Ediacaranand

CambriamBiota

LateProterozoicBuildup(about570mya)
Forallofthe19thcenturyandthefirsthalfofthe20thcentury,
theProterozoiceonhadfailedtoyieldanymetazoanfossils.
Thenin1946,R.C.Sprigg,agovernmentgeologistassessing
abandonedleadminesinsouthernAustraliadiscoveredthefirst
remainsofaremarkablebiotathathastakenitsnamefromthe
region,theEdiacaraHills.
Ediacarananimalsarenowknownfrommanyotherregions
(Mexico,California,Canada,Ireland,Russia,Namibia)andthe
storyseemstobethesamewherevertheyarediscovered.

EdiacaranFauna
Theyrangeinagefrom600to545Mya.

Theyarepreservedasimpressions,typicallyin
sandstone.
Withtwoexceptions,allthemetazoansfromthistime
lackedmineralizedskeletons.
Ediacarananimalscouldbequitelarge,upto50cm
long,butmanywereratherflat.

EdiacaranFaunacont.

Theyexhibitarangeofsymmetries:someareradial(e.g.,
Ediacaria),othersarebilateral(Spriggina).

Ediacaria

(A),aradiallysymmetricalcastof
Ediacariapreservedontheundersideof
asandstonebed,RawnsleyQuartzite,
SouthAustralia.

EdiacaranFaunacont.

Spriggina

Sprigginaispartofacladeofsoftbodied
organismsthatarerestrictedtothe
Precambrian.
SprigginaisknownlargelyfromtheEdiacara
HillsofsouthAustralia,nearAdelaide.The
organismhadacrescentshapedheadand
numeroussegmentstaperingtotheposterior
end;itisonlyaboutthreecentimeterslong.
Sprigginawasdescribedasanannelidbutit
nowappearstoberelatedtothearthropods,
althoughSprigginahadnohardparts,andit
isunclearexactlywhatkindofappendagesit
had.
Spriggina

EdiacaranFaunacont.

Dickinsonia
Dickinsonia is known from Vendian
rocks of south Australia and north
Russia.
It is often considered to be an annelid
worm because of its apparent
similarity to one genus of extant
polychaete, Spinther.
However, in the opinion of some, it
may in fact be a cnidarian polyp.
This specimen is an adult one from the
Ediacara Hills of southern Australia

EdiacaranFaunacont.

Eoporpita
Eoporpitaisoneofthemoststriking
Vendianfossils,notedforitsthick
tentaclessurroundingacentralbody.
Thisspecimenisnearly6cmacrossand
wasfoundattheWinterCoastofthe
WhiteSea;otherspecimenshavecome
fromsouthAustralia.
Untilrecently,Eoporpitawasthoughtto
beachondrophorine,butsome
researchersnowdoubtthisinterpretation
ofEoporpitaandconsiderittohavebeen
abenthicpolypratherlikeaseaanemone.

EdiacaranFaunacont.

Cloudina
ExamplesofgroupswithmineralizedskeletonsincludeCloudina
andsponges
Cloudinahadaskeleton
composedofasimple
calcitetubeandwas
probablysomesortof
polyporganism
Spongesproduceinternal
skeletalsupportscalled
spiculesthathavenow
beenfoundinlate
Proterozoicrocks.
Cloudina

EdiacaranFaunacont.

NextofKin?
Someofthemareclearlyrelatedtojellyfishandothercnidarians.
Someofthebilateralformscouldberelatedtoflatworms.
Thelackofanyevidenceforburrowsthatpenetratefarintothe
sedimentssuggeststhatnoneoftheEdiacarananimalshad
coeloms,sotheyprobablyarenotrelatedcloselyrelatedtothe
otherfossilformingmetazoans(mollusks,arthropods,etc.).

EdiacaranFaunacont.

Energyharvesting?
Theformssimilartojellyfishandseaanemoneswereprobably
carnivores.
SomespecimensofCloudinaappeartohavebeenattackedbya
boringorganism,soatleastoneadvancedpredatormusthavebeen
onthescene.
Thebilateral"wormy"formsprobablyconsumedorganicrich
mud.
Theextremelyflatorganismsthatappeartolackmouthsmay
havehadphotosyntheticendosymbiontslivingintheirtissues.

EdiacaranFaunacont.

Summary
Ediacarananimalsdisappearfromthefossilrecordabout545mya.
Manyofthesespeciesmaywellhavesufferedatthehandsofthe
waveoforganismsthatappearedintheCambrian.
Someseemtohavesurvived,givingrisetomoderncnidariansand
theirkin.
Stromatolitesalsobegintodeclineafterthispoint;possiblydueto
intensifiedpredationandgrazingby"new"metazoans.

