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AcritarchsandChitinozoa

AcritarchsandChitinozoa

Introduction
Acritarchs,thenamecoinedbyEvittin1963whichmeans"of
uncertainorigin",areanartificialgroup.Thegroupincludesany
small(mostarebetween20150micronsacross),organicwalled
microfossilwhichcannotbeassignedtoanaturalgroup.Theyare
characterisedbyvariedsculpture,somebeingspinyandothers
smooth.Theyarebelievedtohavealgalaffinities,probablythe
cystsofplanktoniceukaryoticalgae.TheyarevaluableProterozoic
andPalaeozoicbiostratigraphicandpalaeoenvironmentaltools.
Chitinozoaarelarge(502000microns)flaskshapedpalynomorphs
whichappeardark,almostopaquewhenviewedusingalight
microscope.TheyareimportantPalaeozoicmicrofossilsas
stratigraphicmarkers.

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HistoryofStudy
Theearliestpalynomorphswhichwenowclassifyasacritarchsare
probablythosereportedbyWhitein1862fromtheOrdovicianto
DevonianofNewYork.Intheearly1930'sEisenack,workingon
materialfromtheBaltic,reassignedmanyspeciestotheacritarch
groupandcametoregardthem(aswedonow)asbeingderived
fromphytoplankton.ItwasDownie,EvittandSerjeanthoweverwho
tidieduptheclassification,providingausablesystem,eventhough
wearestilluncertainofthegroupstrueaffinities.
ChitinozoawerefirstnamedbyEisenackin1931,whonotedthe
chitinlikewallmaterial,(theyareactuallycomposedofa
pseudochitinousmaterial).Astherearenolivinganaloguesweare
againuncertainoftheiraffinities,Kozlowski(1963)proposedthey
weretheeggsofannelidworms.Thecooccurrenceofchitinozoa
withscolecodontsandcloselylinkedtrendofabundanceadd
weighttothisproposal.In1970Jenkinsrecognisedanaffinity
betweenchitinozoaandgraptolites,againcloseassociationsof
occurrence,geologicalrangeandtrendsofabundancewerenoted
leadingtohistheorythatgraptolitesandchitinozoaareprobably
geneticallylinked.Paris(1981)dividedchitinozoaintotwoorders,
theProsomitiferaandtheOperculatifera.

Range
TheoldestknownAcritarchsarerecordedfromshalesof
Palaeoproterozoic(19001600Ma)ageintheformerSovietUnion.
TheyarestratigraphicallyusefulintheUpperProterozoicthroughto
thePermian.FromDevoniantimesonwardstheabundanceof
acritarchsappearstohavedeclined,whetherthisisareflectionof
theirtrueabundanceorthevolumeofscientificresearchisdifficult
totell.
ChitinozoafirstappearintheearlyOrdovician(Tremadocstage)
andbecameabundantanddiversebytheAreniganduntilthe
Silurian.ManygenerabecameextinctattheendoftheSilurianand
fewappearintheDevonian,theydisappearintheearliest
Carboniferous.

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Classification
Naturalclassificationsofacritarchshavenotoftenbeenattempted
noraccepted.TheartificialschemeintroducedbyDownie,Evittand
Serjeanthasbeenwidelyacceptedandservesthepurposesof
mostpalynologistssothereislittleincentivetochangeit.Acritarchs
arethereforedividedintothefollowinggroups(mostcommonfirst)
basedontheirmorphology.
Acanthomorphshavesphericalbodieswithspineswhich
usuallyopenintothebody.
Polygonomorphshaveabodyshapedefinedbythenumber
andpositionofspines,theyareoftentriangularorsquarein
outline.
Netromorphshaveafusiformbodywithoneormorespines.
Diacromorphshavesphericaltoellipsoidalbodieswith
ornamentconfinedtothepoles.
Prismatomorphshaveprismatictopolygonalbodiesthe
edgesofwhichformaflangeorcrestwhichmaybeserrated.
Oomorphshaveaneggshapedbodywithornamentconfined
toonepole.
Herkomorphshavearoughlysphericalbodydividedinto
polygonalfields,ratherlikeafootball.
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Pteromorphshavearoughlysphericalcentralzoneoften
compressed,surroundedbyaflangeorwinglamellawhich
maybesustainedbyradialfoldsorprocesses,theyresemble
underthelightmicroscopeafriedegg!
Sphaeromorphshavesimplesphericalmorphology.

