PALEOLECTURE,PAGE1

I.TalesToldbytheDead
A.Paleontologystudyofancientlife
Fossil=anyevidenceofprehistoriclife
1.Paleozoologystudyoffossilanimals
a.Invertebratepaleontologystudyoffossilinvertebrates(animalswithoutavertebral
column)
b.Vertebratepaleontologystudyoffossilvertebrates(animalswithavertebralcolumn)
2.Paleobotanystudyoffossilplants
a.Palynologystudyofpollenandspores(somealsoincludemarineonecelled"plants";i.e.
acritarchs,dinoflagellates,tasmanites,silicoflagellates,diatoms,ebridians,calcareous
nannoplankton/coccoliths)
3.Micropaleontologystudyofsmallfossils(includesmanygroupsmentionedunder
palynologyandalsoforaminifera,radiolaria,chitinozoa,graptolites,pteropods(gastropods),
ostracods(crustaceans),conodonts
B.Objectivesofthepaleontologist
1.Identification
2.DetermineForm(=Morphology)andFunction
3.Associationofplantsandanimalsandenvironmentalreconstruction(paleoecology)
4.EvolutioninVariousOrganisms
5.Dispersalanddistributionofplantsandanimalsthroughspaceandtime(includingstudiesof
paleozoogeography/paleogeographyandbiostratigraphy)
6.CorrelationandDatingRocks
7.StudiesofGeochemistryespeciallychangesinoceanchemistryduetoactionsoforganisms
C.Prerequisites/PreferredConditionsforfossilization:
1.Relativelyabundantorganisms

PALEOLECTURE,PAGE2
2.Presenceofhardparts
3.Avoidchemicalandphysicaldestruction
rapidburial,typicallywithinarelativelylowenergydepositionalenvironment
preservationdependsonEh/pHenvironment;plantsoftenpreservedwithinacidicandreducing
conditions;calcareousshellsandbonestypicallywithinnonacidic
D.TypesofHardParts
1.Plants
a.Cellulosefibrouspolysaccharideformingcellwalls
b.Lignincomplexpolymerbindingcellulosefibers
2.Invertebrates
a.OrganicCompoundsinclude:
a1.Chitin=nitrogencontainingpolysaccharide(carbohydrate)formingfibrousmolecules;
Ex.=arthropods
a2.Scleroproteins=fibrousproteinssuchascollagen(Ex.=graptolites)andconchiolin(Ex.=
molluscs)
b.Mineralsinclude:
b1.Calciumcarbonate=formsintergrowthofcrystalsinanorganicmatrix;includescalcite
(Ex.=echinoderms)andaragonite(Ex.=somemolluscs;aragoniteisachemicallyunstable
mineralandtypicallyrecrystallizestocalcite)
b2.Opalinesilica=oftenoccursasspicules(discreteparts;Ex.=somesponges)orforms
coherentnetwork(ex.=radiolarians)
3.Vertebrates
a.Bone=collagen(ascleroprotein)hardenedbymineralsalts(mostlycalciumphosphate);
withcellularstructure
b.Cartilage=aresilient,partiallyfibrousprotein;usuallynotpreserved
c.Teeth=withdensecalciumsaltsoverlainbyenamel(almostpurecalciumphosphateand
carbonate)

PALEOLECTURE,PAGE3
E.TypesofFossilPreservation
1.Unaltered
a.UnalteredSoftParts
unstableorganiccompoundssuchascarbon,hydrogenandoxygen
rarelypreserved;sometimeswithinpermafrost(Ex.=mammoths)orglaciers,mummification
indrycaves(groundsloths),tanningbyhumicacidsinpeat(Ex.="bogpeople"),within
anaerobicaqueousenvironments(suchasthe"limnicstagnationdeposits"intheEoceneGerman
"browncoal"atMessel),withinoilseeps,andinamber
b.UnalteredHardparts(DuraparticPreservation)
preserveoriginalcalciumcarbonateorcalciumphosphate"hardparts"suchasbone(ex.=La
Breatarpits,California),shells,"coralline"algae;relativelyrare
2.Alteredmoretypicalcase.
a.Petrificationincludes:
a1.CellularPermineralization(Impregnation)=percolatinggroundwaterintroducesminerals
(ex.=silicates,carbonates,ironcompounds,phosphates)intotheporespaces(especially
permineralizecalcareousshellswithcalcite;alsowoodandboneoftenpermineralized)
CoalBalls=permineralizeuncompactedpeatwithcalciumcarbonate;especiallyimportantfor
Carboniferousplantstudiesfrombituminouscoalbeds
a2.Recrystallization=changeformand/orsizeoforiginalcrystalstructure;Ex.=conversion
ofaragonitetocalciteoftendestroysfossildetail
a3.Replacement=percolatinggroundwaterdissolveshardpartsandreplacesthemwith
differentminerals;Mineralsinvolvedincludecarbonates,silicates,ironoxidessuchashematite
and"limonite",pyrite,andcollophane
b.Carbonizationvolatilecomponents(hydrogen,oxygen,nitrogen)decreaseandtheoutline
oftheanimalsispreservedasacarbonfilm;scleroprotein,chitin,celluloseandligninmaybe
carbonized;oftencombineswithpetrification
CoalifiedCompressionsplantcellwallscollapseafterdeposition;causelossofsoluble
materialswithresiduesalteringtoblack,coalydeposits
3.TracesofAnimals
a.Moldsandcasts

PALEOLECTURE,PAGE4
Moldimpressionofskeletal(orskin)remainsinanadjoiningrock
Externalmold=impressionofouterside
Internalmold(steinkern)=impressionshowsformormarkingsofinnersurface
Castoriginalskeletalmaterialdissolvesandcavity(mold)fillswithmaterial
Endocastnaturalinfillingofcranialcavity(maystudybrainevolutioninfossilmammals)
b.Ichnologystudyoftracefossils(Ichnofossils=tracks,trailsandburrowsoforganisms)
c.Coprolitesfossilexcrementofanimals;maycontainundigestedremainsoffood
F.Pseudofossils
manyrocksandrockstructureslooklikefossils,butaren't!
thefollowingrepresentafewsedimentaryfeaturesthatmaybeconfusedforfossils:
1.DifferentialWeathering
weatheringofrockandmineralsurfacesoftenyieldfossillookingfeatures
2.Nodules
formedbyfillingvoidsinthesedimentandincorporationofsedimentarymaterialswithinthe
sedimentarybody
a.ChertNodules
microcrystallinequartz;typicallyfoundalongbeddingplanesinlimestone
b.Septaria
largenoduleswithradialandconcentriccracksintheircenters
Melikariaareboxworkpatternsofmaterialfillingseptariancracks;maybeallthatisleftafter
weatheringoftheseptaria
c.Rosettes
radiatingmacrocrystallinebodiesofdiscoidalorsphericalshape,consistingessentiallyofone
mineral(typicallypyrite,marcasite,barite,orgypsum)
3.Concretions
mineralgrowthwithinsedimentoftenformsstructuresthatresemblebones,turtleshells,logs,
etc.
4.Dendrites
precipitationofmanganeseoxidealongbeddingplanescreatesfernlikepatterns

PALEOLECTURE,PAGE5

II.Rocks,FossilsandAges
A.BiasesoftheFossilRecord
certainenvironmentsandprocessespreferentiallypreservefossils;collectingtechniquesare
alsobiased
1.HardPartssoftbodiedorganismsrarelyfossilize
2.Preferentialenvironments
thosewithrapidburial
a.Aquaticenvironmentspreferentiallypreserved;especiallyshallowwatersofcontinental
marginsandinlandseas,deltas,lagoons,rivers(especiallyfloodplains),coalswampsandlakes
typicallylowerenergy(finergrainsize)environmentswithbestpreservation;Ex.=limestone,
shale,siltstone,chert
b.cavesandfissurefillingsalsogoodforpreservation
c.KonservatLagersttten
fossillocalitiesexhibitingexceptionalpreservation
fossilizationoftentakesplaceunderanaerobicconditionsand/orwithinfinegrainedsediments
someexamplesincludetheBurgessShale(Cambrian;BritishColumbia,Canada),Mazon
Creek(Pennsylvanian;Illinois),SolnhofenLimestone(Jurassic;Germany),Messel(Eocene;
Germany)andLaBrea(Pleistocene;California)
3.PreservationalBiases
mostfossilsareknownfromspeciesthatwerecommon,widespreadandlonglived
4.CollectingBiases
a.bestfossilhuntingisofteninerosionalareassuchas"badlands"ofdesertsandsemiarid
areas(whereyoucanseethefossils)
b.collectingtechniquesmaybebiasedtolargeanimals,smallanimalsoranimalsfromcertain
paleoenvironments
B.GeologicTime
1.RelativeDatingTechniques
sequencegeologicevents
a.Biostratigraphyseesectionbelow

PALEOLECTURE,PAGE6

b.Lithostratigraphycorrelationbasedonrocks
CorrelationisoftenAccomplishedbyUseof:
b1.Key(Marker)Bedsdistinctivebedwhichisnearlythesameageeverywhere;Exs.=
volcanicash,tillite
b2.Unconformitiesdepositsrestingonunconformities(erosionalsurfaces)areofsimilar
age;often"global"unconformitiesareduetomarineregressions[Eustatic(worldwide)lowering
ofsealevel];unconformitiescanbelocatedinsubsurfacebyseismicsurveys
c.FormalLithostratigraphicUnitsrockstratigraphicunits
c1.Formationfundamentalrockstratigraphicunit;withmappabilityandlithologicconstancy
c2.Membersubdivisionofformation;maybemappedlocally
c3.Groupcontainsseveralformationsunitedonbasisofsimilarcharacteristics
c4.Supergroupcomposedofseveralgroups
2.Absolute(Actual)DatingTechniques
yieldsdatesinyears
a.Radioactivity
a1.Isotopesformsofanelementwithsamenumberofprotons,differentnumbersofneutrons
a2.RadioactiveDecayatomschangetoanotherelementbyreleasingsubatomicparticlesand
energy;parentisotopedecaystodaughterisotopeataconstantrate
a3.RadiometricDatingmeasureamountofparentmaterialsrelativetotheirdaughter
products
HalfLifetimerequiredforisotopetodecaytohalfitsoriginalamount
inpaleontologyoftenusepotassiumargon(especiallyonvolcanicrocks)andCarbon14(for
Pleistocene/Holocenedeposits)
Notation:
Kiloannum(plural=Kiloanna;kiloan)=Ka=thousandsofyearsintheradioisotopictimescale
Megannum(plural=Meganna;megaan)=Ma=millionsofyearsintheradioisotopictime
scale;M.Y.(orm.y)=millionsofyears,withoutreferencetotheradioisotopictimescale

PALEOLECTURE,PAGE7
Gigannum(plural=Giganna;gigaan)=Ga=billionsofyearsintheradioisotopictimescale
b.MagneticStratigraphy
b1.Earth'sMagneticFieldduetomotionsofliquid,ironrichoutercore(behaveslikebar
magnetwithnorthandsouthpole)
b2.MagneticReversalreversalofpolarityinearth'smagneticfield;isrecordedinironrich
igneousandsedimentaryrocks(NormalInterval=polaritysameastodays;ReversedPolarity=
polarityoppositetotodays)
b3.haveconstructedPaleomagneticPolarityScalebasedonmagneticreversalsand"tied"with
absolutedates(Ex.=Text,p.34)
Chrons=largerintervalsdefinedbymagneticstratigraphy
3.ChronostratigraphicUnitsbodyofrockrepresentingaparticularintervaloftime
TimeUnit ChronostratigraphicUnit
Eon
Eonathem
Era
Erathem
Period
System
Epoch
Series
Age
Stage
D.GeologicTimeScale
learnTimeScale(LastPageofLectureNotes)
E.Biostratigraphy("StratigraphicPaleontology")
1.Biostratigraphicdistributionsarecontrolledby:
a.Evolution
b.Paleoecologynoorganisminhabitsallenvironments
b1.Faciescontrolledorganisms=restrictedtoparticularsedimentaryenvironments(often
withslowevolutionarychange)
b2.Biofacies=faciesdistinguishedonthebasisoftheirfossils(Ex.=reefbiofaciesmay
havecorals,corallinealgae,stromatoporoids,rudistbivalves)
2.BiostratigraphicUnits
bodyofrocksdelimitedfromadjacentrocksbytheirfossilcontent

PALEOLECTURE,PAGE8
oftenusefossilsforCorrelation(matchingstratigraphicsectionsofthesameage)
a.Firstappearancesoffossilsmaybedueto1)evolutionaryfirstoccurrence2)immigration
FAD=FirstappearancedatumFOD=Firstoccurrencedatum
b.Lastappearanceoffossilsmaybedueto1)extinctionevent2)emigration
LAD=LastappearancedatumLOD=Lastoccurrencedatum
c.Biozone
basicunitofbiostratigraphicclassification
basedonthedistributionofIndexFossils(fossilscharacteristicofkeyformations;shouldhave
shorttimespan,widegeographicrange,independentaspossibleoffacies,abundant,rapidly
changingandwithdistinctivemorphology)
TypesofBiozonesInclude:
c1.AssemblageZonesstratagroupedtogetheronthebasisofanassemblageofforms
OppelZoneintervalofcommonoccurrencesofalloraspecifiedportionofthetaxa
Mammalagegeochronologicunitbasedonanassociationoffossilmammalsconsideredto
representaparticularintervalofgeologictime;importantforcorrelatingCenozoicfossil
vertebratefaunasworldwide
c2.RangeZonesplotstratigraphicrangeoffossil(s)
Teilzone=partial,localrangezone
TaxonRangeZone(Acrozone)totalhorizontalandverticalrangeofataxon
Concurrentrangezoneoverlappingrangesofspecifiedtaxa
TaxonandConcurrentRangeZonesaremostimportantrangezones
c3.AcmeZone(PeakZone,AbundanceZone)
groupedtogetherbecauseofabundanceofcertainforms
3.MajorFossilsusedinBiostratigraphy
bestarepelagic[planktonic(floating)ornektonic(swimming)]forms
a.Macrofossils
ammonites(PermianandMesozoic)
landmammalsandplants(Cenozoic)
b.Microfossils

PALEOLECTURE,PAGE9
mostimportantincludeforaminiferans,radiolarians,palynomorphs(pollen,spores,
dinoflagellates,acritarchs,calcareousnannoplankton),conodonts
4.QuantitativeBiostratigraphy
usestatisticstocomparethedegreeofsimilaritybetweenfossilfaunas
useSimilarityCoefficientsincluding:
a.SimpsonCoefficient=C/(N1+N2)
b.JaccardCoefficient=C/(N1+N2C)
c.DiceCoefficient=2C/(N1+N2)
d.OtsukaCoefficient=C/squarerootofN1N2
WHERE:C=numberofitemsincommon
N1=numberofspeciesinthesmallersample
N2=numberofspeciesinthelargersample
Thelargerthevaluesofthecoefficientscalculatedfromtwofaunaswhencompared,thecloserin
agetheyareconsideredtobe.Butitisdifficulttocorrelatequantitativelywithoutdetermining
therelativevalueoftheindexfossils!
III.ContinentsHaveMovedandClimatesHaveChanged
A.Paleobiogeography
studyoftheancientgeographicdistributionoforganisms
1.Differencesindistributionaredueto
a.Barrierstoorganismdispersalincludephysicalbarriers(Ex.=landandwaterbarriers)and
environmentalbarriers(i.e.latitudinalandtemperaturechanges)
b.HistoricalFactorsevolutionofdifferentorganismsindifferentregions,etc.
2.AncientFaunalProvinces
oftenclassifiedinmodernecosystemsonthebasisofthenumberofendemicspecies(=
organismsconfinedtoonebiogeographicunit)
a.FaunalRealmlargestbiogeographicalunit;over75%endemicspecies
b.FaunalRegionbetween50%and75%endemics

PALEOLECTURE,PAGE10

c.FaunalProvincebetween50%and25%endemics
d.FaunalSubprovincelessthan25%endemics
theseclassificationsnottypicallyusedforfossilassemblages
3.InfluencesofPlateTectonics
oneofmajorpiecesofevidenceforthepresenceofsupercontinentswasthecommon
distributionofplants(EX.=Glossopterisflora)andanimals(theaquaticreptileMesosaurus)on
the"Gondwanacontinent"
a.ClosingOceans
convergentplatemarginsoftencausegreatersimilarityoforganisms
b.OpeningOceans
continentalfragmentationoftenleadstofragmentationofrangesoforganismsandincreasing
evolutionarydissimilaritythroughtime
c.AccretedTerrains
accreted(suspect)terrainsarecausedwheremicrocontinentssuturetoothercontinentalplates
individualaccretedterrainsareoftenrecognizedbytheirdistinctive(exotic)fossilfaunas
d.VicarianceBiogeography
moderndistributionoforganismsislargelydueto"vicariating"(fragmenting)therangesof
organisms(duetoplatetectonics,iceages,etc.)
B.Paleoecology
1.Ecologystudyofthefactorsthatgovernthedistributionandabundanceoforganisms
2.Paleoecologytherelationshipsbetweenspeciesrepresentedinthefossilrecordandthe
environmentsinwhichtheyinhabited
3.Taphonomyallaspectsofthepassageoforganismsfromthebiospheretothelithosphere
a.TaphonomicProcesses
a1.PhysicalProcesses
examplesincludemechanicalbreakdownoforganicmaterialbywavesandcurrents,andburial
bysediments
a2.ChemicalProcesses
examplesincludealterationofshellmineralogy,andleachingofshellsandskeletonsby
groundwater

PALEOLECTURE,PAGE11

a3.BiologicalProcesses
examplesincludedestructionofhardpartsbyscavengers,andbreakdownofskeletonsbythe
actionoforganisms(boringalgaeandsponges,etc.)
b.TaphonomicBiasescertainenvironmentsandtaphonomicprocessespreferentiallypreserve
fossils;collectingtechniquesarealsobiased
b1.PreferentialenvironmentsAquaticenvironmentspreferentiallypreservefossils
b2.PreservationalBiasesmostfossilsareknownfromspeciesthatwerecommon,
widespreadandlonglived
b3.TimeAveragingfossilassemblageswillbelesssimilartothelivingcommunitythe
greaterthetemporalvariationofthelivingcommunityandthelongerthetimeaveragedinthe
fossilassemblage
b4.CollectingBiasesbestfossilhuntingwhererocksandsedimentsareexposed;canavoid
biasesbybulkcollectingmatrixandestimatingproportionsoffossilsbyconstructingquadrants
orlinetransects
c.SizeDistributionoftenfossilsaresizesortedduetocurrentaction;MicromorphFaunas
consistofunusuallysmallindividualsofspecieswhosesizeisduetounusualenvironmental
factors
4.SedimentaryEnvironmentsportionoftheearth'ssurfacewithdistinctivephysical,chemical
andbiologicalcharacteristics
a.Faciesbodyofsedimentorrockswithdistinctivecharacteristics
FaciesModelssummaryofspecificsedimentaryenvironments
b.Walther'sLawtheverticalsequenceofrocksmayreflectthehorizontalsuccessionof
environments/facies
5.BiologicCriteria
mustbecautiousinenvironmentalinterpretationsbasedonfossils
modernecosystemsarecharacterizedbybiocoenoses("lifeassemblages");paleontologistsfind
primarilythanatocoenoses("deathassemblages",ortaphocoenoses)
a.Habitatsenvironmentsinhabitedbylife
b.Speciesrelationships
Ecologicalnichewayinwhichaspeciesrelatestoitsenvironment

PALEOLECTURE,PAGE12
c.EcologicCommunitypopulationsofseveralspecieslivingtogetherinahabitat
paleontologistsdonotobservefossilcommunities("paleocommunities");whattheyobserveare
assemblagesoffossils(fossilsthatoccurtogetherrepeatedlydefinefossilsassemblages)
c1.Ecosystemorganismsandtheirphysicalenvironments
Faunaanimalsofanecosystem
Floraplantsofanecosystem
Biota=flora+fauna
c2.Diversity
numberofspeciesthatlivetogetherinacommunity;tropicalclimatescontainmorediverse
plantandanimalcommunities
Diversity=numberofspecies/numberofspecimens
c3.Foodchainssequenceofnutritionalstepsinanecosystem
TrophicLevelpositioninfoodchain;organismsfromlowertrophiclevelshavemorepotential
forfossilizationthanthosefromhighertrophiclevels(becauseorganismsfromlowertrophic
levelsaremorenumerous)
c4.Foodwebsnutritionalstructureofecosysteminwhichmorethanonespeciesoccupies
eachlevel
Competitiontwospeciesvieforlimitedenvironmentalresources
Autotrophs(Producers)=manufacturetheirownfood;"plants";formlowesttrophicleveland
constitutethebaseofthebiomass"pyramid"

Heterotrophs(Consumers)=feedonotherorganisms;consistof"animals"(muchenergyislost
cyclingthroughhighertrophiclevels,andthereforewithfewerorganisms)
Herbivores=feedonproducers
Predation=effectofapredatoronapreyspecies
Carnivores=feedonotherconsumersbypredation
Parasites=derivenutritionfromotherorganismswithoutkillingthem
Scavengers=feedondeadorganisms

PALEOLECTURE,PAGE13
Commensalism=biologicalassociationbeneficialtoonebutdoesnothurtthehost
Symbiosis=mutualbenefittobothparticipants
c5.Succession=changesduetomodificationoftheenvironmentbyorganisms
StagesofSuccessionInclude:
PioneerStage:withabundant,rapidgrowing,shortlivedspecieswithabundantoffspring(r
strategists)
MatureStage:withthemostdiversity
ClimaxStage:slowergrowing,larger,longerlivedspecieswithfeweroffspring(Kstrategists)
replaceorganismsofearlierstages
6.Limitingfactors
environmentalfactorscontrollingspeciesdistribution
includeschemical,physicalandbiologicalfactors
OrganismDistribution(EspeciallyMarine)DependsUpontheFollowing:
a.SeawaterPropertiesDensityandViscosity
Densityofaquaticorganismstypicallyequalswaterdensity
Viscosityinfluencesshapeandfeeding(therearemany"filterfeeders"inaquaticenvironments,
duetotheviscosityofwaterallowingfoodtobeheldinsuspension)
b.Salinity
usuallymeasuredinpartsperthousand(0/00);averageseawatersalinityis350/00butvaries
from0to2700/00
inGeochemicalStudiesofPaleosalinityuseboron(greaterinsaltwater);othertraceelements;
typeoforganicmatter;carbonandoxygenisotopes[freshwatersdepletedinheavycarbon
(C13)andheavyoxygen(018)]
inBiologicalStudiesusestenohaline(restrictedbysalinity;organismsinternal"salinities"
equalssurroundingwatersalinity;ifrapidchangecellsmaynotfunction)versuseuryhaline
(salinitytolerant)organisms
c.Temperature
watermoderatestemperature
incoldbloodedorganisms,anincreaseintemperatureof10Coftencausesmetabolicactivity
todouble
inwarmbloodedorganismsthereislittlemetabolicchangewithtemperaturechange
temperatureinfluencesreproductivecycles
inGeochemicalStudiesofPaleotemperatureuse18O/16O(lesswithgreatertemperature;most