CambrianFauna
Themodernphylaofmulticellularorganismsshowupina
"flash"atthebeginningofthePhanerozoicEon(=thestartofthe
PaleozoicEraandtheCambrianPeriod).
545Myamarksthefirstappearanceofcomplex,sediment
penetratingtracefossils
Implication:biganimalswithcoelomsareonthescene.
Yettheymusthavebeensoftbodied,aswedon'thaveagood
bodyfossilrecordfromthisinterval.

CambrianTraceFossils
The Cambrian Tapeats Sandstone occurs in the Grand canyon in Arizona.
The half a billion year old sediments represent tidal flat and near shore deposits
during a time when primitive metazoans ruled the earth.
The Tapeats contains a few trace fossils, include Corophioides
Corophioides.Thesearethe
dwellingtracesofalargeU
shapedannelidworm
whichlivedbeneaththe
sedimentsurface.Erosion
hasremovedthetoplayer,
leavingthebottomofthe
burrowasadishshaped
notch.Thiswasthesingle
mostprominentfossil
foundintheTapeatsinthe
GrandCanyon.

TraceFossilscont.

Simpletracefossilsofbilateriananimals,Rawnsley
Quartzite(2cm).

~530Myasmallshellyfossilsbegintoappearintherecord

Theshellsexhibitarangeofmineralogies(calcite,phosphate,
hardorganics,etc.).
Coiledsnailshellswerepresent,indicatingthatthePhylum
Molluscahadappeared.
SpongesspiculesandtheextinctspongelikePhylum
Archaeocyathawerepresent.
Manyoftheshellyfossilsaredifficulttorelatetoanymodern
group.
Recently,fossilizedembryoshavebeenreportedfromthis
intervalaswell,openingthepossibilitythatitmaybepossibleto
studytheevolutionofdevelopmentdirectlyfromthefossilrecord.

Archaeocyaths

Archaeocyaths are an extinct group of

sponges that had a very brief history.
The first appear roughly 530 million
years ago
Diversified into hundreds of species
during this time period, with some
species contributing to the creation of
the first reefs.
Despite their great success in terms of
numbers, the archaeocyaths were a
short-lived group; completely nonexistent by the middle Cambrian,
some 10 to 15 million years after their
first appearance.

Fossilizedembryos
Eggs of presumed jellyfish
(rear) and segmented worm
(front). About 0.5 mm in
diameter.

Phospatizedanimaleggandearly
cleavagestageembryo,
DoushantuoFormation.(250m)

Trilobita
Trilobiteswereamongthefirstofthearthropods;probablydescended
fromsegmentedworms,developinghardexoskeletons,eyesandtheability
togrowbysheddingtheiroldshells.
Trilobiteswereadominantlifeformduringmuchofthe325millionyears
ofthePaleozoic.
Constituteanextinctclassof
arthropods,theTrilobita,madeupof
eightorders,over150families,about
5000genera,andover15,000described
species.
Speculationsontheecologicalroleof
trilobitesincludesplanktonic,
swimming,andcrawlingforms,andwe
canpresumetheyfilledavariedsetof
trophic(feeding)niches,although
perhapsmostlyasdetritivores,
predators,orscavengers.

Trilobitescont.
Trilobitesarethesinglemostdiversegroupofextinctorganisms,andwithin

thegeneralizedbodyplanoftrilobitestherewasagreatdealofdiversityofsize
andform.
Thesmallestknowntrilobitespeciesisjustunderamillimeterlong,whilethe
largestincludespeciesfrom30to70cminlength

Phacops
Acanthopyge

Tricrepicephalus

Brachiopods
Brachiopodsappearedas
wellabout525mya
Brachiopodsaremarine
lophophorates;relatedtothe
BryozoaandPhoronida.
Theyarefilterfeeders,
collectingfoodparticlesona
ciliatedorgancalledthe
lophophore
Commoninverycoldwater,
eitherinpolarregionsorat
greatdepthsintheocean
Thereareabout300living
speciesofbrachiopods.

Brachiopodscont.
Dividedintotwomajorgroups
ClassInarticulata(includinglingulids),andClassArticulatabased
onthepresenceorabsenceofhingeteethandsockets.
Articulatebrachiopod

A lingulate brachiopod

Morphologically
conservative, having lasted
since the Cambrian with very
little change in shape.
This specimen is of
Lingula,a living brachiopod

Echinoderms
Eocrinoidea
Eocrinoids

are among the earliest groups of echinoderms to appear,

ranging from the Early Cambrian to the Silurian.
Most eocrinoids were sessile and
fed with their long brachioles
(the arm-like structures, which in
this specimen are spirally
twisted).
The body was covered by
plates; in early eocrinoids the
holdfast was also covered by
plates, but later eocrinoids
evolved a stalk with columnals,
like crinoids and blastoids.