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Othergroupsonceassignedtotheacritarchsarenowthoughttobe
prasinophytesorothergreenalgae,theseincludesomespeciesof
herkomorphs,pteromorphsandsphaeromorphs.
ChitinozoahavebeenclassifiedbyParis(1996)basedonthe
structureoftheiropeningsandoverallshapeofflaskandneck,
againthisisnotanaturalclassificationbutithasbecomewidely
accepted.TheOrderOperculatiferacontainingonefamilythe
Desmochitina(simplesphericalformsoftenfoundinchains)and
theOrderProsomatiferacontainingtwofamiliestheConochitinidae
(withlittledifferentiationbetweenflaskandneck)andthe
Lagenochitinidae(withclearlydistinctflaskandneck).

Applications

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Acritarchsareextremelyvaluableforstratigraphiccorrelationof
rocksofProterozoicandPalaeozicage,primarilybecausetheyare
theonlymicrofossilscommonlypreserved.Chitinozoaaresimilarly
utilisedinstudiesofPalaeozoicstrata.Bothgroupsarealsousedin
palaeoenvironmentalreconstructions,providingusefulevidenceon
environmentalconditionswherefewotherfossilsexist.

Biology
Acritarchmorphologyformsthebasisoftheirclassification
thereforeitisworthdescribinginsomedetail.Overallbodyshape
rangesfromsphericalthroughfusiform(grainofriceshaped),
stellatetoprismatic.Processesmaybesimpleorcomplex,solidor
hollow,openingintothebodyorseparatedfromit.Ornamentmay
occuronthebodyand/ortheprocesses.Theformofopeningcalled
apylome(throughwhichexcystmentoccurs)rangesfroma
distinctiveopeninginadefinedarea,arandomsplitanywhereon
thebodyorcompletedisintegration.

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Chitinozoaareflaskshapedsacsorvesicles,theirsurfacemaybe
smooth,striated,granular,spongy,spinyorornamentedwith
branchingappendages.Themainchambermayhaveaneckwitha
markedflexurebetweenthetwo,theopeningorpseudostomeis
coveredbyadiscoroperculumwhichmaybecarriedbyaringed
stopperliketubeorprosome.Thebaseofthevesiclemaybe
convex,pointedorflat,sometimeswithacrownofappendages.It
isthoughtthatatleastsomechitinozoawerelinkedtogetherat
thesepointstoformcolonialchains.

Lifecycle
Acritarchsareextantbutlittleisknownabouttheorganismwhich
producesthem.Itisgenerallyacceptedthattheyareprobablythe
restingcystsofmarinephytoplanktonandthereforerepresentonly
onestageofamultistagelifecycle.Thegrossmorphologyof
acritarchs,asingleorganicsacorhollowvesicle,andtheirsize,
generallybetween15to80microns,suggestsderivationfroma
unicellularorganism.Manyshowsimpleexcystmentstructures
whichmaybedividedintotwogroups:linearsutureswhichvaryin
positionandshape,andpylomeswhicharecompletesmallcircular
holesinthevesiclewhichmayhaveaplugwhichfitsinthehole
calledanoperculum.Thesefeaturesarestronglysuggestiveofa
dinoflagellatelikehostorganism.Thefactthattheyareonlyfound
inrocksofmarineorigin,theirgreatageandotherpreservational
associationsallowsustobealmostsuretheyarewhollymarine.
Chitinozoaareextinctmarineorganismsanditisthereforeeven
hardertodrawanyinferencesabouttheirlifecyclefromtheirshape
andstructure.Theyhavebeenassignedtotheprotozoans,
metazoans,protistsandfungi,howeversomeconclusionshave
beenmade.Thechainformingattributeofmanychitinozoaisan
importantclue,thegenerallyacceptedtheoryisthatthechitinozoa
aretheeggsofametazoan,aslongcoiledchainsofeggsare
knowninsomegroupsforinstancepolychaetewormsand
molluscs.Thefactthatthechitinozoavesiclesofeachspeciesare
allofthesamesizehintsthattheyaretheproductofonelifestage.
Therehavebeenreportsof"juvenile"chitinozoabutthesehavenot
beenconvincingandhavenotbeengenerallyaccepted.