PALEOLECTURE,PAGE14

importantfordeterminingpaleotemperatures);boronandbrominegreaterifgreater
temperature;Calcium/MagnesiumandCalcium/Strontiumratiosarelessifthetemperatureis
increased
inBiologicalstudiesofpaleotemperatureusestenothermal(temperatureintolerant)versus
eurythermal(temperaturetolerant)organisms;alsomaylookatspeciesdiversity(greaterin
warmerenvironments)ormorphology(bodyformreflectsenvironmentalfactors)
d.DissolvedGases
concentrationsdependonatmosphericconcentration;solubilityofgas;watertemperatureand
salinity
d1.Nitrogen(N)mostabundantdissolvedgas;requiredbyplantsinionicform
d2.Oxygen(O)entersseabyphotosynthesis,riverwater,atmosphere;allorganismsuse
oxygenduringrespiration;oxygenatmaximumnearsurface,minimumatabout700
1,000mOxygen;approximately6to10ppm;warmer,saltierororganicdebrisrichwater
withlessoxygen
d3.CarbonDioxide(CO2)entersseafromorganismrespiration,atmosphereandrivers;
removedbyplantsforphotosynthesisandusedbyorganismstomakeshells;increasesto
approximately1,000m;increasedCO2leadstoGreenhouseEffect(increasetemperature)
d4.HydrogenSulfide(H2S)producedbyanaerobicbacteria
e.Light
Photiczone=zoneoflightpenetration
Euphoticzone=upperilluminatedlayersofwaterinthephoticzone;receivesufficientlight
tosupportphotosynthesis;usually1060metersbutcleartropicalwatersmaybegreater
than100meters
Aphoticzone=zoneinwhichlightdoesnotpenetrate
f.Pressure
pressureincreasesapproximately1atmosphereper10meters
affectsverticalmigrationoforganisms,bacterialdecomposition,productionofshells(CCD=
carbonatecompensationdepth)
g.Depth
deepwaterstorescarbon,nitrate,phosphate
Paleobathymetrytheancientwaterdepthmaybedeterminedbytypeofbodyfossilsandtrace
fossilspresent
h.Waterenergy,turbidityandsedimentationrates

PALEOLECTURE,PAGE15
affectsdistributionoffoodandnutrients;typesandmorphologyoforganismspresent
amountofsuspendedsedimentespeciallyaffectsfilterfeeders
natureofsubstrateaffectstypeofinfauna(liveinsubstrate)orepifauna(liveonsubstrate;
sessileorvagilebenthonic)present
7.Paleoclimatology
studyofancientclimates
utilizessedimentologic,paleontologicandgeochemicaldatatoreconstructancienttemperature,
windpatterns,precipitationandevaporation
8.ChemicalCyclesinEarthSystemHistory
ChemicalReservoirsbodiesofkeyelementsandcompoundsintheEarthsystemthatshrinkor
expandasfluxesbetweenthemchange
thesereservoirsareinfluencedbythefollowing:
a.PhotosynthesisandRespiration
Photosynthesisprocessbywhichplantsusetheenergyofsunlighttoproducesugarsfrom
carbondioxideandwater;oxygenisabyproductofthisprocess
Respirationoppositechemicalreactionversusphotosynthesis;organismsoxidizesugarsin
ordertoreleasetheirenergy
b.CarbonDioxideandOxygenCycles
ifnodeadplanttissueisburied,itdecomposesandcarbondioxidereturnstotheatmosphere
ifdeadplanttissueisburied(suchasinswampsoranoxicmarineenvironments),itupsetsthe
balancebetweenphotosynthesisandrespiration(withtheamountofcarbondioxideinthe
atmosphereshrinkingandwithincreaseinoxygenlevels)
weatheringofmineralsremovescarbondioxidefromtheatmosphere(enhancedbymountain
building,warmclimates,highratesofprecipitation,andmorevegetation)
theinitialspreadofforestsduringtheDevonianintensifiedweathering,depletedthe
atmosphericreservoirofcarbondioxide;thisreducedgreenhousewarmingandprobably
contributedtothecoolerclimateconditionsandformationoftheLatePaleozoic"IceAge"
c.MethaneCycles
methaneisapowerfulgreenhousegas
whenglobalwarmingmeltsmassesofmethanehydrateontheseafloor,theadditionofmethane
totheatmosphereproducesfurtherglobalwarming
d.NegativeFeedbackinCarbonDioxideandGlobalWarmingCycles
whenclimatewarms,chemicalweatheringaccelerates(extractingcarbondioxidefromthe
atmosphere)andtheamountofevaporationincreasesontheocean(whichfurtheraccelerates
weatheringonland,extractingmorecarbondioxidefromtheatmosphere)

PALEOLECTURE,PAGE16

e.SubmarineVolcanismversusSeawaterChemistry,MineralogyandTypesofOrganisms
seawatercirculatingaroundmidoceanicridgestransferscalciumtotheseawater;magnesiumis
extractedfromthewaterandbecomeslockedintherocks[thereforewithmoreseafloor
spreadingthereisariseinsealevel(morerocksproduced)andadecreaseinthe
magnesium/calciumratio(moremagnesiumextractedfromseawater)]
withincreasedmarinevolcanism,thelowmagnesium/calciteratiosproduce"CalciteSeas",in
whichcalciteformsoolitesandmarinecementsandorganismswithcalciteskeletonsbecome
successfulreefbuilders(thelowestmagnesium/calciteratioofthePhanerozoicwasduringthe
Cretaceous,whichcontainsmuchmore"chalk"thananyothersystem)
whenthetotalvolumeofvolcanicsatmidoceanicridgesislow,aragoniteandhigh
magnesiumcalciteismoreabundant;"modern"typesofcorals,witharagoniteskeletons,are
moreabundantduringtheseperiodsofEarthhistory
IV.Groups,NamesandRelationships
Taxonomyprocessofclassificationandnamingorganisms;typicalclassificationoforganisms
isbytheirrelationshiptooneanother(="natural"classification)
Systematicsgroupingorganismsaccordingtotheextenttowhichtheyarerelated
theclassificationoforganismshastraditionallyusedtheLinnaeanSystem,formulatedby
CarolusLinnaeusinthe1700's(Manybiologistsandpaleontologistarenowabandoningthe
LinnaeanSystem,duetotheinfluenceofCladisticTaxonomy)
A.Taxa(singular=taxon)intheLinnaeanSystem
1.Domain
insomerecentclassifications,constitutesthehighesttaxonomiccategory
oftenincludetheDomainsArchaea/Archaebacteria,Bacteria/Eubacteria,andEucarya
2.Kingdom
inmanyclassificationsisthehighesttaxonomiccategory
therearetypically5to6recognizedkingdoms[Monera(oftenclassifiedasDomainor
KingdomArchaea/ArchaebacteriaandDomainorKingdomBacteria/Eubacteria);Domain
EucaryaincludestheKingdomsProtoctista(Protista),Fungi,Animalia(Metazoa),andPlantae
(Metaphyta)]
3.Phylum
4.Class
5.Order
superordersoftenendinica,ordersinida(formesinmanyvertebrateorders)andsuborders
inina

PALEOLECTURE,PAGE17

6.Family
superfamiliesoftenendinoidea,familiesinidaeandsubfamiliesininae
ToconvertLatinnamestoEnglishusesuffix"id";ifitresemblesagroupuse"oid"
7.Genus
groupofinterrelatedspecies;plural=genera
8.Species
fundamentalunitoftaxonomy
Binomen=genus+speciesnames;theremayalsobesubspecies
a.Therearetwoconceptsofwhatconstitutesaspecies:
a1.BiogeographicGeneticSpecies=populationofindividualsthatcaninterbreedand
produceviableoffspring;arespatiallysegregatedandgeneticallyisolatedfromsimilaradjacent
groups
a2.LinnaeanSpecies=basedondiscontinuityintherangeofvariationintheform
(morphology)oforganisms;istypicallyusedinpaleontologicalstudies
B.NamingaTaxon
useInternationalRulesofZoologicalorBotanicalNomenclature
1.Eachgenus+specieshasanameindependentofchange;namesareLatinized
2.Eachgenus+specieswillhaveseparatenames
Nogenusnamecanbeduplicated
3.Differentnameswillnotbeappliedtoonespeciesorgenus
Synonym=twoormoredifferentnamesgiventothesameanimal;useLawofPriorityto
determinewhichiscorrect
4.Eachgenus+speciesmusthaveaType(=primarynamebearer)
a.Typespeciesofagenuscomposedofoneofitsspecies
b.Typespecimensofspeciesisaparticularspecimen
Holotype=singlespecimenservesasthenamebearer(isnowarule)
Paratypes=otherspecimenswhichservetocharacterizeaspecies

PALEOLECTURE,PAGE18
5.Authorcitations
a.Citeauthorsnameaftertaxonandfirstyearofpublication
b.Ifauthor'sspeciestransferredtoanothergenusthenplaceoriginalauthor'snamein
parentheses
C.TheoriesofTaxonomy
whattaxonomistsseekistheestablishmentofMonophyleticGroups(speciesthatsharea
commonancestrythataregroupedtogethertaxonomically;thegroupingthatincludesthe
ancestoranditsdescendantsisoftentermedaClade)
taxonomistswanttoavoidPolyphyleticGroups(groupevolvedfromtwoormoredistinct
ancestors)andParaphyleticGroups(groupswithacommonancestrybutwithoneormore
descendantgroupsexcluded;includemanymoderngroups)
1.EvolutionaryTaxonomy/Systematics
thisisDarwin'staxonomy
youmustclassifypopulations("taxa")ratherthanindividualsorcharacters
firstyouclassifyaphenotype(thecharacteristics)andthenyouinferthegenotype(itsgenetic
constitution)
2.Phenetics(NumericalTaxonomy)
classifyorganismsbypurelymechanicalormathematicalmeans
selectagroupofcharacterstodescribetheOperationalTaxonomicUnit(OTU),constructa
similaritymatrix,andthendisplaytheresultsasaphenogramordendrogram
Butisita"natural"classificationsystem?
3.CladisticTaxonomy(PhylogeneticSystematics;HennigianSystematics)
formulatedbyentomologistWilliHennig
Hennigstatedthatprimitivecharactersareofnouseintaxonomy(a"character"isanydefinite
aspectofaparticularorganism)
Thelargerthenumberofnonprimitive("sharedderived"orsynapomorphic)charactersshared
bytwosubgroupsthemorecloselyrelatedthesubgroups(character=anyrecognizabletraitof
anorganism;CharacterState=presenceorabsenceoftraitoralternativewaysinwhicha
charactermaybeexpressed)
a.TypesofCharacters
a1.Discretecharactershavealimitednumberofpossiblevalues
BinaryCharactershavetwostates(Ex.=presenceorabsenceofafeature)
MultistateCharactershavemorethan2states(Ex.=blue,greenandbrowneyecolor)

PALEOLECTURE,PAGE19
a2.ContinuousCharacters
measurementsonacontinuousscale(Ex.=sizeoforganisms)
continuouscharactersarehardtouseandinterpret
b.TypesofPhylogeneticTraits
b1.Plesiomorphy=primitivetrait
Symplesiomorphy=sharedprimitivetrait
b2.Apomorphy=specializedorderivedtrait
Autapomorphy=specializedtraituniquetoonegroup
Synapomorphy=specializedtraitsharedbytwoormoregroups
c.SisterGroups
twogroupsunitedbythepresenceofoneormoresynapomorphiccharacters
SisterGroupsshareacommonancestorandareeachother'sclosestrelatives
d.Polarity
directionofevolutionarychange
isdeterminedbythestratigraphicsequence,ontogeny(organismslifehistory,especially
embryologicaldevelopment),outgroupcomparison[charactersarederivediftheydonotappear
incloselyrelated"outgroups"(i.e.taxathatarenotpartofthegroupunderconsiderationbut
shareacommonancestry)],characteranalysiswithingroupsandanatomicalprogression
(Morphocline=charactersthatvaryquantitativelywithinagroup;ex.=bodysize,limb
proportions)
e.Parsimony
thebesthypothesisistheonewiththefewestnumberofprocesses(themajorityofcharacters
rule;also,chooseandweightthecharactersof"mostimportance")
f.Cladogram
diagramthatdepictsrecentnessofdivergenceofsubgroupsbasedonthenumberofshared
derivedcharacters(butisnotbasedontheabsolutetimescale)
g.Whysometaxonomiststhinkcladistsarefullofbull:
Cladiststendtoignorestratigraphyofthefossilrecord
itisverydifficulttoassessrelationshipsfromtheinitialstagesofmajoradaptiveradiations
(plesiomorphic,autapomorphicandsynapomorphiccharactersaredifficulttodifferentiate)
CladistshopethatwhattheydealwithisDivergentEvolution(evolutionarydiversification)and
thatevolutioncannotbereversed(Dollo'sLaw);Dollo'sLawmaytypicallybetrue(butitis
toughtojudge)

PALEOLECTURE,PAGE20

Therearetypesofevolutionthatmayconfusecladists:
ParallelEvolution=twocloselyrelatedorganismsundergoasimilarevolutionarychange
throughtime
ConvergentEvolution(adaptiveconvergence)=closemorphologicsimilarityarisesbetween
twounrelatedgroupsthattakeonsimilarlifehabits(Homeomorphs)
D.MolecularPhylogeny
sequencesofbasesinchainsofgeneticmaterial(DNAandRNA)andaminoacidsequencesof
proteinsarediagnosticforeachorganism(theclosertherelationshipsthegreaterthegenetic
similarities)
randommutationssubstitutevariousaminoacidsinmoleculesthatismoreorlessdirectly
proportionaltotime(thereforetheremaybe"MolecularClocks")
maycomparetheproteinsinpreservedtissuesuptoorcomparetheproteinsoflivingformsto
determinetheirrelationships(latterismostoftenused)
E.AdaptationandFunctionalMorphology
1.Adaptation
the"fitness"ofanorganism;howorganismscopewithchangingenvironmentalconditions,
invadenewenvironments,andfunctionmoreefficientlyinagivenenvironment
a.Whyadaptationoccurs:
a1.Musthavefeaturesthatallowstheorganismtosurviveinparticularenvironments
a2.RedQueenHypothesisformulatedbyLeighvanValen;becauseofcompetitionfor
resourceswithotherorganisms,aspeciesmustcontinuouslyimproveitsadaptationorbecomes
extinct(eveninstableenvironments)
b.EvidenceforAdaptation
b1.Morphologybodypartsofanorganismare"designed"tofittheenvironmentinwhich
theylive
b2.ConvergentEvolutionindependentevolutionofsimilarmorphologiesthatfunction
similarly
c.Organismsarenot"completely"adaptedtotheirenvironmentsbecause:
c1.theyareconstrainedbytheirancestry
c2.therearemultipleusesoforgansbutinchangingenvironmentsthisisnotallbad!!!!

PALEOLECTURE,PAGE21
2.FunctionalMorphology
thestudyoftherelationshipofformtothefunctionsthatorganismortheircomponentparts
perform
a.Morphology(form)iscontrolledby:
a1.Adaptationorganismsmorphologyisrestrictedbyenvironmentalconstraints
a2.Phylogenyorganismsmorphologyisrestrictedbytheirevolutionaryhistory
a3.Growth
IsometricGrowthallpartsgrowatthesamerate
AllometricGrowthsomepartsgrowfasterorslowerthanothers(foundinmosthigher
organisms)
b.StudyingMorphology
b1.TheoreticalMorphology
oftenusesaParadigm(modelorpattern);computerprogramsarehelpfulforstudyingawide
varietyofparameters
b2.ComparisonwithLivingOrganisms
isveryhelpfulbutassumesthatmodernorganismshavesimilarmorphologicalfeaturesand
constraintsastheirfossil"counterparts"(sometimesabadassumption!!!)
F.MechanismsofEvolution
Evolution=(1)historicalchangesinstructure,functionandadaptation(2)geneticchangesand
processesofselectionandpopulationdynamics
1.HistoryofEvolutionaryTheory
a.JeanBaptisteLamarck(17441829)Frenchnaturalist;developedtheorynowknownas
Lamarckism(theoryofinheritanceofacquiredcharacteristics)
b.ThomasMalthus(17661834)Englishclergymanandeconomist;wrote"Essayonthe
PrincipleofPopulation";introducedconceptthatpopulationexhibitsexponentialgrowth,
whereasfoodproductionexhibitslineargrowth;populationexpandstolimitssetbyfamine,war
anddisease
c.AlfredWallace(18231913)codiscovererofthetheoryofnaturalselectionindependentof
Darwin;alsoaprominentzoogeographer
d.CharlesDarwin(18091882)mostnaturalistsofhistimewere"specialcreationists";as

PALEOLECTURE,PAGE22
ship'snaturalistontheH.M.S.Beagle(18311836)developedthefoundationofhistheoryof
evolution;ReadMalthus'EssayonPopulation;Wrote"TheOriginofspeciesbymeansof
NaturalSelection"in1859
Darwin'sfactsanddeductionsinclude:
organismstendtoincreaseinnumbersbyageometricratio
inspiteofthetendencytoprogressiveincreasethenumberofindividualswithinaspeciestend
toremainapproximatelyconstant.
Deduction:Sincemoreyoungareproducedthancansurvivetheremustbeacompetitionfor
survival("StruggleforExistence")
allorganismsvary;somevariationsareinherited
someindividualsfailtosurvive,otherslivetoreproduce(naturalselection)
SummaryofDarwinianEvolutionaryTheory:Newspeciesarisefrompreexistingonesasa
resultofnaturalselectionactingoninheritedvariations
e.TheSyntheticTheory(Neodarwinism)
developedinthe1930'sand1940'sbyTheodosiusDobzhansky,ErnstMayr,GeorgeSimpson
andJulianHuxley
statedthatthedeterminantsoftraitsonwhichnaturalselectionactsaregenes(heritableunitsof
informationgoverningstructure,developmentandfunction)
variationisduetogenemutation(alsonowtheorizedthattandemmultiplicationofnucleotides
andgeneduplicationisimportant)
alsopopulationstructureanddistributionisimportantinthedevelopmentofnewspecies
NaturalVersusSexualSelection
naturalselectiondependsonfitnessoftheorganismtoitsenvironment
sexualselectiondependsonattractivenesstofemales(onemayworkagainsttheother;Ex.=
elaborateplumageofbirds)
f.TheNeutralTheory
developedinthelate1970'sbyMotooKimura
statesthatmostgeneticdifferencesneitherfosternorhinderanorganismssurvivalandtheir
persistenceoreliminationinapopulationisamatterofchance
geneticdifferenceshaveanadaptiveeffectthatsuppliesabundantrawmaterialforthecreative
forceofnaturalselection
g.PunctuatedEquilibriumversusPhyleticGradualism
g1.Phyleticgradualism
ratesofevolutionareregular
isDarwinianEvolutionaryTheory
g2.Punctuatedequilibrium

PALEOLECTURE,PAGE23
firstproposedbyNilesEldredgeandStephenGouldduringthe1970's
saysthatevolutionoccursinfitsandspurtsseparatedbylongperiodsoflittlechange
problemintesting(suddenappearancesinfossilrecordmaybeduetoimmigrationratherthan
rapidspeciation)
probleminclassification(noclassificationsystemcanshowintermediateforms(only
"species")
h.MosaicEvolution
firstproposedbyGavindeBeer
differentpartsoforganismsdonotchangeatauniformrateinthecourseofevolution
Iinterpretitasthemiddlegroundbetweenpunctuatedequilibriumandphyleticgradualism
2.EvolutionaryChanges
a.EvolutionaryChangeisbymeansof(?):
a1.PhyleticEvolution(Anagenesis)processbywhichasinglelineagechangesovertime
a2.Speciation(Cladogenesis)processbywhichasinglespeciesdividesintotwolineages
thatbecomereproductivelyisolatedfromoneanother;preferredbyproponentsofpunctuated
equilibrium
Cladeallorganismsdescendedfromaprogenitorspecies
b.PeripatricSpeciation
Evolutionarychangesaremostlikelywithinsmall,geographicallyisolatedpopulations
(thereforemostimportantgroupsforestablishingevolutionarypatternsandprocessesareleast
likelytobepreserved)
inbreedingleadstohomozygosity(withrecessiveanddominanttraitscommonandtherefore
potentiallyrapidchange)
ifanisolatedpopulationrejoinswiththeparentpopulationagain(sympatry),theymustdiverge
behaviorallyand/orphysicallyevenmoretosurvive
3.MicroevolutionversusMacroevolution
a.Microevolution=processofevolutionwithinasinglelineage
involvesnaturalselectionbetweenindividuals
b.Macroevolution=processofevolutionbetweengroups
involveshighertaxonomiccategories
b1.SpeciesSelectiondifferentialsurvivalamonganumberofspeciesthatdescendedfroma
commonancestor;ifsignificantchangestakeplacethanassumetherewerealargenumberof
speciesinvolvedinthatcompetition
Competitionbetweengroupsisdemonstratedbyconvergentevolution,mimicry(oftennon

PALEOLECTURE,PAGE24
poisonoustypesmimicpoisonousones)andprotectivecoloration(organismscamouflagedto
resembleenvironment)
Evolutionary"ArmsRace"=competitionbeweenpredatorsandpreyleadstosequenceof
evolutionarychangesinbothgroups
b2.TrendsinMacroevolution
initiallywithlownumberoflineagesandabsenceofcompetition(notmuchdifferencebetween
species)
featuresbecomestabilizedandarerecognizedascharactersoffamiliesororders
adaptiveradiationmaygiverisetotremendousincreaseindiversityduringshorttime
b3.Cope'sLaw=sizeincreaseamongwarmbloodedanimals
4.EvolutionandOntogeny
a.Ontogenylifehistoryofanorganism(especiallyembryologicaldevelopment)
b.Embryosdevelopfromgeneralcharacterstowardsmorespecificones
inearlystagesofdevelopmentanimalembryostendtobeverysimilar
c.Haeckel'sLawontogenyrecapitulatesphylogeny
i.e.,initsdevelopmentfromembryotoadulttheindividualpassesthroughtheevolutionary
stagesofitsancestors(=Peramorphosis)
d.Paedomorphosisearlystagesoftheontogenyofancestorsbecometheadultstagein
descendants(Ex.=someadultamphibianshavelarvalikeform)
G.EvolutioninEarthHistory
1.RatesofEvolutionaryChange
a.isprobablyirregular[Simpson,1944,coinedBradytely(slowchange),Horotely(medium
change)andTachytely(rapidchange)terms]
Darwin=unitformeasuringevolutionarychange;d=changebyafactorofepermillionyears,
whereeisthebaseofnaturallogarithms(verydifferentratesforvertebrates;probably0.02to
400din"natural"settings)
b.AdaptiveRadiation
emergenceofnewstructuresandwaysoflife,ormassextinctions,oftenleadsto"adaptive
radiations"withlargeincreaseinnumberofdaughterspeciesevolving
majorchangesinlifestylesoftenduetorelativelyminormodificationofskeletalfeatures
manyclineswithadaptiveradiationsfollowedbyslowdeclinetowardextinction
c.Fordeterminingevolutionarypatternsandratesatthespecieslevelneed(andisrarely