Gogia from the Middle Cambrian House

Range of Utah.

Echinoderms cont.

Helicoplacoids
The

Helicoplacoidea is a small group of fossil

echinoderms known only from the Lower
Cambrian.
In life, they were shaped somwhat like a
slender football or a fat cigar, and were able to
extend or contract the length of their bodies.
Their "skin" was covered in spirals of
overlapping ossicles that functioned like armor;
their "mouth" was a long groove that also
spiralled around their body.
It is thought that helicoplacoids lived in
burrows, extending their bodies outward to feed.
The helicoplacoids are among the oldest
groups of echinoderms to appear in the fossil
record, along with eocrinoids

Fossil of Helicoplacus from the

Lower Cambrian strata of the
White Mountains in California

Reefs
Thereefsofthetimeperiod(~525mya)werecomposedof
archaeocyathids(extinctanimalsrelatedtosponges)

Archaeocyatha

~525to515Mya:BurgessShaleandSimilarDeposits
DiscoveredbyWalcottin1909
TheBurgessshaleformation,locatedhighintheCanadianRockies,
hasbeenproducingastonishingarrayofsoftbodiedandhardbodied
fossilsfromtheCambriansinceitsdiscovery.
Fossilsfromthesitehavebeencollectedandstudiedbyanumber
ofgroups,andnewBurgesstypelocalitieshavebeendiscovered
alongthewestcoastofNorthAmerica,andineasternNorth
America,Greenland,andChina.

SomeoftheBurgessanimalsfitnicelyintoextantphyla
ofsoftbodied(andshelly)animalssuchas:
Sponges
Annelidworms
Priapulidworms
Arthropods(thoughsomeofthearthropods,such
asMarella,don'tlooklikeanylivingarthropod)

CambrianSponges
ThebranchingspongeVauxiawasoneofthemost
commonspongesencounteredbyWolcottinthe
BurgessShale

Vauxiaspp.

CambrianAnnelids
Canadiaspinosa,apolychaeteannelidabout1to2inchesin
length
Theheadboreapairofslendertentacleswhilethebodywas
coveredwithinnumerablesetae(shortbristles).
Thegutcouldbeevertedanteriorlytoformafeedingproboscis.
Canadiacoulduseitslimbs
towalkonthesubstrateor
swimjustaboveit.
Sedimenthasneverbeen
foundinthegut,suggesting
thatthiswormmayhavebeen
acarnivoreorscavenger.
Canadia

Burgessochaeta
A polychaete worm related to Canadia

Burgessochaeta

CambrianPriapulids
Ottoiaprolifica,probablylivedinaU
shapedburrowthatwasconstructedinthe
substrate.

Note the anterior proboscis

(on the left) and the dark trace
of the interior digestive tract.
Ottoia was probably
carnivorous.

Ottoia

CambrianArthropods
Asmall"arthropod"somewhatreminiscent
ofatrilobite,butwithseveraldistinctive
features
Called the "lace crab" by Walcott, Marrella is the
most abundant type of aminal in the Burgess Shale.
More than 15,000 have been collected.

Marrella splendens

UncertainFossils?Wiwaxia
Wiwaxia is believed to be closely related to either polychaetes, but is not thought to be a member of
the group
Longerspinesprojectintworowsalongtheback,andevidentlyprovidedsomeprotectionfrom
predators.
Therestofthedorsalsurfaceiscoveredwithsmall,flat,overlappinghardplates,termedsclerites.
EachoftheselittlescaleswasattachedwitharootlikebaseandweassumeWiwaxiagrewby
moltingtheseplatesfromtimetotime.
Itdidhaveananteriorjawwithtworowsofteethontheventralsurface,suggestingitwasanother
bottomfeeder.

Wiwaxia

UncertainPhyla?Anomalocaris

Largecreatures(60cmlong)

Long,ovalshapedhead,largeeyes,feedingappendagesatthefront
thatlooklikecombs,andacircularmouthunderneath.
Behindtheheadisa"trunk"withlobesunderneathitinpairsfor
swimming.
Nolegsorwalkingappendages;probablyswamallthetime.
Themouthwaslocatedonthefrontendofthehead,underneath;
roundandcylindrical,withmanytinyteethfacinginward.

UncertainFossils?Dinomischus
Dinomischus was a wine glass shaped animal, measuring about one inch long.
It had a bulb at the base of its stem to secure it in the mud.
On its circular upper surface, surrounded by petal-like bracts, was a mouth and
an anus.