PreparationTechniques
WARNING:Pleaserememberallpreparationtechniquesrequire
theuseofhazardousmaterialsandequipmentandshouldonlybe
carriedoutinproperlyequippedlaboratories,wearingthecorrect
safetyclothingandunderthesupervisionofqualifiedstaff.In
particularthepreparationoforganicwalledmicrofossilsrequiresthe
useofpotentiallylethalchemicalsandproceduresandshouldonly
becarriedoutbytrainedpersonnel.
Clays,silts,mudstones,sandstones,limestonesandshalesmay
firstbebrokendownphysicallyintolessthanfivemillimeter
fragments.Thesampleisthenplacedinaplasticbeakerinafume
cupboardandcoveredinhydrochloricacidtoremovecarbonates.
Thesampleisthenremovedfromthehydrochloricacidandplaced
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inhydrofluoricacidtoremovesilicates.Onceremovedfromthe
hydrofluoricacidthesampleischeckedinawetmounttoascertain
thenextprocedurerequired.Ifthesampleiscleanofmineral
matteritcanbeoxidisedeitherusingnitricacidorSchultze's
solution.Ifmineralmatterispresentboilinginhydrochloricacid
mayberequiredbeforeoxidation.Thesampleisremovedfromthe
lastacidtreatmentandcaneitherbeswirledtoseparateorganic
matterfromanyremainingmineralmatterorheavyliquid
separationcanbeusedwhereaheavyliquidwithspecificgravityof
1.62.5isaddedtothesampleandcentrifuged,theorganicmaterial
floatingtothesurface.
Twotechniquesforwashingtheacidoutofthesamplescanbe
used.Thedecantingmethodinvolvesallowingtheacidandsample
mixtosettleandthencarefullydecantingofftheacid,toppingup
withdistilledwaterandrepeatinguntiltheliquidisneutral.Thisisa
timeconsumingmethod.Thesecondmethodistowashtheacid
andsamplemixthroughanacidresistantsievegauzeofamesh
sizesuitabletoretainthepalynomorphs.Themeshsizesoften
usedaretenandtwentymicron,tenmicrongauzesbeingfine
enoughtocatchallbutthesmallestsporesandacritarchs,soifitis
knownthesemaybepresentfivemicronmeshsizemaybeused.
Chitinozoaarelargeenoughtobecapturedintwentymicron
gauzesandareoftenpreparedforseparately.
Stainingofsamples(withSafranin0,aredcolour,forexample)
maybeuseddependingonthenaturalcolourremaininginthe
palynomorph.Coalsand"coallike"samplesareoftensodevoidof
mineralmatterthattheinitialacidtreatmentscanbeomittedand
oxidationonlyisrequired.Thisneedscarefulhandling,however,as
theoxidisingreactionmaybeextremelyvigorous.Kerogenslides
areoftenrequiredandrequireaspecialpreparationtechnique
withoutanyoxidationorsievingtoretainthetotalorganicmatter
preservedinthesample.Onceallacidtreatmentandseparation
processesarecompletethesamplemaybemountedbystrewing
ontocoverslipsandallowingtodry.Theinvertedcoverslipisthen
gluedontoaslideusingNorlandOpticalAdhesiveorPetropoxyor
similarproprietaryglueofsufficientrefractiveindex.

ObservationTechniques
Palynologyslidesareexaminedusingtransmittedlight
microscopes,commonlywithtimesfortydryandtimesonehundred
oilimmersionobjectivelenses.Accuratecoordinatesofindividual
specimensonaslidemayberequiredandareoftengivenusing
eitherthegraduatedscaleonthetraversableslidetableofa
particularmicroscopeor,preferably,bygivingEnglandFinderco
ordinates.AnEnglandFinderisaspeciallymadeslidewhichis
dividedintoagridofsegmentseachonegivenanalphanumeric
reference.Whenaspecimenislocatedonaslidetheslideis
carefullyremovedfromthemicroscopeandreplacedbythe
EnglandFinderandthecoordinatesmarkeddown,aslongasthe
orientationoftheslideisalsonoteditisthenpossibletorelocate
thespecimenusingtheEnglandFinderonanymicroscope.