PALEOLECTURE,PAGE25
accomplished):
sedimentaryrecordcompletewithgapslessthan10,00020,000years
totalsequenceprobablyexceeding100,000years
gooddating(especiallyradiometric)
entirespeciesrangeknownwitharecordofancestralanddescendantspeciesinthearea
geographicalrangeofspeciesknown
significantportionofskeletalanatomyknown
2.TrendsandCycles
a.Trend"unidirectional"changes
Ex.=increaseinoxygencontent
increasedspecializationsoforganismsthroughtime
b.Cycles
repetitivesequences(maygreatlyinfluenceevolution)
b1."Greenhouse"/"Icehouse"Cycles
movingcontinentsandexpansionandcontractionofoceanbasinsleadsto"greenhouse"
(increasedvolcanismproducesmoreCO2,whichleadstoagreenhouseeffectwithgreater
temperatures)
cycles300to500millionyearsinlength
b2.SeaLevelFluctuations
transgressions(relativeriseinsealevel)andregressions(dropinsealevel)mayinfluence
evolution[regressionsmayplacestressesonshallowwatermarineorganisms;"specialist"
organismsusuallywithhighestevolutionaryratesjustaftertheperiodofgreatesttransgression]
Seismicevidencefrompassiveplatetectonicmarginsindicatestherearepatternsofsmall,
mediumandlargescalesealevelcyclesthatproduceunconformities(VailCurves)
b3.AstronomicalCycles
periodsofincreasedbolideimpacts(asteroids,comets)leadtoextinction
3.Extinction
mostlikelyinspecieswithsmallpopulationsandliveinlimitedgeographicareas(population
sizerelatedtotrophiclevelandbodysizewithcarnivoresmostlikelytobecomeextinctand
smallherbivoresleastlikely)
a.TypesofExtinction
a1.BackgroundExtinction
probabilityofextinctionisapproximatelyconstantthroughthelifeofaparticulargroupbut
ratesvaryfromgrouptogroup

PALEOLECTURE,PAGE26
thereforethereisa"normalbackgroundrate"ofextinction
a2.MassExtinction
therearesixPhanerozoicepisodesofmajorextinction
b.Reasonsforextinctioninclude:
b1.Competition
difficulttodetermineevenamonglivingspecies
b2.Predation
alsotoughtotell
b3.EnvironmentalDeterioration
climatechange(Exs.=coolingtrends,dropinsealevel,oxygendepleteddeepoceanwater
risesontocontinentalshelves,violentvolcanism)causesmassextinctions
b4.StochasticProcesses
saysthatoriginandextinctionoforganismsisprobabilistic(likea"flipofacoin")
computerprogramsgenerating"artificial"phylogeniesaremuchlike"natural"clades
Ex.=Extraterrestrialcausesextinctionbyperiodicbolideimpactsorcomets?
b5.Man
importantforpast11,000years(?)
V.Earth'sOldestRemains
A.EarthOrigin
1.AgeoftheEarthbasedprimarilyonExtraterrestrialEvidence:
a.Meteoritesextraterrestrialobjectsthatfalltoearth;mostdatedatapproximately4.6Ga
b.MoonRocksoldestnearly4.6Ga
c.OldestRocksonEartholdestcrustalrocksfromCanadaaredatedat4.04Ga;
metamorphosedsedimentsinwesternAustraliahavezircongrainsdatedat4.4billionyearsold
2.OriginoftheSolarSystemandPlanets
SolarNebulaTheorysolarsystemformedfromcloudofcosmicdust;rotated,becamedisclike
andplanetsaccreted

PALEOLECTURE,PAGE27
3.FormationoftheEarth'sAtmospherewasprobablyby:
a.Outgassingofearth'sinteriorduringArcheantimesbyvolcanicactivity;producedwater
vapor,nitrogen,carbondioxide,etc.
b.Impactbycometaryice
c.PlantPhotosynthesisprovidesoxygen(butprobablywithlittleoxygeninthe
Precambrian)
4.TheOceans
gasescondenseduringEarthcooling;"modern"salinityobtainedinEarlyArchean
theEarlyArcheanoceanwereprobablymuchwarmerthanthatoftodayduetothepresenceof
abundantradioactiveelementsintheEarth'scrustandthe"GreenhouseEffect"
B.Organisms
ordered(i.e.withcellularorganization)livingcreatures
"life"isaseriesofchemicalreactions,usingcarbonbasedmolecules,bywhichmatteristaken
intoasystemandusedtoassistthesystem'sgrowthandreproduction,withwasteproductsbeing
expelled
lifeformspassontheirorganizedstructurewhentheyreproduce
C.OriginsofLife
1.TheEarthDuringtheArcheanEon
equableconditionsforprebioticevolutioncouldhaveexistedonEarthaslongagoas4.4Ga
ArcheanEarthwasdominatedbyoceaniclithospherewithvolcanicislandsandsmall
microcontinents
largeamountsofCO2mayhaveledtoaGreenhouseEffect,withatmospherictemperaturesup
to100Cormore(thereforeNOpolaricecaps;withpermanentlystratifiedstagnantironrich
deepoceanwatersandwindmixedironpoorsurfacewaters)
hotsprings,submarinehydrothermalsystems,andheatedwindmixedlayersoftheoceansmay
havebeenareaswhereprebioticevolutionoccurred
2.OriginsofLife
a.DependsuponthesynthesisofCarbon
oncecarbonissynthesized,allotherbiogenicmoleculesmaybeformed(OrganicMolecules
arecomplex,carbonbasedmolecules)
elementsmostprominentinorganicmoleculesarecarbon,hydrogen,oxygenandnitrogen
b.CellularStructure
b1.Cell

PALEOLECTURE,PAGE28
a"container"filledwithorganicandinorganicmolecules(=Protoplasm);thecellcontains:
b2.Proteins
builtfromaminoacids;proteinsareusedas"buildingmaterials"andforchemicalreactions
b3.NucleicAcids
includesDeoxyribonucleicAcid(DNA)andRibonucleicAcid(RNA);provideinformationfor
thestructureoftheorganismandthemeanstopassonthisinformationinreproduction
DNAcarriesthegeneticcodeofanorganism,providinginformationforitsgrowthand
metabolism;ithastheabilitytoreplicateitselfinordertopassthisinformationonto
subsequentgenerations
RNAhasseveralfunctions(carriesgeneticmessageofDNAtosites;assemblesaminoacids
intoproteins;actsasacatalystforchemicalreactions),andbecauseofthisversatilitywas
probablythenucleicacidpresentwithintheearliestlifeforms(thisearliestecosystemisoften
termedthe"RNAWorld");butRNAwaseventuallyreplacedbyDNAasthegeneticcode(as
DNAisamorestablemolecule)
b4.OrganicPhosphorousCompoundsfoundinsmallamounts;transformlightorchemical
fuelintoenergy
c.TheFormationofProteins
c1.AminoAcids
mixtureofmethane,ammonia,hydrogenandwatervapor(ornitrogen,carbondioxideand
watervapor)inthepresenceofelectricityorultravioletlightleadstotheproductionofamino
acids
somemeteoritesalsocontainaminoacids
productionofaminoacidsmusttakeplaceinananaerobic(devoidoffreeoxygen)environment
c2.Proteins
removingwaterfromaminoacidsyieldsPolypeptides(proteinlikechains)
whenpolypeptidescooltheyformMicrospheres(celllikestructures)
D.KindsofOrganisms
1.Prokaryotes
singlecelledorganismswiththeirDNAlooselyorganizedwithinthecell,arenotboundedbya
membraneintoanucleusandtheylackchromosomes
reproducebysimplenucleardivisionofcells;meiosisabsent
rangefrom0.3to20microns
areoftentermedMonerans
oftendividedintotwoDomains(orKingdoms):

PALEOLECTURE,PAGE29

a.Domain/KingdomArchaea/Archaebacteria
superficiallysimilartoeubacteriabutdiffergreatlyintheirmolecular(especiallyRNA)
sequences
includethemethanogens(tendtobefoundinhighlysalineenvironments),sulfurmetabolizing
bacteriaandsulfatereducingbacteria(foundaroundhydrothermalvents)
probablyincludedtheoldestlifeforms,whichwereprobablythermophilicautotrophs(used
molecularhydrogen,carbondioxideandsulfurcompoundstoproduceenergy,withoptimal
growthattemperaturesfrom70to110C);possibleenvironmentsoforiginincludehotsprings,
heatedoceanwaters,andhydrothermalvents
b.Domain/KingdomBacteria/Eubacteria
containthemostcommonlyrecognizedor"true"bacteriaandcyanobacteria
evolvedboththermophilicautotrophsinheatedenvironmentsandphotoautotrophsinshallow
marineenvironments(seediscussionbelow)
lifeoriginatedatleastasearlyas3.5Gaago,asindicatedby(mostly)Eubacteria
EvidenceIncludes:
b1.MegascopicStromatolites
Stromatolitesarelaminatedstructuresformedbybluegreenalgae(cyanobacteria)
theearliestStromatolitescomefromtheSwazilandGroupofSouthAfricaandthePilbara
SupergroupofAustralia(datedabout3.0to3.55Ga)
b2.PermineralizedMicrofossils
filamentouskerogenrichmicrofossilssimilartocyanobacteriaoccurinchertsofthePilbara
Supergroup(WarrawoonaGroup)ofAustraliadatedatabout3.4Ga
b3.BiologicallyProducedOrganicMatter
organiccarbon13valuesfromthe3.0to3.55GaoldSwazilandandPilbarasedimentsare
similartothoseofmoderncyanobacteriaandphotosyntheticbacteria
2.DomainEucarya
Eukaryotes=singleormulticelledorganismswithchromosomesmadeofDNA,RNAand
proteinscontainedwithinamembraneboundnucleus
a.Characteristics
withspecializedstructures(vacuoles,mitochondria,manywithchloroplasts)
oxidizesugarsasasourceofenergy
meiosispresent=withtwoconsecutivecelldivisionsbywhichthechromosomesarereduced
fromthediploidnumberofsomaticcellstothehaploidnumber(half)characteristicofgametes
andspores
sexualreproductionprovidesmorevariationthatmaypotentiallyenablethespeciestobetter

PALEOLECTURE,PAGE30
surviveenvironmentalchanges
cellsrangefrom3micronstoseveralmillimeters
theDomainEucaryaincludestheKingdomsProtista(Protoctista),Fungi,Plantae(Metaphyta)
andAnimalia(Metazoa)
b.OriginofEukaryotes
nuclearmembraneprobablyformedbyinvaginationofcellmembrane
specializedstructures(chloroplastsandmitochondria)probablydevelopedfromendosymbiotic
prokaryoteslivingwithinthecellmembraneofarchaebacterialprokaryotes
c.TheOldestEukaryotes
asoxygenbuiltupintheEarlyProterozoicatmosphere,duetothepresenceofphotosynthetic
prokaryotes,theconcentrationofdissolvedoxygenincreasedintheupperocean;asaresultmore
nitrogenwasoxidizedtoformnitrate(NO3),whichisanimportantnutrientforeukaryoticalgae
(Cyanobacteriadon'tneednitrates,astheycanusepurenitrogen(N2)fortheirmetabolism)
oldestknownprobableeukaryoteisthecorkscrewshaped,cylindricalmegascopiccolonialalga
Grypania,froma2.1GaoldbandedironformationinMichigan
organicwalledmicrofossilsofeukaryoticphotoautotrophicplankton("acritarchs")occurin
rocksslightlyyoungerthanthosecontainingGrypania
E.AtmosphericOxygen
increaseddramaticallybetween2.2and1.9Gaago
increaseinatmosphericoxygenwasveryimportantforthedevelopmentofmorecomplexlife
forms
1.EvidenceofIncreasedAtmosphericOxygenLevelsIncludes:
a.Paleosols
analysesofironoxidesinsoilsolderthan2.2Gaindicateatmosphericlevelsofoxygenat
0.2%,or1%PAL(presentatmosphericlevelis21%);atmosphericlevelsinyoungerthan1.9
Gaoldpaleosolsindicate15%PAL,orabout3%totalgaseouscompositionofatmosphere
b.RedBeds
oldestknownthickredbedsatover2.2Ga;indicateoxidationofiron
c.WeatheringResiduesinSedimentaryRocks
nowidespreaddepositsofplaceruraninitearefoundinrocksyoungerthan2.3Ga(uraniniteis
unstableinthepresenceoffreeoxygen)
d.MetalsInBlackMarineShales
weatheringofuraniumandmolybdenumonlandinoxygenatedenvironmentwouldleadto
greaterconcentrationsoftheseelementsinseawater
2.52.7Gaoldmarineshalesarelowinuraniumandmolybdenum;thoseyoungerthan2.1are

PALEOLECTURE,PAGE31
enriched(indicatingpresenceoffreeoxygen)
e.BandedIronFormations
alternatingchertandhematite/magnetitelayers
thereforeoxidized(ferric)ironformedinmarinebasins(althoughthereissomedebateastothe
originaloxygencontentinBIF's)
butBandedIronFormationsdisappearedabout1.9Gaagowhenoxygencontentwassupposed
tobeincreasing(BIF'smayalsobeinfluencedbyoceanstratificationandthereforethismaybe
thesourceofconflictingdata,ortheymaynothavecontainedasmuchoxygenassome
geologistshaveclaimed)
2.TheOzoneShield
developmentofozone(O3)preventedlethalradiationfromreachingtheEarthandwasofmajor
importanceinthedevelopmentoflife
F.OriginandDiversificationoftheMetazoa
1.Metazoa(Animalia)
withspecializedcellsformingtissues(=metazoanorganization)
tissuesareunitedintoorgans(exceptinsimplestinvertebrates)
2.VendianBodyFossilsandTraceFossils
a.TheVendianPeriod(610550Maago)orNeoproterozoic
beginningofVendianPeriodwithmostintensiveglaciationinEarthhistory(the
Varanger/VarangianorMarinoanGlaciation)
laterVendianwithrelativelywarmglobalclimate,withmajormarinetransgressionand
developmentofextensiveshallowmarineenvironments,whichledtoagreaterdiversityof
organisms
b.TracksandBurrows
oldestundisputedmetazoantracesfoundinLateProterozoicrocks(680900Ma)
Vendiantracefossilassemblages(Ichnocenoses)includefeedingburrows,dwellingburrows,
crawlingandgrazingtrails;differfromlaterPhanerozoictypeswithVendiantracefossils
smaller,withshallowpenetrationintothesediment,anduseddifferenttypesofperistalticmotion
(withhydrostaticskeletonsormuscularcontractionoftheventralsideofthebody)
c.EdiacaraFauna
originallyfromPoundQuartziteofSouthAustralia;laterfoundinapproximately25Late
Proterozoiclocalitiesworldwide(onallcontinentsexceptSouthAmericaandAntarctica)
Vendianfossilrecordconsistsofmoderatelylarge,softbodiedinvertebratespreservedinwell
aeratedshallowmarineenvironments(unusualtopreserveinthisenvironmentduringlater
Phanerozoictimes,probablymostlyduetoabsenceofpredators,scavengers,depositfeeders,etc.
duringVendiantimes)

PALEOLECTURE,PAGE32

thestructureandrelationshipsofVendianFossilsisgreatlydebated;hypothesesoftheir
relationshipsconsistofthefollowing:
c1.VendianFossilsmaybeplacedprimarilywithinModernSoftBodiedMetazoans
the"classicaltheory"saysthatEdiacarafossilscanbeplacedinthePhylumCoelenterata
(classesHydrozoa,Anthozoa,Scyphozoa,Conulata,medusaeofuncertainsystematicposition,
andproblematicPetalonamae),PhylumAnnelida(ClassPolychaeta),PhylumArthropoda
(SuperclassTrilobitomorphaorCheliceratarepresentedbytheuncertainclassandsuperclass
Crustacea,ClassBranchiopoda),PhylumPogonophora,PhylumEchiurida,aswellassome
formsofuncertainpositionevenatthelevelofphylum
c2.VendianFossilshaveNOrecentmodernAnalogues
AdolfSeilachersuggeststheEdiacarafauna(theVendobionta)haveauniqueorganization
characterizedbyanextensivebodysurfacedevelopedbyaverycomplicatedreliefandbyalow
bodyvolumeduetotheirflatness("pancakeanimals");fossilshaveafoliatedandquilted
constructiontoincreasesurfacearea
flattenedmorphology,quiltingandhighsurface/volumeratioallowedabsorptionofoxygenand
organicmatterdissolvedinthewaterbydiffusionthroughthebodywall;thereforedidNOT
haveamouth,digestive,orrespiratoryorgans
somefossilsmayhavehadinternalsandskeletonsadaptedtolivingonunstablesandybottoms
(Seilacher's"rockinasock"morphology)
3.TheTommotianFauna
oftenclassifiedasthebaseoftheCambrian
firstfossilsofthe"CambrianExplosion";firstabundantrecordofhardparts,withthousandsof
taxarepresented
lateProterozoicandearliestCambrianwithfossiltubesofvariouscomposition
theTommotianFaunaincludes"smallshellyfossils";withdisjunctscleritesofcalcium
carbonateorcalciumphosphate("tommotiids"arephosphaticscleritesthatevidentlyarticulated
toformanexoskeleton),mollusclikeshells(monoplacophorans),calcareousandsiliceous
spiculesofspongesandprobablysoftcorals,arthropodcarapaces,calcareouscupsofspongelike
archaeocyathans,shellsofbrachiopodsandbrachiopodlikeanimals,aswellasvarioustoothlike
objects
hyolithids(CambrianPermian)sometimesclassifiedasmolluscsbutoftenwithoperculaand
"armlike"structures
G.TheCausesofMetazoanDiversification(the"CambrianExplosion")
1.EnvironmentalFactors
a.endoflatePrecambrian(Varanginian)glaciation
b.developmentofextensivecontinentalshelfareasandepicontinentalseas

PALEOLECTURE,PAGE33

c.Oxygenincreasesto610%ofpresentatmosphericlevels;developmentoftheozonelayer
allowsorganismstoleaverestrictedenvironments
2.BiologicalFactors
a.microorganismsincreaseinnumberandthereforewithincreaseinfilterfeeders
b.organismscreatehabitatsforotherorganisms
c.secretionofskeletons
developmentofhardskeletonsoforganic(scleroproteinsandpolysaccharids)orbiomineralized
(silica,carbonate,apatite)materials
forprotection,supportabovethesubstrate,muscleattachment,guidesforfeedingcurrents,
suppliesofcalciumandphosphatenutrients
calciumcarbonateskeletonscouldnotbesecreteduntiloxygenreachedapproximately10%of
modernlevels(about2%totalatmosphericgases)
H.CharacteristicsandtheClassificationoftheModernMetazoans
1.Presence/AbsenceofInternalBodyCavities
a.AcoelomateOrganisms
donothavecoeloms(cavities)forhousingorgans
includeProtozoa,Porifera,Cnidaria,Platyhelminthes(flatworms)
probablyevolvedfromcolonialflagellateprotozoans
b.CoelomateOrganisms
withcoeloms;higherinvertebratesandchordates;molluscswithpoorlydevelopedcoeloms
echinoderms,annelids,arthropods,chordateswithwelldevelopedcoelomiccavities
b1.Pseudocoelomates
spacebetweenthegut(endoderm)andexternalcovering(ectoderm)isnotfilledwith
mesodermalcells;includesthewormlikerotifersandnematodes(nofossilrecord);coelomisto
makethebodyrigidformusclecontraction
b2.Schizocoelomates
coelomformsfirstasasplitinthemesodermaltissue(=schizocoel);characteristicofannelids
andarthropods
b3.Enterocoelomates
coelomformswhenpouchesfromthegutextendintothemesodermalcellsandare"pinched
off"(=enterocoels);characteristicofechinodermsandchordates
schizocoelomatesandenterocoelomatesoriginatedcoelomformetabolicexchanges

PALEOLECTURE,PAGE34

2.Bilaterians(theBilateria)
organismswithabilateralsymmetry
includemostphylaofmetazoans,exceptspongesandcnidarians
3.Metamerism
segmentationoforganisms
formedduetoneedformuscularcontractionsincrawling,burrowing,swimming,etc.
a.Metamericsegmentation
dividesentirebody;ischaracteristicofannelidsandarthropods
b.Oligomericsegmentation
withbodydividedintothreeparts;characteristicofmostothermetazoans
4.DevelopmentoftheEmbryonicMouth
a.Protostomes
invagination("pushingin")ofembryobecomesthemouth;includesmolluscs,annelidsand
arthropods
b.Deuterostomes
invaginationofembryobecomestheanus;includesechinodermsandchordates;echinoderms
probablydevelopedfromtubedwellingworms
chordatesprobablydevelopedfromfreeswimming"larval"echinodermlikeforms;
developmentofnotochordasan"antitelescoping"deviceforswimming
5.Lophophorates
possessalophophore(comblikeorganusedforfilterfeeding);includebryozoansand
brachiopods
VI.AVarietyofProtists
A.KingdomProtista/Protoctista
Unicellorunicellcolonialorganisms;eucaryoticcells(withorganelles,membranebound
nucleus,etc.);Range=PrecambriantoRecent
Acommonclassificationforprotistsisasfollows:
KingdomProtista
SubkingdomProtozoa
PhylumSarcomastigophora
SubphylumSarcodina
SuperclassRhizopoda

PALEOLECTURE,PAGE35
ClassGranuloreticulosa
OrderForaminiferidaforaminiferans("forams")
SuperclassActinopodea
ClassRadiolariaradiolarians("rads")
*SubkingdomAlgae
DivisionPyrrhophyta
ClassDinophyceaedinoflagellates,ebidians(?)
?ClassIncertaeSedis(uncertaintaxonomicposition)=acritarchs
DivisionHaptophyta
ClassCoccolithophyceaecalcareousnannoplankton
DivisionChrysophyta
**ClassBacillariophyceae(Diatomaceae)diatoms
ClassChrysophyceaesilicoflagellates,archaeomonads(?)
ClassXanthiophyceae
DivisionRhodophytaredalgae
DivisionChlorophytagreenalgae
DivisionPhaeophytabrownalgae
*Note:SubkingdomAlgaeisoftenplacedwithintheKingdomPlantae.**Diatomsareoften
placedinaseparateDivision(asinyourlab).
B.DivisionRhodophyta
redalgae
1.?PhylloidAlgaeleaflike;formedsmallcarbonatebanksinCarboniferousseas
2.SuperfamilyCorallinaceae(LatePaleozoic?JurassicRecent)corallinealgae;most
importantrhodophytesinmodernoceans;skeletaltissueformstwolayeredcellularstructure
C.DivisionChlorophyta
greenalgae;ancestraltolandplants
codiaceanalgae(JurassicRecent)withtinyaragonitespicules;importantinformationof
carbonatesedimentsinmoderntropicaloceans
receptaculitids(OrdovicianPermian)withsaclikeform;oftentermed"sunflowerfossils";the
classificationofthisenigmatictaxoniscontroversial,withsomescientistsplacingthemwithin
the"pleosponges"(kintoarchaeocyathans)
D.DivisionPyrrhophyta/Dinoflagellata
dinoflagellates;importantindexfossils
1.Morphology
during"restingstage"shelldifferentiatedintotwolayers[outerlayer(theca)composedof
celluloselikesubstance;innerlayer(testorcyst)madeoftough,resistanteasilyfossilized
organicsubstance]