UncertainPhyla?Opabinia
Acreaturewithfiveeyesandalongflexibleproboscistippedwith
graspingspines
Italsopossessespaddlelikeprojectionsattheposteriorendofthe
body.
Opabiniaisthoughttohavelivedinthesoftsedimentontheseabed,
althoughitpresumablycouldhaveswumafterpreyusingitssidelobes.
Superficially,Opabiniaresemblesacrustacean,butlacksimportant,
distinguishingdetails.
Itremainsunassignedtoanyotherextinctorcurrentlyliving,major
group.

UncertainPhyla?Aysheaia

Possesses an unusual assembly of spines and grasping arms at the

head end. Its mouth lies in the center of a ring of six finger-like
projections.
The limbs of this animal are not jointed; instead, they are tapered,
lobe-like appendages, ten pairs in all
Aysheaia may have been a parasite living on sponges since it is
commonly found in association with their remains (spicules).
Presumably, the spiny parts at its head were designed for grasping
and feeding on its prey.

Aysheaia

UncertainPhyla?Hallucigenia

Oldinterpretation

Newinterpretation

Note the paired spines (now

interpreted to stick up on the
dorsal side - or back), and the
slightly curved legs; caterpillarlike.
Thoughttorelatedtothe
onychophoransor"velvetworms

CambrianChordatesPikaia
Itisbelievedtobeoneoftheearliestknownrepresentativesofthe
phylumChordata
It has a well defined notochord near the dorsal surface.
Also possesses rib-like features which are believed to be muscles.
Pikaiaprobablyswamabovetheseafloorusingitsbodyandan
expandedtailfin.

ExplanationsfortheCambrianExplosion
A.EnvironmentalExplanations
OceanChemistry
Changeinoceanchemistryallowingshells.
SoftbodiedBias
Anapparentexplosion,duetothestrongbiasagainstfindingsoft
bodiedforms
ChangeinOxygen
Suddenoxygenbuildupthatallowsbigbodiesandperhapsskeletons.
Maybe.Thereisstronggeochemicalevidenceforhigheroxygen
levelsattheProterozoicCambrianboundary.

A.EnvironmentalExplanationscont.

PositionsofContinents
Between750and570mya,thecontinentsweregroupedtowardthe
SouthPoleandtherewereseveralepisodesofcontinentalglaciation
effectingmanyareasoftheworld
However,thiswassoonfollowedbymovementofthecontinents
awayfromoneanother
Thisresulted
inhigher
temperatures
andanincrease
incoastlineand
continental
shelf,
augmentingthe
rightconditions
formarinelife

B.BioticExplanations
ArmsRace

Therapiddiversificationmayrelatetoapredator/preyarmsrace.
Predationwouldcertainlyfavoranimalswithskeletons.

HoxGenes

Evidencefromdevelopmentbiologyindicatesthattherapid
developmentofcomplexbodyplans,withmanydistinctcelltypes
andanatomicalstructurescanoccurthroughtheactionofHoxgenes

HoxGenes

Hoxgenesevolvedfromamoreinclusivegroupofgenesthe
homeoboxgenescodingforspecificproteinsthatactivateother
genes
Hoxgenesareuniquelyarrangedinalinearsequencealongthe
chromosome,whichcorespondswithboththelinearandthetemporal
sequenceoftheiractivationalongtheanterioposterioraxisofthe
embryo

HoxGenescont.

ThenumberofHoxgenesarrangedinaclusteralonga
chromosomeisbroadlycomparabletothedegreeofcomplexityof
theorganism

Oneinsponges,4to
5incnidarians,610
inmostofthehigher
metazoa,andupto39
arrayedin4Hox
clustersondifferent
chromosomesin
mammals

HoxGenescont.

Thesegenescontrolthepositionandtheexpressionofmajor
structuralfeaturesofthebody,includingtheelementsoftheheadand
thesequenceandnatureoftheappendages
Hoxgenesactasswitchestocontroltheexpressionofavarietyof
genes,whichinturncontroldifferentstructuresandcelltypes
Theoriginofmulticellularityandcomplexbodyplansamong
animalswasauniquephenomenon,dependentontheevolutionof
HoxgenesneartheendofthePrecambrian
Onceevolved,theirsubsequentduplicationanddivergentchangein
adaptivelydistinctlineagesestablishedthebasisfortheradiationof
themanymetazoanphyla

HoxGenescont.

Mostphylahaveapparentlyretainedarelativelyconstantnumber
ofHoxgenessincetheCambrian
Also,wecanrecognizeahierarchyofchangeassociatedwithHox
genesbetweenandwithinphyla

CambrianFaunaandEcosystemFunction