Images

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Thefollowingimagesareofarepresentativeselectionofacritarchs
andchitinozoaaimedatgivingageneraloverviewofthedifferent
morphotypes.Eachspecimenisgivenagenericandifpossiblea
speciesnamefollowedbyitsagerange,thesitelocationfrom
whichthesamplewasobtainedandthemagnificationatwhichthe
imagewastakenoritssizeinmicrons.Clickonanimagetoviewa
largerversion.
Multiplicisphaeridiumsp.
Silurian
Rushbury,Wales
LM
colonyofLeiosphaeridiumsphaeromorph
acritarchs
Silurian
Rushbury,Wales
LM
Crassiangulinatesselita
Silurian
Rushbury,Wales
Neoveryhachiumcarminae
Silurian
RoseHill,NewYorkState,USA
LM
Fungochitinasp.
Silurian
RoseHill,NewYorkState,USA
length250micronsLM
Fungochitinasp.
Silurian
RoseHill,NewYorkState,USA
length250micronsLM
Fungochitinasp.
Silurian
RoseHill,NewYorkState,USA
length250micronsLM
Lagenochitinasp.(right)Conochitinasp.(left)
Silurian
RoseHill,NewYorkState,USA
Lagenochitinasp.
Silurian
RoseHill,NewYorkState,USA
LM
chitinozoa(aboralview)
Silurian
unknown
Veryhachiumsp.
Silurian
Rushbury,Wales
19micronsexcludingprocesses
Cymatiosphaeraoctoplana
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Silurian
Rushbury,Wales
25microns
Leiofusatumida
Silurian
JoslinHill,NewYorkStateUSA
25micronsexcludingprocesses
Veryhachiumcf.lairdi
Silurian
JoslinHill,NewYorkStateUSA
9micronsexcludingprocesses
Veryhachiumcf.lairdi
Silurian
JoslinHill,NewYorkStateUSA
13micronsexcludingprocesses
Multiplicisphaeridiumsp.
Silurian
JoslinHill,NewYorkStateUSA
20micronsexcludingprocesses
Leiofusasp.
Silurian
JoslinHill,NewYorkStateUSA
7micronsexcludingprocesses
Gerongracilis
Silurian
JoslinHill,NewYorkStateUSA
12micronsexcludingprocess
Gerongracilis
Silurian
JoslinHill,NewYorkStateUSA
12micronsexcludingprocess
Veryhachiumlimaciforme
Silurian
JoslinHill,NewYorkStateUSA
25micronsexcludingprocess
Multiplicisphaeridiumsp.
Silurian
JoslinHill,NewYorkStateUSA
20micronsexcludingprocess
Veryhachiumtrispinosum
Silurian
JoslinHill,NewYorkStateUSA
12micronsexcludingprocess
Diexallophasisdenticulata
Silurian
JoslinHill,NewYorkStateUSA
12micronsexcludingprocess
Veryhachiumlairdi
Silurian
JoslinHill,NewYorkStateUSA
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12micronsexcludingprocess
Multiplicisphaeridiumfisherii
Silurian
JoslinHill,NewYorkStateUSA
18micronsexcludingprocess
Leiofusabernesgae
Silurian
JoslinHill,NewYorkStateUSA
10micronsexcludingprocess
Geronsp.
Silurian
JoslinHill,NewYorkStateUSA
21microns
Veryhachiumsp.
Silurian
Cyrenaica,Eastern,Libya
40micronsexcludingprocesses
?Diacrodium?sp.
Tremadocian(LowerOrdovician)
32micronsexcludingprocesses
?acritarch?
Tremadocian(LowerOrdovician)
32micronsexcludingprocesses
?acritarch?
Tremadocian(LowerOrdovician)
43micronsexcludingprocesses
acritarch
Tremadocian(LowerOrdovician)
45micronsexcludingprocesses
acritarch
Tremadocian(LowerOrdovician)
29micronsexcludingprocesses
PriscothecacomplanataDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Shropshire,UK
40micronsexcludingprocesses
StelliferidiumcuvillieriDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
38microns
CymatiogaleamembranispinosaDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
30micronsexcludingprocesses
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VulcanisphaerabrittanicaRasul
Tremadocian(LowerOrdovician)
Tremadoc,Wales
30micronsexcludingprocesses
AcanthodiacrodiumangustumDownie
Tremadocian(LowerOrdovician)
Tremadoc,Wales
36microns
VulcanisphaeraimparilaRasul
Tremadocian(LowerOrdovician)
Tremadoc,Wales
32micronsexcludingprocesses
VulcanisphaeraafricanaDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
48micronsexcludingprocesses
TrichosphaeridiumannovolaenseTimofeyev
Tremadocian(LowerOrdovician)
Tremadoc,Wales
46microns
StelliferidiumcuvillieriDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
32microns
CymatiogaleaconvexaRasul
Tremadocian(LowerOrdovician)
Tremadoc,Wales
31microns
PriscothecacomplanataDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
33microns
SaharidiafragileCombaz
Tremadocian(LowerOrdovician)
Tremadoc,Wales
68microns
VulcanisphaeraafricanaDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
43microns
LeiofusasquamaDeunff
Tremadocian(LowerOrdovician)
Tremadoc,Wales
80microns
ArcheohystrichosphaeridiumfunisRasul
Tremadocian(LowerOrdovician)
Tremadoc,Wales
32microns
LeiofusasimplexRasul
Tremadocian(LowerOrdovician)
Tremadoc,Wales
45microns
ImpluviculusmiloniVanguestaine
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Tremadocian(LowerOrdovician)
Tremadoc,Wales
10microns

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