PALEOLECTURE,PAGE36
2.MorphologictypesofDinoflagellates
Hystricospheres(JurassicRecent)withlongspines(processes;i.e.,choratecysts);usually
foundinopenmarineenvironments
ProximateTestswithspinesshortorabsent;mostoftenfoundinnearshoreenvironments
3.Ecology/Paleoecology
mostmarineplanktonic;somebenthonic;othersfreshwater,symbiotic(zooxanthellaeofcoral
reefs),orparasitic
produce"redtide"duetoproductionof"paralyticshellfishpoison";someluminescent
E.DivisionBacillariophyta
diatoms
1.Morphology
pillboxlikesiliceousskeleton(frustule);epitheca(largervalve)overlapsthehypotheca
2.MorphologicaltypesofDiatoms
centricdiatomswithradialsymmetry
pennatediatomsbilaterallysymmetricalandwitharaphe(longitudinalslitinthevalvealongits
apicalaxis)
3.Ecology/Paleoecology
widevarietyofenvironments;freshwater,marine;planktonicorbenthonic
asexualreproductionwithreductioninsizeoftests(returnstonormalsizeduringsexualstages)
F.DivisionHaptophyta
calcareousnannoplanktonorcoccoliths;EarlyJurassicRecent
withskeletonconsistingofminutecalcareousshields(usuallyfallapartafterdeathofalgae)
coccoliths(individualellipticaltocircularshields)1to15micronsindiameter
asteroliths(discoasters)oftenstarshaped
G.PhylumProtozoa
singlecellsorcolonialaggregatesorcellswithoutdifferentiationoffunction;reproducemostly
asexually;movebymeansofpseudopodiaorcilia
SubphylumSarcodina=nonflagellate,withpseudopodia
1.OrderForaminiferida
a.ForaminiferanBiology
a1.Nutrition
Pseudopodia=flowingprotoplasmicextensions;movesparticlesinoroutofinnerprotoplasm
inconveyerlikemotion(Streaming)

PALEOLECTURE,PAGE37
Somebenthicandplanktonicforamscohabitwithsymbioticalgae(Zooxanthellae);
photosynthesisprovidesfoodforforams
a2.Movement
Benthicforamssessileorvagile(movebypseudopodia)
Planktonicforamsseveralmigratethroughwatercolumnintosurfacezone(probablyby
changesingascontentofprotoplasm)
a3.Reproduction
Heterophasicwith2types(phases)ofreproduction(=alternationofgenerations);onlyin
benthics
Schizogony=asexualphase;withlargerinitialchamber(proloculus)=Macrospheric
Generation)
Gamogony=sexualphase;smallerproloculus(=Microsphericgeneration)
b.ForaminiferanTestMorphology
Test(the"shell")consistsofasecretedoragglutinatedcovering
b1.WallStructurecementgrains,mineralizecarbonatesorcombinationofthesetwo
AgglutinatedWallStructureoldestgeologically,cementparticlesonlayeroftectin(organic
compound)

Microgranularwallstructure=evolvedinPaleozoic;microganularcalcitegivestesta"sugary"
appearance
CalcareousWalls
Calcareoushyaline=calciteoraragonite;withminuteperforationsinthetestwall
Radialhyaline=calciteoraragonitearrangedwith"C"axisnormaltotestsurface
Granularhyaline=crystallitesrandomlyoriented
Calcareousporcelaneous=shiny,smoothappearanceoftestduetoorientationof
submicroscopiccalcitegrains(randomlyarrangedorbricklike)
b2.Chambers=testcavityanditssurroundingwall
maypossessoneormorechambers
Chamberarrangement=uniserial(1rowofchambers);biserial(2rowsofchambersaddedin
eachwhorl);triserial[3(or2and3)chambersaddedineachwhorl]
Degreeofcurvatureofrowsofchambers=rectilinear(straightseries);arcuate(curvedrowof
chambers);planispiral(spiralliesinsingleplane);trochospiral(spiraldoesnotlieinsingleplane
butwithanaxisofcoiling);streptospiral(trochospiralcoilinginseveralplanesofcoiling)
b3.Degreeofinvolution=involute(majorityofpreviouscoilshidden)andevolute(majority
ofpreviouscoilsvisible)

PALEOLECTURE,PAGE38
b4.CoiledTestmorphology=spiralside(sideshowingtracesofcoil);umbilicalside
(oppositesideofspiral,mayhaveumbilicus=axialspacebetweeninnermarginsofchambers
belongingtosamecoil)
b5.Apertures=primaryopeningsofthetesttotheoutsideenvironment
b6.Pores=round,slitlikeorirregularopeningsapproximately56mminsize;foundin
agglutinatedandhyalineforams
b7.Ornamentation=protrusions;thickeningorsharpeningofchamberperipheriestoform
keels;alsoribs,ridges,striae,furrows,spines
c.MajorMorphologicalGroupsofForaminiferans
approximately100families;1200genera;27,000speciesofforams
over35classificationschemeshavebeenused
c1.Basisofclassification(mostimportantaregroupedfirst)
1)wallcompositionandmicrostructure
2)chamberarrangementandseptaladdition
3)aperturecharactersandmodifications
4)chamberform
5)lifehabitsandhabitats
6)protoplasmiccharacteristics
7)ontogeneticchanges
8)reproductiveprocesses
9)GeologicRanges
c2.OutlineofClassification(MajorGroups)
SuborderAllogromiina=singlechambered,tubular,round,orflaskshaped;testpseudochitinous
(tectinous),agglutinatedmaterialinsomegenera(PaleozoicRecent)
SuborderTextulariina=arenaceousoragglutinatedtests(PaleozoicRecent)
SuborderFusulinina=primitiveforams;calcareousmicrogranulartestswhichlacka

PALEOLECTURE,PAGE39
crystallographicorientation(PennsylvanianPermian)
SuborderMiliolina=calcareous,porcelaneoustests(TriassicRecent)
SuborderRotaliina=hyalineperforatecalcareoustest(TriassicRecent)
d.FusulinidForaminifera
d1.Morphology
smalltoverylarge(upto10cm);oftenlooklike"wheatgrains"
studiedbymeansofaxial(mostimportant),sagittalandtangentialthinsections
evolutionarytrendsareincreaseinsizeoftheinitialchamber(proloculus);increaseinoverall
size;increaseincomplexityofthewallstructure;increaseinintensityoftheseptalfluting(i.e,
increaseincorregationsofthesepta,whichdividethetestintochambers)
d2.Biostratigraphy
FusulinidRange=LowerPennsylvanianUpperPermian
InUpperPaleozoicwith10fusulinidbiostratigraphicassemblagezones
d3.Paleoecology
mostweremarine,benthonicandassociatedwithrelativelyshallow,wellaerated,carbonate
depositionalenvironments;oftenassociatedwithcoralsandalgae
e.PaleozoicAgglutinatedForaminiferans
SubordersAllogromiinaandTextulariinawithapproximately67generainthePaleozoic
typicallyfoundinsiltyshalesand/orfinegrainedshalysandstonesandfinegrainedmudrich
limestones
Carboniferouswith14biozonesbasedonagglutinatedandmicrogranularforams
f.GiantForaminiferans
importantthroughouttheCenozoicintropicalseas;typicallyplanispirallycoiled;Ex.=
Nummulites(Camerina)
g.PlanktonicForaminiferans
JurassicRecent
importantsedimentformersandimportantinMesozoicandCenozoicbiostratigraphy
withornamentation(spines,keels)orbulbouschamberstoassistinflotation
2.PhylumActinopoda(Radiolaria)
radiolarians
a.Morphology
shellsofsilicaorstrontiumsulfate

PALEOLECTURE,PAGE40
b.Classification
dividedintofourclassesonthebasisoftheconstructionofthenucleusandtheshapeand
compositionoftheshell(onlyClassesSpumellariinaandNassellariinaimportant;bothwith
shellsofopalinesilica)
spumellarianswithsphericalsymmetry;nasellarianswithtripodshapeorringlike,orshells
elongate,multichamberedandlatticed
c.Ecology/Paleoecology
exclusivelymarineandplanktonic;liveatalloceanicdepthsandwithcosmopolitan(world
wide)distibution
VII.Sponges,TrueandProblematical
A.PhylumArchaeocyatha
sometimesplacedwithinthesponges
1.Biology
a.Calcareousskeleton,usuallyconical
usuallydoublewalledwithspace(intervallum)inbetween(somelackinnerwall);vertical
partitions(septa)andhorizontalpartitions(tabula)dividetheintervallumintoloculi
withcentralcavity;skeletonperforatedbylargeandsmallpores
b.Reproductionbyasexualbuddingandfission;alsosexual?(probablywithplanktoniclarvae)
2.Paleoecology
shallowmarine,sessilebenthonic(asadults),filterfeeders
livedinaggregatesorcommunitiesdominatedbycalcareousalgaeoralgallikeorganismsof
uncertainaffinities;oftenfoundinreeflikecarbonatebuildups
3.Biostratigraphy
Range:LowertoUpperCambrian(inNorthAmericaonlyLowerCambriantypesareknown)
B.PhylumPorifera
sponges
1.Biology
a.SoftParts
Nointernalorgans,nervoustissue,circulatoryordigestivesystem(Nomouthoranus)
Dermalpores=externalapertureswhichbringinfoodandoxygen;linedwithcollared
flagellatecells(choanocytes)thatproducewatercurrentsandtrapfood
Cloaca(spongocoel)=centralcavity

PALEOLECTURE,PAGE41
Osculum=letsoutwaterfromthespongocoel
b.Reproduction
Asexualbybuddingorbranching
Sexualmostimportant;sphericallarva(amphiblastula)attachthemselvestosubstrateandgive
risetoasponge
2.Skeleton
simpletypeswithnoskeleton
a.Spongin=hornyorganicsubstance;notfoundfossilized
b.Spicules
interlockingminutesiliceous(opalinesilica)orcalcareous(calcite)needles,hooksorplates
embeddedinthetissues
withTwosizesofspicules=Megascleres(largerspiculesupto0.3mmlong;formsmainmass
ofskeleton);Microscleres(smallerspicules0.010.1mm;serveasreinforcement,especially
aroundpores)
KindsofspiculesincludeMonaxonspicules(withsingleaxis;calcareousorsiliceous);Triaxon
spicules(siliceous;3axes);Tetraxonspicules(4axesnotinthesameplane);Polyaxonspicules
(withseveralequalraysdivergingfromapoint;siliceous)andDesmas(siliceousspicule;withno
orderedarrangement)
3.Typesofbodystructure
a.Ascon(asconoid)type
simplest;vaselike;waterentersbodybymeansofdermalporesandwaterexitsthroughlarge
roundedvent(osculum)attopofsponge
b.Sycon(syconoid)type
withinfoldedwall(forstrengthandincreasedigestivearea)

c.Leucon(Leuconoid)type
chambersbecomesubdividedandconcentrated;typicallylackawidecentralchamber
d.Rhagon(rhagonoid)type
mostadvancedsponge;derivedleucontypeinwhichflagellatedcellsareconfinedtospherical
chambersburieddeeplyinbodywall
4.SpongeClassification
Thefollowingisatypicalspongeclassification:
PhylumPorifera

PALEOLECTURE,PAGE42
SubphylumSymplasma
ClassHexactinellida
SubphylumCellularia
ClassCalcarea
OrderPharetronida
?OrderHeteractinida
Otherminororders
ClassSclerospongea*
?OrderStromatoporoidea*
?OrderChaetetida*
?ClassArchaeocyathida*(seeabove)
ClassDemospongia
OrderLithistida
OrderSphinctozoa
OrderKeratosa
Otherminororders
*Note:TheSclerospongea(includingStromatoporoidea),ChaetetidaandArchaeocyathidaare
oftenreferredtotheDemospongia.Archaeocyathans,asnotedabove,areoftenclassifiedasa
separatephylum(asfollowedherein).Sclerosponges,chaetetidsandstromatoporoidsareoften
classifiedseparately(asinyourbook).ReceptacultidsandArchaeocyathidsareoftenclassified
togetheras"pleosponges",althoughIhavetentativelyclassifiedreceptaculitidsasalgae!
5.ClassDemospongea
includesapproximately95%modernmarinesponges
a.Morphologymaylackspiculesorhaveskeletonsofsponginorsilicaorspongin+silica;
(oftensiliceousspiculeswithraysmeetingat60or120angles);bodystructure:rhagon
b.FossilRepresentativesLithistidsmostimportantPaleozoicdemosponges;skeletonof
siliceousdesmas
c.Ecology/Paleoecologyincludesallofknownfreshwaterforms;approximately95%of
modernmarinesponges;marineenvironmentsrangefromwarm,shallowsubtidal,highenergy
toquiet,coldoceanicdeeps
d.BiostratigraphyMiddleCambriantoRecent;recordistypicallydiscontinuous,localized
andrepresentsapoorsamplingofspongediversity(allspongesoflittleusebiostratigraphically)
6.ClassCalcarea(Calcispongea)
a.Morphologydiscreteorunitedcalcareousspicules;spiculetypes=monaxon,triaxons
(mostcommon),tetraxons;bodystructure=ascon,sycon,rhagon

PALEOLECTURE,PAGE43
b.FossilRepresentatives
"Pharetronids"(probablyapolyphyleticgroup)areCalcareawithrigidskeletons(Lower
PermianRecent)andrepresentmostabundantgroup
OrderHeteractinidawithpolyaxonsofoctactine(mostabundant),sexiradiateandpolyactine
morphology(seelabmanual);spiculesaresolidandformdiscreteunits;MiddleCambrianto
Permian;Ex.=Astraeospongium
c.Ecology/Paleoecologymostlyshallowmarine
d.BiostratigraphyRange=Precambrian?;CambrianRecent(alsoseeaboveforindividual
groups)
7.ClassHexactinellida
include"glasssponges"
a.Morphologytubularbodies;sixrayeddiscreteorunifiedsiliceousspicules;raysdeveloped
alongthreemutuallyperpendicularaxes;microscleresalwayspresentandeitherhexasters(small
hexactinalspicules,oftenflowerlikeends)orbirotulates(smallmonaxonswithumbrellalike
ends);axialcanalofmegascleresissquareincrosssection;bodystructure:Rhagon
b.Ecology/PaleoecologyPaleozoictypesmostlylivedonorganicrich,softmudbottoms,
probablyinquietwater;Mesozoicforms(especiallyCretaceous)ofteninchalkfacies;Cenozoic
formslittleknown;moderntypesmostlyinupperbathyalzone(2002000mwaterdepth)
c.BiostratigraphyRange:LowerCambrianRecent
8.ClassSclerospongiae
a.Morphologywithathreelayeredskeleton(baseofcrystallineandaspicularcalciteor
aragonite;middleoflivingtissues;upperwithsiliceousspiculesandcollagenousfibers);
probablyrelatedtodemosponges(andoftenclassifiedwiththem)
b.Fossilsmayincludethestromatoporoidsandchaetetids(seebelow)
c.Ecologymoderntypesofteninshadedcrevices,cavesandtunnelsoncoralreefs
d.BiostratigraphyCambriantoRecent(ifstromatoporoidsareincluded)
e.OrderStromatoporoida
e1.Morphology
calcareousskeleton(coenosteum)withhorizontal(plates,laminae)andvertical(pillar)
structures;outwardlybranchingcanalsmaybepresent;growth=laminar(sheetlike),massive,
cylindrical,dendroid;withsmallporespresentinlaminaeorpillarsandlaminae

PALEOLECTURE,PAGE44

e2.Paleoecology
typicallylivedinclearwarm,shallowwater;occuredonreefs(reefbuilders)orascommensals
ofcoralsinquietwaterafewfeetdeep
Bioticassociation=mostoftenfoundwithtabulatecorals,alsobryozoa,algae,crinoids,
brachiopods
e3.Biostratigraphy
CambrianthroughCretaceous(abundantincarbonateenvironmentsfromMiddleOrdovician
LateDevonian)

f.OrderChaetetida
f1.Morphology=skeletoncomposedofnumerousdistinct,narrowtubes(calicles);wallsof
tubesjoined;caliclessmall(0.10.6mm),dividedbysepta(verticalpartitions)andtabulae
(horizontalpartitions)
f2.Paleoecology=firmsubstrate,mostinshallow,warm,sunlitwaters;oftenassociatedwith
algaeandmayformbiohermsorreefs
f3.Biostratigraphy=OrdoviciantoMiocene(locallyabundantinLowertoMiddle
Pennsylvanian)
VIII.SimpleCoelenterates:theCnidarians
PhylumCnidariawelldevelopedbodytissuesbutsimplyorganized;includescorals,sea
anemonies,jellyfish
A.GeneralCharacteristics
1.Multicellular,mostwithtentacles(foodgraspingprojections)whichsurroundmouth
2.radialsymmetryaboutanaxisbetweenthemouth(oralpole)andbase(aboralpole)
3.Tissuegradeofconstruction;bodywallandtentacleswithectodermandendodermseparated
bymesoglea(connectivetissue)
4.Withcentralcavity(coelenteron/enteron)joiningmouth(intakeandejectionoffood);central
cavityoftendividedbyradialfoldsandpartitions
a.Systemspresent=digestive,muscular,nervous,reproductive,elementarysensory

PALEOLECTURE,PAGE45

b.Systemslacking=respiratory,excretory,circulatory
5.Nematocysts(stingingcapsules)presentforuseindefenseandfoodcapture
6.Asexualbuddingproducingsolitarypolypsorpolypoidcolonies;sexualreproduction
producesciliatedlarvae(planula)whichattachtobottomandformapolyp
7.Withtwostages(polymorphism)afreeswimmingmedusoidstageandfixedpolypstage;in
Hydrozoabothstagesmayoccurormayhavemedusoidstageonly
8.Somewithendoskeletons,someexoskeletonswhichmaybecalciteoraragonite(hydrozoans,
anthozoans)orchitinandcalciumphosphatic(conulatids);veryrarelyhornlikefossilsof
hydroidsandaremorelikeanthozoans
9.Chieflymarine
B.ClassHydrozoa
includesHydra(freshwater)andmanymarinetypes(staghorncoralandsiphonophores)
1.Characteristics
solitaryorcolonial;enteronlacksgullet(leadsfrommouthtogastrovascularcavity);mostwith
chitinousskeletons(somecalcareous);mostlymarine(fewfreshwater);oftenpolypand
medusoidgenerationsalternate
2.Range:LateProterozoic?toRecentnotimportantasfossils
C.ClassScyphozoa
1.Biology
solitary;radialsymmetry;Medusastagedominant;Polyp(attachedforms)muchreduced,
highlymodifiedorlost;coelenteroninsometypesdividedbymesenteries(radiallyarranged
wallsoftissue);livingformswithsexesseparate(malesperm,femaleeggs)
2.Classification
a.SubclassScyphomedusae
jellyfish(truemedusae=freefloatingforms);nektonic;hardpartslacking(gelatinousbelland
tentacles);marineonly;Range:UpperProterozoic?Recent

b.?SubclassConulariida(conulatids,conularids)
formerlyreferredtotheworms,molluscs,coelentratesandhemichordates;Skeletonchitinand
calciumphospate;elongatepyramidalform;attachedbyapexofpyramidorfreeswimming;
tentaclespresent

PALEOLECTURE,PAGE46
Paleoecology=Marinesessilebenthonicornektonic;foundinalllithologiesbutmostlyindark
carbonaceousshale
BiostratigraphicRange:MiddleCambrianEarlyTriassic
D.ClassAnthozoa
includesseaanemones,corals,seafans,seapens,seafeathers
1.Biology
solitaryorcolonial;polypsonly(nomedusae);oralendbearstentacles;gulletpresent(leads
frommouthtogastrovascularcavity);coelenteronpartitionedbymesenteries(radiallyarranged
walloftissue;willhavemineralizedseptaofaragonitewithinandbetweenmesenteriesinmost
groups)
2.Classification
classifiedonwhetherthemesenteriesarepairedornot;2subclassesbutonlyimportantoneis
theZoantharia
SubclassZoantharia
includescorals,seaanemones,seafans,seapens,tabulateandrugosecorals
classificationbasedonarrangementanddevelopmentofmesenteriesandpresenceorabsence
ofaskeleton
Characteristics=solitaryorcolonial;calcareousexoskeleton;paired(coupled)mesenteries
a.OrderRugosa(Tetracoralla)
Biology=solitaryorcolonial;calcareousskeleton(corallite)withepitheca(calcareouswall),
septa(radialplatefromwalltoaxisofcorallite)andtypicallywithtabulae(transversepartitions)
anddissepiments(smallcurvedplatesformingavesicle);6primarysepta(protosepta;
correspondto6pairedmesenteries);secondaryseptadevelopin4of6interseptalspaces
[therefore4fold(biradial)symmetryandcalledTetracoralla)
Paleoecology=marinesessilebenthonic;mostinshallowwater;warm,welloxygenatedwater
withnormalsalinity;bestinareasofslowdeposition(limestonebedsratherthanshaleor
sandstone);Solitarytypeswithsmallattachmentandoftenknockedover(twistedskeletons);
oftenfoundonreefsalongwithtabulatecorals,stromatoporoids,brachiopodsandbryozoans
BiostratigraphicRange:OrdovicianPermian;RugosebiozonesinMississippianthrough
PermianinNorthAmerica,BritainandAustraliaandtoalesserextentinOrdovicianSilurian
b.OrderTabulata
Biology=colonial;calcareousskeletonwithepitheca(outerwall)andtabulae(transverse
partitions);septa(radialpartitions)typicallysmallorabsent;whenpresentmostcommon
numberis12
Paleoecology=reefbuildersinwarm,shallow(lessthan50mwaterdepth),clear,well
oxygenated(agitated,gentlycirculating)water;bottomfreeofsilt(ifsiltywouldpreventlarval
attachment)
BiostratigraphicRange:OrdovicianPermian

