglasPalmerpublishedbyMarshallEditions,London
(whichformspartofTheOpenUniversitycourseS193LifeinthePalaeozoic)
andgratefulthanksareextendedtothemandtoallrightsownersfortheir
permission.

Thanksarealsocreditedintheoriginalpublicationfortheirkindhelpduring
productionto:DrAlanSmith,DepartmentofEarthSciences,Universityof
CambridgeandDrNormanMacleod,NaturalHistoryMuseum,London.

Pictures

58l,58r,59t,59b,62t,63t,64,65l,66bSimonConwayMorris,Universityof
Cambridge;60,61DrPeterGrimes;62bRoyalOntarioMuseum;63b,65tr,65br
DrDerekE.G.BriggsUniversityofBristol;66tGaryBell/PlanetEarth;67
NHPA;68tWilliamCampbell/DRKPhoto;68bJensRydell/BruceColeman;69t,
72bMartinLand/SciencePhotoLibrary;70RichardAldridge,Universityof
Leicester;71tBD/IPR/197BritishGeologicalSocietyNERC.Allrights
reserved;71b,73t,73b,76b,77bTheNaturalHistoryMuseum,London;72t
ManfredSchauer;74l,75r,80bSinclairStammers/SciencePhotoLibrary;74r
DepartmentofEarthSciences,CardiffUniversity,Wales;75lKajRSvensson/
SciencePhotoLibrary;76tBreckPKent/OxfordScientificFilms;77tEricand
DavidHosking/Corbis;78ZigLeszczynski/OxfordScientificFilms;79,82r
UniversityMuseumofZoology,Cambridge;80tGIBernard/NHPA;81tSarah
Finney;81bTheManchesterMuseum,TheUniversityofManchester;82l
AndrewMounter/PlanetEarthPictures;83NationalMuseumsofScotland.

Illustrators

60/61,74/75,JimChannell;64/65,82/83ColinNewman;78/79,RobinBouttell,
MarshallEditionsDevelopmentLimited

Maps

ThreedimensionalmapsproducedbyLovellJohnsLimited,Witney,using
illustrationbyHardlinesLimited,Charlburyandbasedondatasuppliedby
PaleomapServicesLimited,Cambridge.

TwodimensionalmapsbyEugeneFleury,MarshallEditionsDevelopment
Limted