PALEOLECTURE,PAGE47

c.OrderScleractinia
c1.Morphology
septainmultiplesofsix(hexacorals)
c2.Paleoecology
mostimportantmodernreefbuilders[liveinsymbioticrelationshipwithdinoflagellates
(zooxanthellae)]
shallowwater(maximumwaterdepthfiftymeters);welloxygenated,agitated,wellcirculated
waterinphoticzone;preferwatertemperatureofabout65F;substraterelativelyfreefromsilt
accumulations
IX."MossAnimals",orBryozoans
PhylumBryozoa4,000livingand15,000fossilspecies;liveincoloniesorafewindividuals
upto10cmdiameter;mayappearbushlike,fungiformorencrusting;oftencarpetlike("moss
animals");Sessile,mostwithcalcareousskeleton
A.Biology
1.Softparts
Zooid=(individualanimal)consistsofpolypide(softparts)andzoarium(skeleton)
polypideenclosedinacarbonateskeletonwithopening(orifice,zooecialaperture)
withlophophore[thislinksbryozoanswithtubicolcousworms(PhylumPhoronida)and
Brachiopoda;all3probablyevolvedfromawormlikeancestor]=foodcatchingorganism
consistingoftentacularcrown(with8tomorethan100tentacles)arrangedinacirclearoundthe
mouth;tentacleswithciliawhichproduceawatercurrenttosweepfoodintothemouth;food
consistsofmicroorganisms,bacteriaandorganicdetritus
Mouthleadsintoacompletealimentarycanal
Norespiratoryorcirculatorysystem
Polymorphism=manygroupswithordinaryfeedingindividuals,somemayhavewierdbird
headlikeanimalsforfeedingordefense,otherswithbristlestomovefoodandlarvae,others
withthickenedwallsforattachment
2.ReproductionandLifeCycle
Coloniesusuallyhermaphroditic(withmaleandfemalezooids;rarelycoloniesconsistofall
malesorallfemales)
Larvasettlestosubstrateinafewhoursandmetamorphosesintoaprimaryzooid;Colony
formsbyasexualbuddingofdaughterzooids
3.Skeleton
mostwithskeletonofCaCO3(calciteorcalciteandaragonite)

PALEOLECTURE,PAGE48
studyusuallybythinsections(1longitudinalsectionand1tangentialsection(perpendicularto
tubes)=especiallystudyWallStructure
TypesofWallStructureincludeamalgamatestructure=wallsofadjacenttubescoalesced;
integratestructure=tubewallsofzooeciumdistinctfromotherzooeciumwalls
Acanthopore=slenderdensewalledtube
Zooecium(Autopore)largetubeorchamberoccupiedbyoneofmainzooids
Mesopore=tubeparalleltoautopores;usuallysmallerandmoreangularwithnumerous
diaphragms;probablyoccupiedbyspecializedzooids
Cystiphragm=calcareousplateextendingfromzooecialwallpartwayacrosstube;surface
coned,convexupwardandinward
Diaphragm=calcareousplateextendingtransverselyacrosswidthofzooecialtube;surfaceflat
orgentlycurved
Coenosteum(coen)=vesicularordenseskeletalmaterialbetweenzooecia
B.ClassificationofBryozoans
1.ClassStenolaemata
zooeciacylindricalwithcalcifiedbodywall;newzooeciaproducedinacommonbudby
divisionofsepta;marine;ovicelllarge;OrdovicianRecent(approximately550genera)
a.OrderTubuliporata(Cyclostomata)=calcareoustubularchamberwithlidlesscircular
aperture;EarlyOrdoviciantoRecent
b.OrderCryptostomata=shortcalcareoustube;colonyperipheralwallsthick;Ordovician
Permian
c.OrderCystoporata(Fistuliporids)=tubularzooeciaisolatedbycystosetissue;Early
OrdovicianTriassic
d.OrderFenestrata(fenestrates,fenestellids)=sheetspiercedbyholes;EarlyOrdovician
Triassic
e.OrderTrepostomata=longcurvedcalcareoustubeusuallyintersectedbypartitions
(immatureandmaturepartsofcolonydistinct);OrdovicianTriassic

2.ClassPhylactolaemata
noncalcareousbodywall,freshwater;LateTertiaryRecent(fossilrecordpoor)

PALEOLECTURE,PAGE49

3.ClassGymnolaemata
mostsuccessfulgroupofmodernbryozoans;almostexclusivelymarine(butsomefreshwater)
havingacircularrowoftentaclessurroundingthemouth;mostcomplexformswithelaborately
calcifiedzooids,skeletonsoftenwithbotharagoniteandcalcitelayers;OrdovicianRecent
a.OrderCtenostomata
zooidsenclosedinagelatinouschamber;comblikeprocessescloseaperturewhententaclesare
retracted;fossilstypicallyconsistofchemicalboringsproducedbysoftbodiedcolonieswithin
calcareoussubstrates(andoftenclassifiedastracefossils);OrdovicianRecent
b.OrderCheilostomata
zooidsenclosedinshortsaclikechitinousorcalcareouschamber;hingedchitinousoperculum
(lid)enclosesaperturewhententaclesareretracted;about1000genera,constitutingmostknown
fossilandlivingspeciesofbryozoans;JurassicRecent
C.Ecology/PaleoecologyofBryozoans
1.Sessilebenthonic;mostnormalmarine[afew(12genera)arefreshwater]
2.Lithology=mostincalcareousrocks(limestone,calcareousshalesandshellymarls);rarein
blackshales,dolomitesandquartzoseclasticrocks
3.BioticAssociation
Paleozoic=mostwithsessilebenthonicorganisms(solitarycorals,articulatebrachiopods,
echinoderms)
PostPaleozoic=mostlywithmollusks,sponges,octocorals
4.Inbothancientandrecentsedimentsmostoftenfoundinsedimentsofcontinentalshelves
andaroundcoralreefs(oftencontributetoreefbuildupbytrappingsediments)
5.Factorscontrollingdistribution
generallyhardsubstrates(attachedtoinvertebrateshells,stonesandlargealgae)
waterturbulencecontrolsupperbathymetriclimit;delicate,leafliketypesincalmwater;
branchingformsusuallyinlowerwatersandmassive,encrustingtypesinmoreturbulentwater
ratesofsedimentation=Bryozoanotmuchaffected
salinity=normalseawater(salinity=35o/oo)
D.BiostratigraphyofBryozoans
bryozoansmaybeusefulbiostratigraphically;microorganisms,widespread,rapidevolution
LowerPaleozoicwithmostlystonybryozoans(withrobustbranchingzoariabutoccurin
massiveorglobularcolonies)

PALEOLECTURE,PAGE50
UpperPaleozoicmostlyslenderbranchingcoloniesandlaceliketypes(fenestellids)
duringUpperCretaceoustheCheilostomataexpandedtoapproximately100genera,wherethey
arewellpreservedwithinchalkdeposits
LowerTertiarywithrapidandabundantchangesinmorphology
UpperTertiarywithmanygeographicshiftsindistribution,butlessoriginationandextinction
versusLowerTertiary
X.TheSturdyBrachiopods
PhylumBrachiopodalophophorebearingmarinecoelomates;relatedtobryozoansand
phoronidworms;1700fossilandextantgenera,30,000extinctspecies;approximately260extant
species
A.Biology
1.SoftParts
a.Lophophore
complexciliated,feedingorgansurroundingthemouth;consistsofpairofcoiledarms
(brachia)withciliated,tentaclelikecirri
b.Mantle
twofoldsofbodywallthatlinetheinnersurfaceofthevalves;outerfoldsecretesshelland
innerfoldseparates2cavities
c.Cavities
Body(coelomic)cavityinposterior1/3ofshell;mainbodyoccupiesbodycavity
Mantle(brachial)cavityoccupiesanterior2/3ofshell;largelyoccupiedbylophophore
suspendedbetweenmantles
d.Systems
Digestive=articulatesintestinewithblindterminus(fecalpelletsvoidedthroughmouth);
inarticulateswithanus
Primitivecirculatoryandnervoussystem

Muscular=3setsofmusclesinarticulates[1toopenvalve(diductor),1toclosevalve

PALEOLECTURE,PAGE51
(adductor),pedicle(muscular)attachment];Inarticulateswith12adductors(closevalves),3
pairsofobliquemuscles;musclescarmorphologyisimportantinbrachiopodtaxonomy
2.ReproductionandLifeCycle
Sexesseparate
commonlywithfreeswimminglarvalstagewhoseheadattachestobottomtoformbenthonic
adultform
3.Skeleton
a.Bilaterallysymmetrical(linemustbedrawnacrossvalves;pelecypodwithlinedrawn
betweenvalves);valvesinbrachiopodsunequal
b.Bivalved
Dorsal(brachial)valve=containslophophore
Ventral(pedicle)valve=largervalvewhichhasmuscularpedicleatposteriorend
Valveshinged(articulates)orunhinged(inarticulates)
c.Morphologyofarticulatebrachiopods:
Foramen=pedicleopening
Beak=pointedextremityofvalvewhereshellgrowthbegins
Commissure=junctionbetweenedgesofvalves
Hingeline=edgeofshellwherevalvesarticulate
Cardinalextremity=lateralterminusofhingeline
Beakridge=ridgeextendingfrombeaktocardinalextremity
Delthyrium=openinginpediclevalveadjacenttohingeline;servesforpassageofpedicle
Fold=elevation(uparch)ofavalve(usuallyonbrachialvalve)alongthemidline
Sulcus=depressionofavalvealongthemidline(usuallyonpediclevalve)
Plications(plica)=radialridgesanddepressionsinvolvingentirethicknessofshell
(corrugationsoninnerandoutersurfaces)

PALEOLECTURE,PAGE52
Umbo=relativelyconvexportionofvalvenextto(anteriorto)beak
Interarea=planeorcurvedsurfacebetweenbeakandhingelineoneithervalve

B.Brachiopodclassification
1.ClassInarticulata
shellchitinophosphaticorcalcareous;shellpunctate(perforatedbyfinetubesorporesfrom
interiortoalmosttheoutersurface;forrespirationwhenvalveisclosed)orimpunctate(solid
layers);valvesneverarticulatedbyteethandsockets;musclesandbodywallholdvalves
together;attachmentbypedicleornoattachment;Range:LowerCambrianRecent
MajorordersincludePaterinida,Lingulida,Acrotretida,andObolellida
a.OrderLingulida
CambriantoRecent;shellsprimarilycomposedofcalciumphosphate;shellsbiconvex,and
ovaltosquarishinoutline;lingulidsburrowintosoftsediment,wheretheyareanchoredbytheir
longpedicle
b.OrderAcrotretida
CambriantoRecent;shellsgenerallysubcirculartocircular,unequallybiconvex,andoften
haveapedicleopening;craniaceanshavenopedicleandcementtheirventralvalvetothe
substrate
c.OrderObolellida
Cambrianonly;valvescirculartooval;ventralvalvewithapseudointerareaandwithapedicle
opening
2.ClassArticulata
Shellcalcareous,punctate(perforatedbyfinetubesorporesfrominteriortoalmosttheouter
surface),impunctate(solidlayers)orpseudopunctate(lackporesbutfibrouslayerwithrodlike
calcite;differentialweatheringgivesappearanceofbeingpunctate);valvesarticulatedbyhinge
teethandsockets;LowerCambrianRecent
a.OrderOrthida
probablyancestraltootherarticulatebrachiopods;generallyunequallybiconvexshellswith
radialribs(costae);relativelywidestraighthingelinesandwithinterareasonbothvalves;shell
impunctate,rarelypunctateorpseudopunctate;2suborders;LowerCambrianUpperPermian
b.OrderStrophomenida
planoconvextoconcaveconvex,lesscommonlybiconvex;interareashighlyvariable,hinge
linetypicallylong;pedicleopeningmuchreducedorabsent;typicallypseudopunctate;Lower
OrdovicianTriassic;4suborders(twoimportant)
SuborderStrophomenidina(Strophomenacea)=interareawelldevelopedononeorbothvalves;

PALEOLECTURE,PAGE53
pedicleforamenveryminuteorlacking;LowerOrdovicianTriassic
SuborderProductidina(Productacea)interareasreducedorlacking,spinesdistributedover
shellsurface;DevonianPermian
c.OrderPentamerida
probablyevolvedfromtheOrthida;withstronglybiconvexvalvesthataresmoothorfinely
costate;withrobustpediclespondylium(curvedplateinmidlineofbeakonpediclevalvefor
muscleattachment);opendelthyrium(openinginpediclevalveadjacenttohingeline;not
coveredbyadeltidium);impunctateshell;interareascommonlysmall;hingelineshortor
moderatelylong;2orders;MiddleCambrianUpperDevonian
d.OrderRhynchonellida
probablyevolvedfrompentamerids;shelltypicallybiconvex;interareas,pedicleandhinge
lengthhighlyvariable;punctateorimpunctate;spiralbrachidium;MidOrdovicianRecent;3
Suborders(somesaysuperfamilies)
e.OrderSpiriferida
shelltypicallybiconvex;interareas,pedicleandhingelengthhighlyvariable;punctateor
impunctate;spiralbrachidium"points"towardcardinalextremities;MidOrdovicianJurassic;4
Suborders
SuborderSpiriferidina(Spiriferacea)=impunctateshells,longhingelines;surfacesmarkedby
ribs(costae)andplications(internalandexternalcorrugations);MiddleSilurianEarlyJurassic
f.OrderTerebratulida
mostabundantmodernbrachiopods;typicallybiconvexshellswithshorthingeline;shell
surfacesmoothorfinelycostate;interareaonpediclevalveonly;punctate;functionalpedicle;
complexloopedbrachidium(calcareoussupportforlophophore);Range:LowerDevonian
Recent
C.EcologyofBrachiopods
1.Exclusivelymarine,benthonicepifaunalandgregarious
2.Food=diatomsanddinoflagellates(the#1and#2primaryproducersinmodernseas)
3.Waterdepth=mostlyshallow,continentalshelfforms
4.Distribution=cosmopolitanfromArctictoAntarctic
5.Salinity=typicallynormalmarinebutmodernterebratulidsandlingulidscansurvivelower
salinities

PALEOLECTURE,PAGE54
6.Substratemaypreferhardsubstrates
D.PaleoecologyofBrachiopods
1.Paleobathymetry
waterdepth
Paleozoic=essentiallyallshallowwaterforms;Cambrian/Ordovicianarticulatesand
inarticulatesfoundmostlyinsandyandshellyfaciesindicativeoflittoralzone;Manyassociated
withreeffacies(Ex.=PermianDelawareBasin,Texas);Productidslivedintheshallowbasin,a
fewontheshelf;afewassociatedwithgraptoliteswereprobablypelagic(attachedtofloating
seaweeds)
MesozoicTertiary=EssentiallyalldeeperwaterformsoriginatedduringMesozoicpost
Mesozoic
Essentiallyallinarticulatesatshallowdepth(mostlessthan60feet)
2.BioticAssociation
Paleozoicwithveryabundantbrachiopodsandprobablyveryimportantlinkinfoodchain
somewithpossiblysymbioticrelationshipswithcorals(?)
3.Attachment
Fossilbrachiopodsvariouslyattachedpedicles;somefirstattachedbypediclebutlaterpedicle
atrophiesandleavesshellfreeonsubstrate(Strophomenida);attachedbycementationtohard
substrates;originallycementedbutlaterfreeonsubstrate(commonlywithconcaveconvex
shellssuchasproductids;anchoredorattachedbyspines);sometypeswithbyssuslikethreads
attachingtosubstrate
E.BiostratigraphyofBrachiopods
Cambriandominatedbytrilobitesandinarticulatebrachiopods
threemaximafornumbersofgeneraOrdovician,Devonian(mostdiversity)andPermian
threegroupsdominatePaleozoic=orthids,strophomenids,spiriferids
Ordovicianwithmarinetransgressionandopeningofnewniches
Devonianbrachsinallmarineenvironments;manyimportantindexfossils
MississippianPennsylvanianwithdecline(duetoabundantcoalswampenvironmentsand
coolingseas?)

PALEOLECTURE,PAGE55
TremendousdecreaseinnumbersinLatePermianEarlyMesozoic
FewMesozoicforms(mostlyrhynchonellidsandterebratulids;Terebratuloidsfound
throughoutTertiaryworldwide)
XI.Worms,Burrows,Trails,andOtherProblematica
A.WormsofVariousPhyla
anumberoflargelyunrelatedforms(the"Vermes"ofolderclassifications)havebeen
consideredas"worms";theseincludethepriapulidworms(Nemertea),flatworms
(Platyhelminthes),peanutworms(Siphunculoida),roundworms(Nematodaor
Nemathelminthes),horsehairworms(Nematomorpha),andsegmentedworms(Annelida;the
onlyphylumcommonlypreservedasfossils)
1.PhylumAnnelida
segmentedworms,includingearthworms(Oligochaeta),leeches(Hirudinea)andmarine
bristleworms(Polychaeta)
a.AnnelidBiology
composedofmanyringlike,similarsegments,andwithsegmentationofinternalstructures
includingnerves,muscles,circulatory,excretoryandreproductiveorgans
b.AnnelidPaleontology
membersofafewpolychaeteorderspossesspharyngealjaws(scolecodonts),includinga
ventralpair(mandibles)forchewing,aseriesofasymmetricalpairedmaxillaeformanipulating
food,andabasalpair(carriers)thatsupporttheposteriormaxillae;somepolychaetessecrete
agglutinatedtubesbycementingsandgrains,shellfragmentsordebriswithmucus,orsecrete
calcareousshellsthatarecommonlypreservedasfossils(Exs.=Spirorbis,Serpula);about150
generaofpolychaetesknown;Proterozoic?,CambriantoRecent
B.Ichnology
studyoftracefossils(ichnofossils,lebensspuren;tracks,trails,andburrowsoforganisms)
maybedescribedbymeansofdescriptivegeneticclassification,Ethologicalclassificationor
bymorphology
Ichnogenusmorphologicaltypeofatracefossil(reflectsenvironmentratherthanthecreator)
Ichnofaciescorrelationbetweendepositionalenvironmentsandtracefossilassemblages
1.Tracefossilsusefuldueto:
a.longtimerange
b.narrowfaciesrange

PALEOLECTURE,PAGE56
c.nosecondarytransport(thereforegoodindicatorsoforiginalsedimentologicalconditions)
d.preferclasticsediments
tracefossilsareoftenformedinenvironmentshostiletothepreservationofbodyfossils,such
asbeachsandsanddeepmarineshales
e.notusuallyaffectedbydiagenesis
2.Sedimentologicalusesoftracefossils
a.Rateofdeposition
Slow,continuousdepositiontypicallywithcompletebioturbation
Rapid,continuousdepositionwithnotracefossilsexceptescapestructures
b.Substrateconsistency
Siltyandmuddysubstrateswithdepositfeeders
Clean,wellsorted(highenergy)sedimentswithspecializedsuspensionfeeders
Softsubstratessoftbodiedforms
Firmsubstratescrustaceans(withclawmarks)
Hardsubstratesmechanicalorchemicalborers
c.Paleobathymetrydeterminingancientwaterdepth;themajoruseoftracefossils
IchnofaciesInclude:
c1.ScoyeniaIchnofacies
curvilinearrodswithwrinkledorstriatedsurfaces;horizontaltovertical;insectburrows?;
foundinnonmarineenvironments
c2.TrypanitesIchnofacies
boringsintofullyconsolidatedsubstratesonrockycoasts,inbeachrock,reefsand
hardgrounds.
c3.SkolithosInchofacies
intertidalenvironments(zoneofsuspensionfeeders);vertical"piperocks"insandysediments
c4.GlossifungitesIchnofacies
earshapedspreitedevelopedonintertidalfirmmudbodies
c5.CruzianaIchnofacies
crawlingtraces(Cruziana)orinclinedspreitefilledUshapedburrows(Rhizocorallium)
developedonthecontinentalshelf(zoneofgeneralizedsedimentfeeders)
c6.ZoophycosIchnofacies

PALEOLECTURE,PAGE57
largespreitefilledfeedingloopsfoundbelowwavebaseandaboveturbiditezone(zoneof
churners;alsowithwormlikePhycosiphoninmuds)
c7.NereitesIchnofacies
zoneofsystematicgrazersandfarmerswithdenselymeanderinghorizontalfeedingtraces
(Nereites)orleafliketraces(Oldhamia);zoneofsystematicgrazersandfarmers
3.DescriptiveGeneticClassificationofTraceFossils
a.TracksandTrails
a1.Trackimpressionleftinunderlyingsedimentbyapodiumorfoot
a2.Trackwaysuccessionoftracksindicatingdirectedlocomotion
a3.Traildirectedlocomotionproducesasuperficialgroovemadebycontactoftheanimal's
bodyORcontinuoussubsurfacestructuremadebyamobileendobenthic/infaunalorganism
b.BurrowsandBorings
b1.Boringexcavationmadeinconsolidatedorfirmsubstrates(e.g.,shell,rock,bone,wood,
etc.)
b2.Burrowexcavationinloose,unconsolidatedsediment
b3.BurroworBoringSystemhighlyramifiedorinterconnectedburrowsorborings,typically
involvingshaftsandtunnels
b4.BurrowCastssedimentsinfillingaburrow
c.BioturbationTexture
textureorfabricduetodisturbanceofsedimentsbyorganisms(bioturbation);oftenconsistsof
dense,contorted,truncatedorinterpenetratingburrowsorothertraces,fewofwhichremain
distinctmorphologically
d.Spreite
bladeliketosinuous,Ushaped,orspiraledstructureconsistingofsetsorcosetsofclosely
juxtaposed,repetitiousparallelorconcentricfeedingordwellingburrowsorgrazingtraces
4.EthologicalClassificationofTraceFossils
a.Cubichnia
restingtraces;oftenshallowdepressionsmadebyanimalssettlingordiggingintothesubstrate

PALEOLECTURE,PAGE58
b.Repichnia
crawlingtraces;trackways,superficialtrails,etc.
c.Pascichnia
grazingtraces;grooves,pits,andfurrowsduetomobiledepositfeeders("stripminers")ator
nearthesurfaceofthesubstrate
d.Fodinichnia
feedingstructures;temporaryburrowsduetodepositfeeders("undergroundminers");often
formsingle,branchedorunbranchedshaftsorUshapedburrowsORspreitestructures
e.Domichnia
dwellingstructures;burrowsortubesrepresenting"permanent"domiciles,mostlyof
suspensionfeeders;formsimple,bifurcatedorUshapedstructuresperpendicularorinclinedat
anglestothebeddingORbranchingburrowsystems
f.Fugichnia(EscapeStructures)
substratedegradationoraggradationresultsindisplacementofanimalsupwardordownward
withrespecttotheoriginalsubstratesurface;oftenformsverticallyrepetitivetraces
XII.AnimalsinThreeParts:theTrilobites
A.PhylumArthropoda
1.Biology
a.Segmentedexoskeleton(somites,metameres);pairedjointedlegsonmostsegments;
bilaterallysymmetrical;bodyconsistsoftwoormoredistinctregions,termedTagma(most
differentiatedintohead,thoraxandabdomen)
b.Externalskeleton(Exoskeleton)chitinousandjointed
toincreasesizetheexoskeletonisshedandreplaced(thisprocessistermedmolting,orecdysis)
c.BodySystems:
Nervous=highlyorganized
SensoryOrgans=manywithocelli(compoundeyes);organsfortaste,smellandtouch,
"hearing"(antennaeandantennules=sensorybristles)
Digestive=welldeveloped;mouthonundersideofhead

Respiratory=subaqueouswithgills(onlimbsorconfinedtospecialgillchambers);airbreathers

PALEOLECTURE,PAGE59
withbranchingtubularprocesses(tracheae)
Circulatory=heartondorsalside,witharteriesbutnoveins
d.ReproductionandOntogeny
sexesusuallyseparate,fertilizationusuallyinternal;afterhatchingpronouncedchangesinbody
form(metamorphism)occurinmanytypes
2.Classification
Thereismuchdiversityintheclassificationofarthropods.Someauthorsconsiderthe
ArthropodaasasuperphylumthatissubdividedintothePhylumUnirama(Classes
Onychophora,MyriapodaandInsecta),PhylumChelicerata(ClassesXiphosura,Eurypterida,
ScorpionidaandArachnida),PhylumCrustacea(ClassesBranchiopoda,Cirripedia,Malacostraca
andOstracoda)andPhylumTrilobitomorpha(ClassesTrilobitaandTrilobitoidea).Ihave
chosenaclassificationsystemthatisabitmoreconservativeandtraditional.Thenumbersof
describedfossilgeneraaregiven(inparentheses)andthegeologicrangesofeachtaxongiven.
PhylumArthropoda
SuperclassTrilobitomorpha:Proterozoic(?);CambrianPermian
ClassTrilobita(1401);CambrianPermian
Class(es)Uncertainincluding"Trilobitoidea"(16);Proterozoic(?);
CambrianDevonian
SuperclassCrustacea:Proterozoic(?);CambrianHolocene
ClassBrachiopoda(119);Proterozoic(?);CambrianHolocene
ClassMalacostraca(586);CambrianHolocene
ClassOstracoda(1900);CambrianHolocene
ClassCirripedia(107);SilurianHolocene
ClassEuthycarcinoidea(2);Triassic
ClassCopepoda(2);MioceneHolocene
ClassCephalocarida(1);CamprianHolocene
ClassMystacocarida;Holocene
ClassBrachiura;Holocene
SuperclassChelicerata:CambrianHolocene
ClassMerostomata(89);CambrianHolocene
ClassArachnida(289);SilurianHolocene
SuperclassMyriapoda:SilurianHolocene
ClassArchipolypoda(8);SilurianPennsylvanian
ClassArthropleurida(1);Pennsylvanian
ClassDiplopoda(23);PennsylvanianHolocene
ClassChilopoda(5);CretaceousHolocene
ClassSymphyla(1);OligoceneHolocene
ClassPauropoda;Holocene
SuperclassHexapoda:DevonianHolocene

PALEOLECTURE,PAGE60
ClassCollembola(25);DevonianHolocene
ClassInsecta(approximately5,000);PennsylvanianHolocene
ClassProtura;Holocene
ClassDiplura(2);TertiaryHolocene
SuperclassOnychophora:CambrianHolocene
Class(es)uncertain(2);CambrianHolocene
SuperclassPentastomida:Holocene
ClassLinguatulida;Holocene
SuperclassTardigrada:Holocene
ClassEutardigrada;Holocene
ClassHeterotardigrada;Holocene
SuperclassPycnogonida:DevonianHolocene
ClassPantopoda(1);DevonianHolocene
InthissectionIwillreviewthecharactersofthemajorsuperclasses(orwhatevertheyare)of
paleontologicalimportance.
B.SuperclassTrilobitomorpha
aquaticarthropodswithantennae;noappendagesspecializedasmouthparts;GeologicRange:
CambrianPermian
1.ClassesUncertain
thetaxonomyofCambrianarthropodlikefossilsisveryconfused;inthepasttheyhavebeen
lumpedintothe"Trilobitoidea";somespeciessuchasAnomalocarisarenowplacedwithinthe
Protarthropoda(separateandconsideredtobemoreprimitivethanthetrueArthropoda)
theMiddleCambrianBurgessShalefaunaofBritishColumbiahasfossilswithpreservedsoft
partanatomy;mostformshavethefirstpairofappendagesmodifiedintoantennaewiththeother
appendagesundifferentiated
2.ClassTrilobita
a.SoftParts
a1.Appendagesbiramous(differentiatedintoagillandleg)ORuniramous,(forming
antennae)
a2.Bodysystems
Sensoryapparatusappendagesmodifiedasantennae;lateralpairedeyeseithercompound
(holochroal)withcloselypackedhexagonallensesORaggregate(schizochroal)eye;some
specieswitheyereducedorlost;alsowithventralstifftactilebristles
Muscularanddigestivesystemscomplex
Respirationbyexternalgills
Reproduction=sexesseparate
3larvalstages(planktonic);adultsusuallybenthonicbutsomeplanktonicornektonic(spinous
forms);moltingprobablyoccurred2030timesduringlifeofindividuals

PALEOLECTURE,PAGE61

b.HardParts
b1.Exoskeletonmineralized,chitinous
b2.Bodydividedtransverselyintocephalon(headshieldortagma),thorax(seriesofseparate
articulatesegments);pygidium(tailshieldortagma)
b3.Bodylongitudinallytrilobedwith1axiallobe(probablycontainedinternalorgans)and2
pleurallobes(lateraltoaxiallobeandprobablytoprotectappendagesorforhydrofoils;pleurae
=lateralthoracicsegments)
b4.Facialsutures
linealongwhichexoskeletonofheadsplitwhentrilobitemolted;maybelimitedtomarginof
cephalonorpassasfinelinealongdorsalsurfaceofcheek
Opisthopariansuture=crossescheek,passesalongmedialborderofeyeandintersects
posteriormarginofcephalonmedialtogenalangle(Genalangle=posteriorlateralcornerof
cephalon)
Propariansuture=crossesdorsalsurfaceofcephalon,passesalongmedialedgeofeye,
intersectslateralborderofcephaloninfrontofgenalangle
Gonatopariansuture=bisectsgenalangle
Marginalsuture=sutureentirelyalonglateraledgeofcephalon
OtherFeaturesImportantinClassificationInclude:
Cranidium=centralpartofcephalonincludingaxiallobeandboundedbythefacialsuture
Genalspine=spineextendingposteriorlyfromposteriorlateralcorner(genalangle)of
cephalon
Glabella=elevatedaxialportionofcephalon;representsanteriorpartofaxiallobe
Hypostoma=plateonundersurfaceofcephalondirectlyinfrontofmouth
Occipitalring=narrowposteriorpartofglabellasetofffromrestofglabellumbyanoccipital
furrow
Rostralplate=median,ventralplateincephalonbetweendoublureofcranidiumand
hypostoma

PALEOLECTURE,PAGE62

c.ClassificationoftheClassTrilobita
typicallyinseventoeightorders(someauthorshaveasfewattwo!),13suborders,30
superfamiliesbasedprimarilyon1)cephalicaxialcharacters(glabella,shape,furrows,etc.),2)
patternoffacialsutures,3)morphologyofpygidium(especiallycaudalization)
c1.OrderAgnostida=smallesttrilobites;eyeless(except1family)trilobiteswithsubequal
cephalonandpygidium;possessonly23thoracicsegments;probablyplanktonic;2
suborders;LowerCambrianUpperOrdovician
c2.OrderRedlichiida=firsttrilobites;largeandspinose;large,semicircular,relativelywide
cephalon;typicallylargegenalspines;numerousthoracicsegments;facialsutures
opisthoparianorfused;Glabellarsegmentstypicallydistinct;pygidiumsmallto
rudimentary;eyescommonlyformelongatecrescents;3suborders;LowertoMiddle
Cambrian
c3.OrderCorynexochida=subelliptical,typicallywithlargepygidium,cephalonsemicircular
(commonlywithlargegenalspines);glabelladistinct,expandsforward;eyeselongate
andnarrow;Opisthopariansutures;rostralplatefusedwithhypostomaorrudimentary.
Thoraxwith511segmentswithspinosepleurae;LowertoUpperCambrian
c4.OrderPtychopariida=typicallyopisthoparian,morerarelyproparianorwithmarginal
sutures;threeormorethoracicsegmentspresent;pygidiumsmall(early)tolarge;eyes
moredistinctfromeyeridgesthaninRedlichiida.Glabellatypicallytapersforward
(thoughttobecloselyrelatedtoRedlichiidaandancestraltoallpostCambrian
trilobitesexceptagnostids,butmaybeapolyphyleticgroup);5suborders,26
superfamilies;CambrianOrdovician
c5.OrderPhacopida=typicallyproparianorgonatoparian,rarelyopisthoparian.Glabella
eitherexpandingortaperingforward.Pygidiumtypicallymediumtolarge;3
suborders:LowerOrdoviciantoUpperDevonian
c6.OrderLichida=glabellabroad,extendingtoanteriorborder;glabellarfurrowselongated
longitudinally;occipitalringtendstofusewithglabellarlobe.Opisthoparian.
Pygidiumlarge,includesleaflikeorspinosepleurae;LowerOrdoviciantoUpper
Devonian.
c7.OrderOdontopleurida=Stronglyconvexcephalon.Glabellawidestatoccipitalring;ring
elongateposteriorlyandcommonlybearingtuberclesorspines.Opisthoparian.Large
genalspinestypical;smallspinesonanteriorborderofcephalon;Pleuraeeachbearpair
ofspines,posterioronelong.Pairedspinesonpygidium;LowerOrdoviciantoUpper
Devonian
c8.OrderProetida=glabellalargeandvaulted,welldefined,usuallywithgenalspines,

PALEOLECTURE,PAGE63
narrowandbackwardlytaperingrostralplate,opisthoparian,eyesholochroaland
usuallylarge,longhypostome.Thoraxwitheighttotensegments.Isopygous
pygidium,withpygidiumusuallyfurrowedandnotspiny;OrdovicianPermian.
d.Paleoecology
Marineonly
Larvaeprobablymostlyplanktonic/nektonic,adultsprobablymostlybenthonicbutmanyin
otherenvironments
Adaptationsversusenvironment:
Vagilebenthonictendenciesincludereductionorlossofpygidia
someOdontopleuridawithventralcephalicspinestopropthemselvesonseafloor
Burrowing(fossorial)trilobiteswithpygidiawithlargesurfaces(manyincurved);loseexternal
sculpture;eyesreducedorlost(oreyestall);prominentpygidialspines(alsoserveasmooring
function)
Nektonictrilobiteswithpygidiawithlargesurfaces;bodieslightlyconstructed;largeeyes;
expandedglabella(mayhavecontainedfat);ovatebodyshape
Planktonictrilobiteswithveryspinoseforms;expandedglabella;ovatebodyshape
e.BiostratigraphyimportantindexfossilsintheCambrianwithanumberofbiozones;also
usedintheOrdovician,SilurianandDevonian
D.SuperclassOnychophora("Velvetworms")
wormlikearthropodsthatrepresent"missinglinks"betweenthesegmentedworms(annelids)
andthejointleggedsegmentedarthropods
Fossilonychophoransaremarine;Modernonychophoransareusuallyfoundindensetropical
rainforestsbeneathrocksandbouldersandinhumus
CambrianHolocene
XIII.Crustaceans
PhylumArthropoda,SuperclassCrustacea
A.Biology
Headcomposedofseveralfusedcephalicsegments(somites);2somitespreoralandbear2
pairsantennae;1pairofantennulespresent;3somitespostoralandbear2maxillaand1
mandible;thoraxwith2(ostracodes)tomorethan40(somebranchiopods)somites;sometwo
branched(biramous)appendagespresent;withnaupliuslarva(amicroscopic,freeswimming
larvalstage,typicallywiththreepairsofappendages)
B.Classification
9classes;2unknownasfossils
1.ClassCirripedia
barnacles

PALEOLECTURE,PAGE64
freeswimminglarvalstagesettlesdownonhead,losebivalveshell;attachbyantennules;
secreteoverlappingcalcareousplates;foodbroughttomouthbybiramousandfringedlegs;5
orders;SilurianHolocene
marine,mostlyshallowwater;foodplanktonandorganicdetritus,someparasitic
2.ClassBranchiopoda
"fairyshrimp","brineshrimp","waterfleas"
mostprimitivecrustaceans;largenumberofbodysegments(somewithmorethan40somites);
small,shrimplikeanimals;leaflikeappendages;carapaceunivalvedorcarapaceabsent;Range:
Proterozoic(?);CambrianRecent
freshtobrackishwater;supersalinewater(brineshrimp),rarelynormalmarine;tolerateswide
varietyoftemperatureanddryingandfreezingofpools
3superordersand47orders;only1order(conchostracans)importantasfossils
OrderConchostraca=chitinous+calciumcarbonatecarapaceconsistsof2flapscontinuous
acrossthedorsalregion;usefulinmicrobiostratigraphy(mainlybecausepreservedinhighstress
environmentsglaciallakes,supersalineconditions)
3.ClassCopepoda
theCopepodsarethemostabundantzooplanktonintheWorld'soceans(therearealsoinclude
benthonicandparasiticrepresentatives);theyarecommonintemperateandsubpolarwaters
copepodsareabout0.3to8millimeterslong,withajointedexoskeletonand2bristlyantennae
thatformafilterfeedingdeviceinfrontofthemouth
4.ClassMalacostraca
lobsters,crabs,crayfish,shrimp,alsosandhoppersandpillbugs(sowbugs)
Mostwithbodycomposedofapproximately20segments(6fusedtoformhead,8thorax,6
abdomen;also1telsononendofabdomen);appendagespresentonallbodysegments;anterior
antennulesuniramous,1pairbiramousantennae;1pairheavymandibles,2lighterpairs
maxillae;2superorders;58orders;Range:CambrianRecent
marine,freshwater,terrestrial;mostdecapodsinlittoralzone;estuarine,swampandlagoonal
environmentsoftenfavoredbyfossilforms(Ex.=Pennsylvaniancoalswamps);decapods
mostlycarnivorous,othersfilterfeeders
a.SubclassEumalacostraca
Carapacecoversentirethoraxandfusedtoallthoracicsegments;abdomenwithsixsegments;
includesthedecapods

OrderDecapodacrabs,lobsters,shrimp,prawns,crayfish;Range:TriassicRecent
OrderPhyllocarida"leafshrimp";CambrianRecent
5.ClassOstracoda
usuallysmall(0.54mmlong)butmayexceed20mminlength;bodydividedintoheadand

PALEOLECTURE,PAGE65
thorax;carapacebivalved,withhingedarticulation;insideofshellwithcentralanddorsalmuscle
scarfields(musclescarsusedinclassification)
omnivores,herbivores,somescavengersandafewcommensalorparasitic;freshwater,
brackish,marine,someterrestrial(foresthumus);almostalllatitudesanddepths(butmostabove
photiczone);mostpelagicandbenthicfromshorelinetoseveralthousandmeters
usefulforbiostratigraphy(abundantinmanyenvironments)
6orders(1oftheseuncertain);LowerCambrianRecent;thefollowingtwoarethemost
important
OrderPalaeocopida(Beyrichicopida)=carapacewellcalcified;long,straightcardinalmargin;
manywithlobesandsulci;OrdovicianTriassic,Tertiary(?)Recent(?);includesmost
Paleozoicostracodes(500genera)
OrderPodocopida=carapacewellcalcified;adductormusclescars(closevalves)well
developed;includesallrecentfreshwaterformsandmostmodernmarine)GeologicRange:
OrdovicianRecent;incluesmostMesozoicandCenozoicostracodes(1200genera)
XIV.ArthropodsfromShoalstoAir
A.PhylumArthropoda,SuperclassChelicerata(Cheliceriformes)
bodyconsistsofprosoma(cephalothorax)andopisthosoma(abdomen;usually12segments);
Prosomawith6pairsofappendages(24jointedpincers=chelicerae,chelae;pedipalps
(orginallyforlocomotionbutmodifiedforgrasping,sensing,orchewing);noantennae;aquatic
orterrestrial;scavengersorcarnivores;CambrianRecent
1.ClassMerostomata
bodytypicallydividedintotwoparts(acephalothorax/prosomaandanabdominal
tagma/opisthosoma);withanteriorclaws(chelicerae);opisthosomalappendagesbiramous,with
onebranchservingasagill;with150200thinleafgills;terminalsegmentbearsaspine(telson);
marine,freshwater;CambrianRecent
a.SubclassXiphosura("horseshoecrabs")
typicallywideandshortbodywithlongtelson;opisthosomalsegmentstypicallyfused;6pairs
legs(1pairchelicerae,4pairswalking,lastpair"pushers"forforcinganimalthroughmud)
Classification=3orders
Ecology/Paleoecologyallmodernxiphosuransmarine;migratedfrommarineto
brackish/freshwaterduringDevonian;PermoCarboniferouswithmanyfreshwaterforms
Limulus(horseshoecrab)foundinshallowwater;eatwormsandsoftmolluscs;TriassicRecent
b.SubclassEurypterida
Biology=opisthosomawith12freesegmentsexcludingtelson;6thpairofappendages
typicallyoarlike;sexualdimorphism(malewithclaspers);uptoapproximately3meterslong

PALEOLECTURE,PAGE66
Classification=approximately19genera
Paleoecology=foundinfreshorbrackishwaterbutNOTgenerallyfoundwith"normal
marine"invertebrates;possiblypreferredlagoonalwithhighandlowsalinities;nektonic;
carnivorous
Biostratigraphy:OrdovicianPermian(mostcommonSilurianDevonian)
2.ClassArachnida
includesscorpions,spiders,ticks,mites
a.Biology=bodywithcephalothoraxandabdomen(fusedinticksandmites);cephalothorax
bears6pairsappendages(noantennaeormandibles);firsttwoappendagesmodifiedforfeeding,
lastfourforlocomotion;airbreathing,withrespirationbybooklungs(leaflikeplatescontaining
bloodvessels)ortracheae(ramifiedtubules)
b.Classification=5ordersbasedprimarilyonsegmentation(numberofsegments;fusedor
unfused)andformofchelicerae(scorpionswithpincers,spiderswithout)
c.Ecology/Paleoecologyterrestrial,solitary,carnivorousorparasitic
d.Biostratigraphy:SilurianRecent
Scorpions=UpperSilurianRecent
Spiders=MiddleDevonianRecent(approximately250fossilspecies)
Ticksandmites=DevonianRecent;verysparsefossilrecord
B.PhylumArthropoda,SuperclassMyriapoda
1.Biology
wormlike,withmultiplesegmentsbearinglegs;singlepairantennae;mandible"toothed"for
chewing;skeletonchitinous(maybestrengthenedbycarbonates);airbreathingthroughtracheae
2.Classification6classes;relativelyrareasfossils
a.ClassChilopoda
centipedes
allbodysegmentswithsinglepairoflegsexceptfirst(withapoisonclaw)andlast];
CretaceousHolocene
b.ClassDiplopoda
millipedes
somebodysegmentswith2pairslegs
originallyfreshwater,oneoffirstinvertebrategroupstoventureontoland;Pennsylvanian

PALEOLECTURE,PAGE67
Holocene
3.Ecology/Paleoecology
terrestrial;Chilopodacarnivorous;Diplopodaprimarilyeatdecayingvegetation;fossil
representativesprobablysimilarinhabits
C.PhylumArthropoda,SuperclassHexapoda,ClassInsecta
Hexapodaincludesfourclasses;onlyClassInsectawithextensivefossilrecord
1.Biology
Bodywiththreedistincttagma(head,thorax,abdomen);headwith1pairantennae;compound
eyes;3pairsmouthparts(1pairmandibles,2pairsmaxillae);3thoracicsegmentswithpairof
jointedlegsoneach;wingsoftenpresent;abdomenusuallywith11segmentsorfewer(68);
respirationbytracheae;sexesgenerallyseparate;manywithmetamorphosis
2.Classificationapproximately25to30orders;over1millionmodernspeciesand
approximately13,000fossilspecies
a.Apterygota
small,winglessinsects(includethebristletailsandtheirkin);DevonianRecent
b.Palaeoptera
primitivewingedinsectsthatlacktheabilitytofoldtheirwings
thePalaeopteramaybeaparaphyleticgroup
b1.Protodonata"ancestraldragonflies"(oftendifferentiatedintothePalaeodictypteraand
Megasecoptera);wingspreadsupto0.7to0.75metersandbodies0.3to0.4meterslong;
PennsylvanianPermian
b2.Odonata(dragonflies)=EarlyPermianRecent
c.Neoptera
wingedinsectsthatcanflextheirwingsovertheirabdomens
c1.Orthoptera(grasshoppers,crickets)=PennsylvanianRecent
c2.Hemiptera(bugs,aphids)=EarlyPermianRecent
c3.Coleoptera(beetles)=LatePermianRecent
c4.Diptera(flies,mosquitoes)=TriassicRecent
c5.Hymenoptera(bees,wasps,ants)=TriassicRecent

PALEOLECTURE,PAGE68
c6.Lepidoptera(butterflies)=CretaceousRecent
c7.Blattodea(cockroaches)=PennsylvanianRecent
3.Ecology/Paleoecology
essentiallyallenvironments;primarilyterrestrialbutmanywithaquaticlarvae
4.Biostratigraphy
sometimesusefulinmicrobiostratigraphy;GeologicRange:MiddleDevonianRecent
Note:Themillipedes,centipedesandinsectsareoftenplacedwithina"phylum"or
"subphylum"Uniramia,baseduponthesharedpresenceofuniramousappendagesinthese
groups
XV.SnailsandTheirKin
A.PhylumHyolitha
sometimesgroupedwiththemolluscs
1.Biology
operculate,shellelongateandtapering;apicalportioncommonlywithsepta;mostspecieswith
elongatepairedstructures(props)
OrderHyolithidawithaligula(anteriorshelflikeextensionoftheaperture)andahelens(thin,
scimitarshapedlateralappendagesbetweentheoperculumandconch;forstabilization?)
2.Paleoecology
originallybelievedtobeplanktonic/nektonicbutnowbelievedtobevagilebenthonic(used
propsto"pole"animalacrosssubstrate?)orsessilebenthonic;mayhavebeendepositfeeders
3.Biostratigraphy
mostabundantduringCambrian,thendeclinedrapidly;extinctinLatePermian
B.Criconarids
conelike,probablyplanktonicorganismssometimesplacedwiththemolluscs;Ordovician
Devonian
themajorgroupofcriconaridsaretheTentaculitids,typicallyrepresentedby180mmlong
straightconesornamentedwithtransverseringsorstriae
C.PhylumMollusca
Compriseapproximately11%ofallextantandfossilspeciesofinvertebratesandvertebrates
1.MolluscBiology

PALEOLECTURE,PAGE69

a.SoftParts
a1.Unsegmented(exceptMonoplacopherans)butprobablysharecommonancestrywith
"segmented"arthropodsandannelids
a2.bilateralsymmetry(exceptgastropods)
a3.Bodyregions
Head=withtentaclesandeyes(lostinbivalves)
Foot=ventralandmuscular
VisceralMass(visceralhump)=coiledingastropods;internalorgansconcentratedinthismass
Mantle(Pallium)=dorsalsoftskinorsheetoftissueovergrowingvisceralmass;secretes
calcareousshellwithanorganicmatrix
Mantlecavity=spaceatposteriorendofvisceralmasscontainingpairedgills(Ctenidia)usedfor
breathingand(inbivalves)forfilterfeeding;anus,excretoryandreproductivesystemsopeninto
themantlecavity
a4.DigestiveTract
withmouth,commonlywithjaws;buccalcavitywithradula(hornyribbonforrasping)
a5.Organsandorgansystem
usually3chamberedheart,usually1pairofgillsorwithlung,sexesusuallyseparate;
hermaphroditism(bothsexesinsameanimal)widespread;usuallytrochophorelarva(free,
swimmingciliatedlarva),followedbyveligerlarva(topshapedwithequatorialflange(velum)
bearingcilia);orNOlarvalstages
b.Hardparts
mostwithexternalcalcareousshell(aragoniteandcalcite)
b1.Shellstructure
shellswith5basicstructures;mostcommonareprismaticandcrossedlaminar;bestseenunder
petrographicmicroscope
Prismaticstructure=singleorcompoundcrystalswithlongdimensionsatrightanglestothe
planeoftheshell;oftendullorchalkyluster
Crossedlaminarstructure=laminaeinclinedinoppositedirectionstoincreasestrengthofshells
(usuallymadeofaragonite)

PALEOLECTURE,PAGE70

Foliatedstructure=micalike
Nacreousstructure=thickaragonitelayersseparatedbyorganicsubstance
Homogeneousstructure=nostructureunderplanelight
2.Classification
ClassAplacophora=noshell,wormlike;nofossilrecord
ClassMonoplacophora
ClassPolyplacophora
ClassScaphopoda
ClassGastropoda
ClassRostroconchia
ClassBivalvia(Pelecypoda)
ClassCephalopoda
D.PhylumMolluscaClassMonoplacophora
1.Anatomy
a.SoftParts
1livingspecieswithsegmentation(pairedgills,musclesandotherstructures)
b.Hardparts
cap,spoonshapedorarchedsingleshell;shellaragoniteoraragonite+calcite
2.Classification4orders;bellerophontidsarethemostimportantfossils(CambrianEarly
Triassic)
3.PaleoecologymostEarlytoMidPaleozoicinepicontinentalseas;duringLatePaleozoic?
orEarlyMesozoicmovedtodeeperwater;mostwereprobablydepositfeeders
4.BiostratigraphyLowerCambrianTriassic(fossils);1moderngenus
E.PhylumMolluscaClassPolyplacophora(Amphineura)
chitonsandtheirrelatives
1.Biology
a.SoftParts
withencirclinggirdle;flatfootandventralsoleadaptedforcreeping;380pairsofgills;low
visceralhump;rudimentaryhead;radulausuallypresent

PALEOLECTURE,PAGE71
b.HardParts
aragoniteshellcomposedofeightoverlapping,articulateplates(valves)inseriesalongmid
dorsallineorNoshell
fossilsofchitonstypicallyconsistofloose,disarticulatedvalves
2.Classification2ordersdifferentiatedbyvalvestructure
3.Ecology/Paleoecology
marine,vagilebenthonic(sluggishbottomcrawlers);liveonorunderrocks,typicallywithin
theintertidalzone;herbivores,omnivores
4.Biostratigraphy
LateCambriantoRecent;mostabundantinCenozoic
F.PhylumMolluscaClassScaphopoda
"tuskshells"
1.Anatomy
a.Softparts
elongatebody;headandappendagesprojectfromwideranterioropening(aperture);otherend
ofshell(apex)extendsaboveseafloorsurfaceandproducesexhalent/inhalentcurrents;head
withmanyprehensileprocesses(captacula)protrudingfromlargegillswhichcollectfood
b.Hardparts
3layeredtusklikearagoniteshell;apexmaybesimple,slittedornotched;tubeupright
2.Classification
twofamiliesbasedprimarilyonornamentation
3.Ecology/Paleoecology
marine,semiinfaunalbenthonicdepositfeedingburrowers;usuallysublittoralorbathyal
4.Biostratigraphy
Range:MiddleOrdoviciantoRecent;mostlyMesozoicCenozoic
G.PhylumMolluscaClassGastropoda
1.Biology
a.SoftParts
distincthead,fusedwithsolelikefoot(forcreeping);radulanormallypresent;torsionpresent
(reorientorgans;twistssothatthemantlecavityliesalongsidethehead;balancesanimalover
thefoot)

PALEOLECTURE,PAGE72

b.HardParts
wherepresentshell(conch)issingle(univalve),calcareous(mostlyaragonite;mostlylamellar
andnacreousstructure)
b1.TypesofCoiling
Planispiralcoilinginasingleplane
Pseudoplanispiralshellcoiledinasingleplane,butcannotbedividedintosymmetricalhalves
bythisplane
Conispiralcoilingdeviatesgreatlyfromasingleplane;spirallywound
mostconispiralshellsarecoiledclockwisedownthecone(dextral),butafewarecoiledina
counterclockwisedirection(sinistral)
b2.Morphologyofconchs
Apex=tipofshell;pointofbeginningofshellgrowth
Whorl=singlecompleteloopofaspiralshell
Suture=junctionbetweentwowhorls
Spire=coiledgastropodshellexclusiveofbodywhorl
Bodywhorl=lastformedsinglecompleteloopofaspiralshell
Operculum=platelikestructureattachedtothesnail'sfoot;protectstheanimalbysealingthe
aperture(opening)
2.Classification3subclassesbasedontorsionandgillposition
a.SubclassProsobranchiaalmostallcommonlyobservedsnails;shellcapshapedor
conispiral;withtorsion;primarilymarine,somefreshwaterandterrestrial;Geologic
Range:LowerCambrianRecent
a1.OrderArchaeogastropodafullytorted;rightgillusuallylost;shellsusuallylowspired
andequidimensional;innerlayersofshellsusuallynacreous;8suborders;includes
essentiallyallPennsylvaniangastropods;Range:LowerCambrianRecent
a2.OrderMesogastropodawithsingleleftgill;usuallyconispiralandoftenwithinhalent
notch;shelloftenporcelaneous;Range:MiddleOrdoviciantoHolocene;includemany
commonMesozoicandCenozoictypes

PALEOLECTURE,PAGE73

a3.OrderNeogastropodasingleleftgill;shellsconispiralwithsiphonalnotchorcanal;
carnivores;Range:CretaceoustoRecent(thesearethedominantCenozoicandmodern
snails)
b.SubclassOpisthobranchiawithdetorsionandonegill;shelltendstobereducedorabsent
(thereforepoorfossilrecordexceptforthepteropods=marinenektonic;oftenthin
shell;oftenoperculate;GeologicRange:U.CretaceousRecent)
c.SubclassPulmonatamostshellbearingbutlackoperculum;withdetorsion;Mantlecavity
actsasalung;Mostterrestrialandfreshwater;GeologicRange:PennsylvanianRecent
3.Ecology/Paleoecology
mostshallowmarine,herbivorous,vagilebenthonic
Prosobranchia=mostmarine,somefreshwater,fewterrestrial
Opisthobranchia=onlyimportantfossilispteropods(exclusivelymarine)
Pulmonata=terrestrialorfreshwater
4.Biostratigraphy
Range:LowerCambriantoRecent
UsefulonaspecieslevelandforlocalbiostratigraphyinCarboniferous,Permian,Cretaceous
butnospecificbizones
XVI.Bivalves:Clams,Mussels,andOysters
PhylumMolluscaClassBivalvia(Pelecypoda)
A.Biology
1.SoftParts
Bodybilaterallysymmetrical;compressed
Mantlegrowsdownandcompletelyenvelops"head",footandvisceralmass
Headreducedtorudimentarycondition
Footusuallywedgeshaped(movementbyplowing)
Twogillswithabundantcilia
Inhalentandexhalentsiphonspresent(forinflowandoutflowofwaterandbodywastes)
Complexnervoussystem;closedcirculatorysystemanddigestivesystem
2.HardParts

PALEOLECTURE,PAGE74
Shellbivalved,ofcalciumcarbonate(calcite,aragoniteorboth);shellwith23layers
ShellMorphology:
Dorsal=directiontowardpartofshellcontaininghingeline
Beak=sharppointedprojection;initalpointofshellgrowth
Anterior=partofshellcontainingmouth(beaksofmostpelecypodspointanteriorly)
Growthline=concentriclinesparalleltoshellmargin
Ventral=liesoppositehingeline
Umbo=stronglyconvexportionofvalveadjacenttobeak
Hingeline=edgeofvalvealongdorsalmarginthatisinpermanentcontactwithoppositevalve
Lunule=depressedplaneorcurvedareaalonghingelineinfrontofbeak,equivalenttoanterior
partofcardinalarea
Escutcheon=depressedplaneorcurvedareaalonghingelinebehindbeak,equivalentto
posteriorpartofcardinalarea
Planeofcommissure=surfaceapproximatelycoincidingwithvalvemargins
Hingeplane=edgeofvalvealongdorsalmarginwhichisinpermanentcontactwithopposite
valve
Hingeteeth=projectionsfromhingeplateforarticulationofvalves
Lateralteeth=projectionsfromhingeplatenearlyparalleltohingeline,situatedinfrontor
behindcardinalteeth
Cardinalteeth=projectionsverticalorobliquetohingelinedirectlybeneathorcloselyadjacent
tobeak;theyfitintosocketsofoppositevalve
Socket=depressioninhingeplateforreceptionofahingetoothofoppositevalve
Auricle(ear)=forwardorbackwardprojectionofshellalonghingelineinsomepelecypods
Plica=ribsinvolvingentirethicknessofshell
Costa=ridgeformedbythickeningofshellsurface

PALEOLECTURE,PAGE75

Ligamentarea=portionofsurfacealonghingelinetowhichligamentisattached
Posterior=shelldirectiontowardanusandsiphonalopening(usuallyoppositebeakinclination)
Adductorscar=impressiononinsideofvalvemadebyattachmentofmusclewhichfunctions
forclosureofvalve
Pallialsinus=inwarddeflectionofposteriorpartofpallialline,definingspaceforretractionof
siphons
Pallialline=ventrallineardepressiononinsideofshell
Ligamentgroove=lineardepressionincardinalareaorligamentareamaskingattachmentof
ligamentfibers
Cardinalarea=planeorcurvedsurfacebetweenbeakandhingeline
Valve=partofshelllyingoneithersideofhingeline
B.ClassificationofBivalves
1.Mostimportantfeaturesforclassifyingfossilbivalveshellsare:
a.Dentition
importantatalltaxonomiclevels;mostimportanttypesofdentitionareEdentulous(simplest
hingetype;withligamentbutlacksteeth),Taxodont(manysmallshortteethofvariableshape,
whichmostoftenoccupymostofthelengthofthedorsalmargin),Actinodont(dentition
composedofpseudocardinalandpseudolateralteeth),Cyrtodont(lackscardinalteeth),
Parallelodont(posteriorlateralteethparalleltodorsalmargin),Isodont(twosubequalprominent
hingeteethononevalveandcorrespondingsocketsintheother),Heterodont(teethareseparated
byanedentulousspace;teethareofdistincttypewithcardinalsbeneathbeakandlateralsinfront
orbehindorboth)
b.Formofligamentinsertiononthevalves(especiallyimportantatthefamilyorsuperfamily
level)
c.Stateofadductormusclescars(usuallyattheordertosuperfamilylevel)
d.Stateofpallialline(mainlyatthefamilylevelandbelow)
e.Shellshape(alllevelsfromsuperfamilydownwards)
f.Shellmineralogyandmicrostructure(mainlyatthesuperfamilylevel)

PALEOLECTURE,PAGE76

2.BivalveTaxonomy
superfamilylevelofbivalveclassificationismoststableandisoftenusedtodividethebivalves
intomajorgroups;withmorethan50superfamilies(butwewillonlycoverthefollowing
subclasses)
a.SubclassPalaeotaxodonta
EarlyCambriantoHolocene
stratigraphicallyoldestandanatomicallythemostprimitivepelecypods
allmarineandinfaunal;mostdetritusfeeders;palaeotaxodontnuculoidsburrowintosediment
andgatherfoodparticleswiththefleshyextensionsoftheirlabialpalps
noadultbyssus(threadlikeattachmentstructures);usuallywithequivalvedshellandtaxodont
teeth(withnumerous,approximatelyequal,undifferentiatedhingeteeth)
b.SubclassIsofilibranchia
EarlyCambriantoHolocene
mostshallowmarineorbrackish,somefreshwater;mainlyepibenthic,sedentaryandattachby
awelldevelopedbyssus
withsimpletoothstructureoredentulous(lackhingeteeth);typicallyheteromyarian(posterior
adductorscarlarge)andintegripalliate(palliallineformsasimplearc);anteriorportionofvalve
typicallygreatlyreduced
c.SubclassPteriomorpha
EarlyOrdovicianHolocene
mostshallowmarine,someestuarine,fewfreshwater;allsuspensionfeeders;byssateor
cementedtosubstrateasadults
mostwithreducedanteriorendanddeveloplobes,auriclesand/orwings;manyare
inequivalved;typicallyisomyarian(adductorscarsareequalsize)andlackwelldeveloped
inhalentandexhalentsiphons;dentitionvariable
includearcshells,oysters,penshells,fileshellsandscallops
d.SubclassHeteroconchia(Heterodonta)
EarlyOrdoviciantoHolocene
mostshallowmarinebutalsoincludemostlivingfreshwaterspecies;mostareburrowers
(infaunal)andsuspension(siphon)feeders
mostwithheterodontdentition[hingeteethdifferentiatedintoverticalcardinalssituatedbelow
thebeak(umbo)andwithsubhorizontallateralteethoneitherside]andareusuallyisomyarian
includefreshwatermussels,cockles,piddocks,giantclams,coquinaclams,surfclamsand
rudists(SuperfamilyHippuritacea;JurassicCretaceous;importantCretaceousreefformers)
e.SubclassAnomalodesmata
MiddleOrdovicianHolocene
marine;mostsuspensionfeeders;shellsusuallyedentulous(lackhingeteeth)andisomyarian,
siphonscommonlypresent,exteriorshellornamentationoftenwithgranulesorconicalspines

PALEOLECTURE,PAGE77

C.ClassRostroconchia
earliestCambrianthroughlatestPermian;probableancestorsoftheBivalviaandScaphopoda
usuallywithsmallshells(lessthan2centimeters)butsomeDevoniantypeswithshellsupto15
centimeters
consistofbivalveshellsthatwerejoinedpermanentlyacrossthetop(formeda"tacoshell"like
structure);shellapparentlybrokeperiodicallyalongthemargintoallowgrowth
posteriorportionofshelltypicallydevelopedintoanelongatetube
XVII.FeetBeforeHeads:theNautiloidsandTheirRelatives
A.PhylumMolluscaClassCephalopoda
Probablythemoststudiedgroupoffossilinvertebrates;havebeenusedextensivelyasindex
fossils
1.Biology
a.SoftParts
Body=bilaterallysymmetrical;mostplanispiral
Head=welldeveloped;anteriorinposition;mouthwithcrownofmobileandprehensile
tentaclesandcontainsradulaandupperandlowerjaws;eyeswelldeveloped;hoodlocated
abovehead(tough,fleshyoperculumforclosureofaperturewhenheadisretractedintotheshell)
Foot=highlymodified;withhyponome(muscularfunnelorsiphonforpowerfulexhalent
watercurrentejectionforjetpropulsion)
Gills=singlepairindibranchiates(ex.Octopus);twopairsintetrabranchiates(ex.Nautilus)
BodySystems=heart23chambered;Respiratorysystemcomplex;Nervoussystemhighly
organized
Reproduction=sexesseparate;developmentdirect(fromembryotoadultwithouttrochophore
orveligerlarvalstate);internalfertilization
b.HardParts
Shellcalciumcarbonatearagonite/nacreousORshellreduced/absent
b1.Shellmorphology:
Whorls=shellcoils

PALEOLECTURE,PAGE78
Livingchamber=largeopenreceptacleforsoftparts
Camera=gaschamberinshell
Phragmocone(phrag)=chamberedportionofconchbehindlivingchamber
Adapertural(adoral)=portionofconchnearestaperture
Adapical=portionofconchnearestapex
Venter=partofwhorlfarthestfromaxisofcoiling
Dorsum=portionofconchwhichisuppermost
Septum=curvedcalcareouspartitiondividingshellintochambers
Suture=lineofjunctionofseptumwithwallsofshell
Saddle=forward(adapertural)flexureofsuture
Lobe=flexureofsuturetowardrear(adapically)
Protoconch=initialchamberofshell
Siphuncle=tubeextendingfrombackoflivingchamberthroughseptatoprotoconch
Connectingring=calcareousringformingwallofsiphunclebetweensepta
Umbilicus=depression(orinsomeanopening)inaxisofcoilingformedbydiminishingwidth
ofwhorlstowardsaxis
Annulus=linealongwhichanimalisattachedtowalloflivingchamber
Septalneck=flexureofseptumalongsiphuncleformingashorttubeorfunnel
Siphuncular(siphonal)deposit=calcareousdepositsalongsiphuncle;insomenautiloidsof
considerablethickness
Cameraldeposit=calcareousdepositsonseptaand/orwallsofcamera
b2.TypesofShells:
Cyrtocone=curved,slendershell

PALEOLECTURE,PAGE79
Orthocone=straight,slendershell
Lituiticone=coiledinearlystage,straightatmaturity(namedafterLituites)
Brevicone=short,bluntshell
Longicone=longshell
Ascocone=withslendercurvedearlystageandshort,bluntmaturestageinwhichgas
chambersoverlielivingchamber(namedafterAscoceras)
Gyrocone=looselycoiledshell(likeGyroceras)
Advoluteshell=coiled,whorlstouching
Involute=typeofcoiledshellinwhichpartofouterwhorlextendsintowardcenterofcoiland
coveredpartofadjacentinnerwhorl
Convoluteshell(nautilicone)=outerwhorlsembracinginnerones(likeNautilus)
Conispiral(trochoid)=coiledlikeascrew
Planispiral=typeofcoiledshellinwhichwhorlslieinasingleplane
Heteromorph=ammonoidshelltypeinwhichshellisNOTplanispiraland/orinwhichwallsof
coilarenotincontact
b3.TypesofSutures:
Goniatitesuture=typeofsuturecharacterizedbysimpleflutingconsistingofsingleseriesof
lobesandsaddles(i.e.,withfirstorderlobesandsaddles);flexuresundivided(Devonian
Permian;1Triassicgenus,1Cretaceousgenus)
Ceratitesuture=typeofsuturecharacterizedbypresenceofsmalllobesandsaddlesonmajor
lobes(secondorderlobesandsaddles)(MississippianTriassic;afewCretaceous;arethe
predominantTriassicammonoids)
Ammonitesuture=typeofsuturecharacterizedbycomplexfluting;smallersecondaryand
tertiarylobesandsaddlesdevelopedonlargerprimaryset[PermianCretaceous(especially
JurassicCretaceous)]
2.ClassificationofCephalopods
a.SubclassNautiloidea

PALEOLECTURE,PAGE80
smalltolargeconchs,shellsorthoconic(straight),Cyrtoconic(curved,slender),coiledinvolute
(partofadjacentinnerwhorlscovered);siphunclepositionanddiametervariablewithlongor
shortsegments;septalflexureofseptumatsiphuncleformingshorttubeorfunnelnecksusually
straight,cylindricalandextendingonlyshortdistancetoprecedingseptum(butsometimes
curvedconcaveoutward);GeologicRange:UpperCambrianRecent;8orders,700genera
OrderNautilida=coiledshells;OrdovicianRecent
b.SubclassEndoceratoidea
Mediumtoverylargeconchs[upto10meterslong!];usuallyorthoconic(generally
longiconic);rarelycyrticonic(curved,slender)orbreviconic;siphunclesmediumlarge,usually
locatedventrally;siphunculardepositsconsisttypicallyofconicalsheaths(endocones);Geologic
Range:UpperCambrian?;LowerOrdovicianSilurian;2orders
c.SubclassActinoceratoidea
Mediumtoverylargeconchsupto6meterslong,usuallyorthoconic;cameralandsiphuncular
depositspresent;GeologicRange:LateCambrian?,MiddleOrdoviciantoLateMississippian;1
order
d.SubclassBactritoidea
Orthoconictoweaklycryptoconic(shellshiddeninsidetissue);longiconicorbreviconic;septa
concaveanteriorly;NOsiphuncularorcameraldeposits;siphunclenarrowandincontactwith
ventralwall;suturessimplewithatleastonesmall,shallowVshapedventrallobe;probably
ancestraltoallrecentcephalopods(cuttlefish,squid,octopus)exceptNautilusandpossibly
ancestraltoammonoids;GeologicRange:DevonianUpperTriassic;1order
e.SubclassAmmonoideaseediscussionbelow
f.SubclassColeoideaseediscussionbelow
3.Ecology/PaleoecologyofPrimitiveCephalopods
a.SubclassNautiloidea
a1.Locomotion:
gas(argon/nitrogen)providesbuoyancy;additionofcameraeandcameralandsiphuncular
depositsaddballast;cameraeandseptabraceagainstwaterpressure
coilingresultedforsolvingproblemsofhydrostaticinequilibrium
ancestralbreviconesandcyrtoconesmayhavebeenvagilebenthonic(asindicatedbypreserved
"color"patterns;excessive"ballast")
somegroupsnektonic(eggshapedwithYorTshapedaperture)
coiledshelledformseithercrawlersorswimmers
a2.Distribution:

PALEOLECTURE,PAGE81
mostfossilorthoconesprobablyinclear,shallowwater
faunasrichinnautiloidswithsparseammonoidsandviceversa
Feedingprobablymostwerecarnivores
b.SubclassEndoceratoidea
presenceofsiphunculardeposits(endocones=conicalsheaths)enabledhorizontallocomotion;
mostbenthonic?
c.SubclassActinoceratoidea
Cameralandsiphunculardepositspresent,probablymanybenthonic
d.SubclassBactritoidea
Nosiphuncularorcameraldeposits;probablynektonic
XVIII."Ammon'sStones"andNakedCephalopods
A.PhylumMollusca,ClassCephalopoda,SubclassAmmonoidea
1.AmmonoidBiology
shellusuallycoiled(tightplanispiralwithbulbous,calcareousprotoconch);siphunclesmalland
marginal;NOsiphuncularorcameraldeposits;sutureswithseverallobesand(typicallywith
secondarylobesandsaddles);GeologicRange:LowerDevonianUpperCretaceous;8orders;
1800genera(lessthan10%Paleozoic,over90%Mesozoic)
mostPaleozoicammonoidswere"goniatites"belongingtotheOrderGoniatitida
PermianthroughCretaceousformshadammoniticsutures(bothsaddlesandlobesare
crenulate);mostMesozoicformswereplanispirallycoiledbut"heteromorph"typeswithwide
varietyofshapes;Cretaceousformswithshelldiametersupto3meters
2.Biostratigraphy
Ammonoidsnearlybecameextinctthreetimes(endofDevonian,Permian,andTriassic)before
theirtotalextinctionattheendoftheCretaceous;aftereachnearextinctioneventunderwenta
majoradaptiveradiation
EarlyDevonianammonoidswithprimarilyagoniatiticsutures(withafewsimpleundivided
lobesandsaddles,withnarrowmidventrallobesandbroadlaterallobes);tendencytoward
developmentoftightercoiling
UpperDevonianwith30families,withshortrangesandusefulbiostratigraphically
goniatiticsuturesdominatedMississippian(25families)andPennsylvanian(30families)
duringPermian27familiespresent;decreaseingoniatiticsutures,withextinctionofgoniatitic
lineagesatendofPermian
threefamiliessurviveintotheTriassic,withgreatdiversicationoftheceratites(80Triassic
families,almostallceratites)
JurassicandCretaceouswithammoniticsuturesdominating;with46Jurassicand85
Cretaceousfamilies

PALEOLECTURE,PAGE82

b2.SubclassColeoidea(Dibranchiata)
alllivingcephalopodsexceptNautilus
withtwoarborescentgillsinthemantlecavity,8to10suckerorhookbearingarms(twoof
them,when10arepresent,beingdevelopedintolongtentacles),andinternalshells;somelack
externalshell,otherswithunchamberedexternalshell
OrderTeuthida(Teuthoidea)includesthetruesquids(JurassicRecent);withtenarms;
chitinousshell(thepen)liesabovethevisceralmass;fossilstypicallyconsistofpens,withafew
rareimpressionsofsoftparts
OrderSepioidea(Sepiidae)includethecuttelfish;flatteneddorsoventrally;10armspresent;
usuallybottompredators;Spiruladriftsthroughdeepwaters,hasaninternalchamberedshell
OrderOctopodaoctopuses;witheightarmsandabulbousbody;earliestfossilsareimpressions
ofsoftpartsfromtheUpperCretaceous
OrderBelemnoidea(Belemnitida)include"belemnites"(UpperMississippiantoUpper
Cretaceous);withinternalshellconsistingofachamberedcone(phragmacone),proostracum
andaguard(sheath,rostrum);wellpreservedfossilsshowthepresenceofarmhooksandink
sacsinbelemnoids
XIX.MostlyStemmedEchinoderms
PhylumEchinodermataincludesholothurians(seacucumbers);echinoids(seaurchins);
asteroids(starfishorseastars);crinoids(sealilies)andalsoextincttypes(examples=blastoids,
cystoids)
A.EchinodermBiology
1.Mostwithpentameral(5rayed)symmetry;bilateralsymmetryinsomefossilgroupsandin
echinodermlarvae
2.definiteheadorbrainlacking
3.watervascularsystempresent=networkoftubeswhichopenfreelyintosurroundingsea
waterthroughhydroporesoramadreporite(sieveplate);originalfunctionrespiratorybuttube
feet(podia)usedforrespiration,locomotion,sensing,foodgatheringanddigging(build
burrows)
4.Endoskeletonconsistsofseparateplates,spicules(holothurians)orpiecesofcalcium
carbonate

PALEOLECTURE,PAGE83
5.OrgansystemsNoheart,nodefinitevessels;nervoussystemprimitive;respiratorysystem
absent;sexesseparate
B.Classification
Theclassificationofechinodermsisbasedprimarilyontheskeletalmorphologyoftheadult
stages.Asinmostclassifications,thereissomecontroversyastothepreciserelationshipsofa
numberofgroups.However,thefollowingclassificationistypical.Numberofgeneraisgiven
(inparentheses),alongwiththerangeofeachtaxon.
PhylumEchinodermata
SubphylumCrinozoa*
ClassCrinoidea(1005);MiddleCambrian,EarlyOrdovicianHolocene
ClassParacrinoidea(1315);EarlyOrdovicianEarlySilurian
SubphylumBlastozoa
ClassBlastoidea(95);MiddleOrdovician(?),MiddleSilurianLatePermian
ClassRhombifera(60);EarlyOrdovicianLateDevonian
ClassDiploporiata(42);EarlyOrdovicianEarlyDevonian
ClassEocrinoidea(3032);EarlyCambrianLateSilurian
ClassParablastoidea(3);EarlytoMiddleOrdovician
SubphylumAsterozoa(Eleutherozoa)
ClassAsteroidea(430);EarlyOrdovicianHolocene
ClassOphiuroidea(325);EarlyOrdovicianHolocene
SubphylumHomalozoa
ClassStylophora(32);MiddleCambrianMiddleDevonian
ClassHomoiostela(1213);MiddleCambrianEarlyDevonian
ClassHomostelea(3);MiddleCambrian
ClassCtenocystoidea(2);MiddleCambrian
SubphylumEchinozoa
ClassEchinoidea(765);LateOrdovicianHolocene
ClassHolothuroidea(200);MiddleCambrian(?),MiddleOrdovicianHolocene
ClassEdrioasteroidea(35);EarlyCambrianMiddlePennsylvanian
ClassOphiocistiodea(6);EarlyOrdovicianEarlyMississippian
ClassHelicoplacoidea(3);EarlyCambrian
ClassCyclocystoidea**(8);MiddleOrdovicianMiddleDevonian
ClassEdrioblastoidea(1);MiddleOrdovician
*CrinozoaandBlastozoaaresometimesplacedwithintheSubphylumPelmatozoabothhave
radialsymmetry,acupshapedbody(theca)enclosingtheviscera,withfoodgathering
appendages,andtypicallywithajointedstemattachedtothesubstrate

PALEOLECTURE,PAGE84

**CyclocystoideaissometimesplacedwithintheEdrioasteroidea
C.SubphylumCrinozoa
withglobular,tightlysuturedcalyces;mostwithlongerectarms,manyarmspinnulate.Most
withthecaofseveralcircletsofplatesshowingwelldevelopedpentameralsymmetry.Mouth
nearlycentralonuppersurface,anuslateral;typicallyattachedtosubstratebylongcolumnal
bearingstemsorbycirri.MiddleCambriantoHolocene;approximately1025livingandfossil
generaintwoclasses.
1.MorphologyoftheCrinozoaandBlastozoa
Aboral=directionoppositepositionofmouth
Ambulacrum=narrowtractorgrooveextendingradiallyfrommouth(Typicallytissueoverlying
groovethicklyciliatedandunderlainbyradialcanalofwatervascularsystem;maysubdivide
andextendontoappendages)
Ambulacralplate=calcareousplatesformingonfloorofambulacraltract
Ambulacralpore=poreinorbetweenambulacralplates(forpassageofpodiumorfor
connectionofpodiumtoampulla(muscularsacswhichconnecttocylinderswhichconnectto
podia)
Arms=appendageonoralsurfacewhichbearsextensionofambulacrus(crinoid)
Calyx(plural,Calyces)=partofcrinoidwhichcontainsthesoftparts;hardpartofcrinoid
exclusiveofthefreearmsandpelma
Cirrus(plural,Cirri)=jointedappendageofcrinoidstemsattachedtonodalsortocentrodorsals;
alsorootlikebranchatbaseofsomecrinoidcolumns
Columnals=circular,ellipticalorpolygonal,discoidplatesthatformthestem
Crown=crinoidcalyxandarms
Dorsalcup=portionofcalyxbelowthearm
Interambulacral=areabetweenraysofambulacra
Madreporite=multipleopeningsofwatervascularsystemtotheexterior
Oral=towardmouthoronsamesurfaceasmouth

PALEOLECTURE,PAGE85
Pelma=crinoidcolumnwithallitsappendagesandanchoragestructures
Periproct=areasurroundinganus
Peristome=areasurroundingmouth
Pinnule=slender,unbranchedjointedbranchetsofanarm(crinoid)
Stem(column)=seriesofdisklikeplatesmountedoneontopoftheotherandattachedtoaboral
endoftheca;typicallyterminalendisfastenedtothesubstrate
Tegmen=ventralsurfaceofcalyx(coveredbynoncalcareousintegumentorbyambulacralsand
irregularlyarrangedinterambulacrals)
Theca=skeletonofcalcareousplates
Thecalplate=calcareousplateforminganelementinthetheca(usuallydistinguishedfrom
ambulacralorarmplates)
2.ClassificationoftheCrinozoa
a.ClassCrinoidea
"sealilies";approximately800livingspecies(butmorethan6timesthatnumberoffossil
species);pentamerous(normallyandprimitivelywith5brachialprocesses(multiplearmswell
developed,plated,movable;pinnulate,uniserialorbiserialarms);commonlywithlongstalkand
sessile(secondarilystalklessandfreeliving);cuplikecalyxofcalcareousplatesradially
arrangedin3ormorecirclets;uppersurfacewithmouthandanus;thecalplateswithoutpores;4
subclasses,13orders;MiddleCambrian,EarlyOrdovicianRecent
strictlymarine(salinity2436o/oo);cosmopolitan(tropicstoarctic);gregarious;feedon
phytoplanktonandzooplankton;attachmentbyrootsystemorcement(butmanyJurassicRecent
crinoidsnektonic);mostprefershallowwater;armnumbervarieswithtemperature(tropical
many;cold/deepwaterswithfewspecies)
b.ClassParacrinoidea
maybeanartificialgroupofformsintermediatebetween"cystoids"andcrinoids;calcareous
thecalplateswithirregularpatternandvariableinnumber;thecalplateswithpores;brachioles
uniserial;2orders;GeologicRange:EarlyOrdovicianEarlySilurian(2ordersbasedon
whetherarmsattachedorarmsfreeoftheca)
D.SubphylumBlastozoa
withglobular,tightlysuturedthecae,bearingerectbrachioles;mostwithhydropore,gonopore
andaccessorythecalrespiratorystructures.Mostattachedtosubstratebystemcomposedof
columnalsorholdfastcomposedofmanyplatesinearlyforms.EarlyCambriantoPermian;
approximately233generainfiveclasses

PALEOLECTURE,PAGE86

1.MorphologyoftheBlastozoa
forotherfeaturesseeCrinozoa
Biserial=arrangementofbrachioleplatesininterlockingdoublerows;foundinmany
blastozoans
Brachiole=freeappendageofablastozoan,whichbearsafoodgroove(alongwhichfoodwas
transportedtothemouth)
Diplopore=pairedarrangementofthecalporesinsomeblastozoans
Gonopore=oneormoresmallopeningsinthecaofmanyblastozoans,generallylocatedbetween
themouthandanus,presumablyfordischargeofeggsandsperm
Hydropore=generallyaslitlikeopeningadjacenttogonopore,observedinmanyblastozoans;
interpretedasinletforawatercirculatorysystem
Hydrospire=infoldedthinwalledrespiratorystructureofblastoidsbeneathborderofan
ambulacrumorintersectingradialanddeltoidplatesparalleltoanambulacrum
LancetPlate=elongatespearshapedplateoccupyingcentralareaofanambulacrum;itsouter
surfaceismarkedbyamedianlongitudinalgrooveandmanytransversegrooves,whichare
concealedbycoveringplatesinperfectlypreservedblastoids
2.ClassificationoftheBlastozoa
a.ClassBlastoidea
withconical,budshapedorglobularthecawithfourcircletsofplatesinwelldeveloped
pentameralsymmetry;Fiveshorttolongambulacra,underlainbylancetplates,eachbearinga
longerectbrachiole;eighttotengroupsoffoldlikehydrospiressuspendedincoelomiccavity.
Stemusuallylongbutwithsmalldiameter.MiddleOrdovician(?),MiddleSilurianLate
Permian;about95genera.
filterfedonphytoplanktonandorganicdetritus;sessilebenthonic(fixedtosubstratebyflexible
stalk;moderatewaterenergylevels);planktonic(somenektonic)larvalstage;usuallyfoundin
finegrained(shalesandmicrites)facies;salinityprobablynormalmarine
b.ClassRhombifera(RhombiferaandDiploporitaareoftentermed"cystoids")
mediumsizedbrachiolebearingechinodermswithrespiratoryrhombsdevelopedinadjoining
plates;rhombsarerhombicinshapeandconsistofrowsofslitlikeorporelikeopenings;with
globular,elongate,lensshapedorflattenedthecawithnumerousplatesusuallyarrangedinfour
tofivewelldefinedcirclets.Stemofcolumnalsusedforattachmentorpossiblyforswimming.
EarlyOrdovicianLateDevonian

PALEOLECTURE,PAGE87
c.ClassDiploporita
mediumsizedblastozoanechinodermswithpairedpores(diplopores)thatpenetratesomeor
mostofthethecalplates;small,erect,brachiolelikeappendages;stemmedorstemless;Early
OrdovicianEarlyDevonian
"cystoids"weregregarious;rareingeneralasfossilsbutlocallymaybeveryabundant;filter
feeders(brachioleswithfoodgrooves);mostsessilebenthonic(cementationorbymeansof
prehensilecolumn),fewplanktonic?(globoseformslackingstem)
d.ClassEocrinoidea
radiallysymmetricalthecalplatesarrangedincircularrows;thecabearinglongbiserial
brachioles;stempresentorabsent;suturalporespresentinearlyforms;laterformswithverythin
thecalplates;tenfamilies(basedonplates,pores,brachioles);GeologicRange:Lower
CambrianLateSilurian
Numberofbrachiolespossiblycorrelatedwithtemperature/waterdepth(morebrachiolesin
warm/shallowwater,fewerincold/deeper)
E.SubphylumHomalozoa
includesfoursmallclasses;withbodyflatteneddorsoventrally;asymmetricorbilaterally
symmetricskeleton;fewornoarms;plated"tail"or"stem"(steleoraulacophore);thecaof
calcareousplates
1.Classification
originallyplacedwiththecystoids(asClassCarpoidea);plateornamentationandarrangement
maysuggestthatsomeorders(CornutaandMitrata)ofClassStylophoraareprimitivechordates
ancestraltotunicates,amphioxus,andthevertebrates(butmuchdebatedechinoderm
characteristicsofcarpoidsincludecalciteplateswithanechinodermlikemicrostructureand
mostbearafoodgroove);ancestraltootherechinoderms?(modificationoftentacleswouldlead
towatervascularsystem)
Benthonic?orNektonic?orNektobenthonic?;Tail(stele)usedforattachmentandfeeding?
(prehensile?);feedingbycilia,stele,protusibletentacles?
2.Biostratigraphy
GeologicRange:MiddleCambrianMiddleDevonian
XX.Stars,Urchins,andCucumbersoftheSea
A.SubphylumAsterozoa
"starfish"(seastars)andbrittlestars;withstarshapedbody,openwatervascularsystem;
marineandusuallyavagilebenthicmodeoflife
1.ClassAsteroidea
starfish;hollowarmscontaininglargelobesofbodycavityandenclosedorgans;Early
OrdoviciantoRecent

PALEOLECTURE,PAGE88
carnivorous(abletoextrudestomachthroughmouthforexternaldigestion);abletowithstand
manymarineenvironments
2.ClassOphiuroidea
brittlestars;slender,whiplikearms;EarlyOrdoviciantoRecent
manydetritusfeeders,somecarnivorous
B.PhylumEchinodermata,SubphylumEchinozoa
globose,withoutarmsorbrachioles;radialsymmetry(typically5fold)
1.ClassHelicoplacoidea
spirallypleated,expansible,flexibletest;2spirallyarrangedambulacra;GeologicRange:
LowerCambrian;probablydetritusfeedersandmovedinaccordionfashion
2.ClassEdrioasteroidea
Discoidtheca;ambulacralsystemconfinedtoupperhemisphereoftest;5ambulacraradiate
frommouth(maycurveclockwiseorcounterclockwise);smallirregularplatesininterambulacra;
usuallyattachedonlowersurface;anusandcentralmouthonupper;GeologicRange:Lower
CambrianMiddlePennsylvanian
food=plankton;usuallyfoundinlittoralzonesoncalcareous,arenaceousandmicaceousmud
bottoms
3.ClassOphiocistioidea
Pentaradiate,freelivingformswithdomeshapedbody;ventralmouth(peristome)surrounded
by5verylargearmoredpodia;2sizestubefeet[largerforlocomotionandfeeding(directfood
towardmouth)andsmallerforsensory];carnivorous;GeologicRange:LowerOrdovician
LowerMississippian
4.ClassCyclocystoidea
sometimesplacedwithintheEdrioasteroidea
Small,discoidaltheca;probablyattachedtoflattenedaboralside;numerousplatesarrangedin
rings;branchingambulacra;GeologicRange:MiddleOrdovicianMiddleDevonian;mayhave
beenfreeliving;food=filterfeeder
5.ClassHolothuroidea
seacucumbers;saclikebodywithsmall,looselyattachedplates(sclerites)inleatheryskin;
mouthatoneendsurroundedbyringoftentaclesformedfrompodia;anusatotherend;Geologic
Range:MiddleCambrian(?),MiddleOrdovicianRecent
foundatalldepthsbutbetween40008500mconstitute5090%oftotalbiomass;sessileor
vagile;somefossorial(burrowing);normalsalinitybuttolerateswidevarietyoftemperatureand
resistsdesiccation;predominantlyscavengers;microorganismsandorganicdetritus
6.ClassEchinoidea

PALEOLECTURE,PAGE89
a.EchinoidBiology
globeshapedarmoredanimalswithasubdermalskeletoncomposedofmanyplatesandwith
movablespines(forlocomotionanddefense)
regularechinoids(OrdovicianRecent)withanuslocatedatthe"northpole"ofthesphere
withinacircletofplates;vagrantbenthonic
irregularechinoids(JurassicRecent)withanuslyingoutsideofandposteriortotheringof
plates;mostwerefossorial(burrowing)forms
b.EchinoidBiostratigraphy
Paleozoicechinoidsarerareasfossilsandareoflittlebiostratigraphicuse;allechinoidsexcept
cidaroidsbecameextinctattheendofthePaleozoic
duringJurassicregularechinoidsdevelopedlongerspinesandmoretubefeet,increasingtheir
abilitytorespireandgatherfood;developmentoffossorial(digging)adaptationsduringthe
JurassicandCretaceousresultedinanadaptiveradiationwithmanynewformsevolving
(thereforeechinoidsmaybeutilizedforbiostratigraphicstudieswithinrocksofJurassic,
CretaceousandTertiaryage)
XXI.Nets,Wrigglers,and"Teeth"WithoutJaws
A.PhylumHemichordata
oftenplacedwithinthePhylumChordatabutdonotappeartobecloselyrelatedtotrue
chordates;embryoandearlylarvaofhemichordatesandechinodermssimilar(deuterostomes);
suggestcommonorigin
formerlybelievedtopossessanotochordbutstructureisactuallyanelongatetubularpouch
connectedtothedigestivetract;butmostwithpairedgillslits
includesthemodernpterobranchsandacornworms;onlyimportantfossilrepresentativesare
thegraptolites(aretentativelyconsideredhemichordates)
1.ClassGraptolithina
fossilcolonialmarineorganisms;proteinaceousskeleton;formedcomplexlybranchedcolonies
orsimplelinearseriesofinterconnectingtubes;zooidscommonlyinlinearseriesorinseriesof
clusters,connectedtoeachotherbystolons;CambriantoPennsylvanian;approximately240
generaand1800species
a.MorphologyofGraptolites
Apertureopeningofoneofcolonialcupsortubes
Autothecalargestofthreecupsortubesproducedbyeachactofbudding
BasalDiskchitinouspatchatendofnemaforattachmentofcolony
Bithecasmallcuportubeaccompanyingautotheca

PALEOLECTURE,PAGE90

Nemachitinousthreadliketubewhichterminatesinbasaldiskatoneendandsiculaatotherin
dendroids;ingraptoloidswasthreadlikerodbywhichcolonywassuspended
Rhabdosomeentiregraptolitecolony,developedbybuddingfromasinglesicula
Siculacupbelongingtoinitialzooidofcolony
Stipebranchofcolony,consistingofoverlappingthecae
Stolonchitinoidtubeextendingthroughsuccessivestolothecaeofastipe,andsendingoff
branchstolonstobaseofeachautothecaandbitheca
Stolothecacuportubeofeachsetofthreethecaefromwhichasucceedinggenerationofthree
thecaeisbudded
Thecaanycuportubeofcolonythathousedthezooid
Zooidindividualgraptoliteanimal
b.Classification
6orders(Dendroidea,Camaroidea,Crustoidea,Stolonoidea,Tuboidea,Graptoloidea);
DendroideaandGraptoloideamostimportant
b1.OrderDendroidea
attached,branchinggraptolitescharacterizedbytwosizesofthecae(largerautothecaeand
smallerbithecae);inmostspecies,stolonsincludedhard,black,organicsubstance,probably
proteinaceous;mostlivedinrelativelyshallowwaters;MiddleCambriantoPennsylvanian;
approximately30genera
b2.OrderGraptoloidea
planktonicgraptolitescharacterizedbyonetypetheca(equivalentotautothecaofother
graptolites)onfewstipes;insomecoloniesautothecaeofprogressivelydifferentshapesand
sizesalongstipes;OrdoviciantoEarlyDevonian;about185genera
B.PhylumChordataSubphylumVertebrata(Craniata)Class"Agnatha"Subclass
Conodonta
conodontswerepreviouslyplacedwithintheirownphylum(theConodonta),butarenow
typicallyconsideredtorepresentamemberofthejawlessfishes;preservationofsoftbodied
specimensindicatethattheywereelongate,eellikeanimals
1.RelationshipsofConodonts
conodontchordatesynapomorphiesincludethepresenceofanotochord,dorsalnervecord,
myomeres(muscleblocksalongthelengthofthebody),atail,andamidlinetailfin

PALEOLECTURE,PAGE91
Conodontvertebratesynapomorphiesincludethepresenceofacraniuminfrontofthe
notochord,pairedsenseorgans,extrinsiceyemusculature(absentinhagfishes),andacaudalfin
withradialsupports
basedonthepresenceofdentineandenamelliketissues,eyemusculature,etc.theconodonts
appeartobemorederivedvertebratesthaneitherhagfishesorlampreys
seebelowformorecharacteristicsoftheChordata,Vertebrata,etc.
2.MorphologicalGroupsofConodontelements
skeletalpartsconsistofmicroscopicmineralizedElementsarrangedinpatterns(Apparatuses);
microwearstudiesindicatethatthesehardpartswereusedforsecuringandchoppingupprey;
moreorlesscompleteapparatusesaretermedNaturalAssemblages
a.Coneshaped(Coniform)Elements
coneshapedstructuresconsistingofabaseandacusp
b.Bladeshaped(Ramiform)Elements
structuresthatincludeamaincuspflankedbyposteroanteriorand/orlaterallydirected
processesthatcommonlypossessdenticles
c.Platformshaped(Pectiniform)Elements
structuresthatcommonlybeardenticlesonplatformsoflaterallyexpandedprocesses
3.ClassificationofConodonts(basedonisolatedelements)
a.OrderParaconodontida
typicallyconiformwithadeepbasalcavity,afewwidelyspacedlamellae(layersobserved
withinskeletalcrosssections),considerableorganicmatterwithintheskeleton;approximately
15genera;LateProterozoicMidOrdovician
b.OrderConodontophorida
withconiform,ramiform,andpectiniformelements;numerouscloselyspacedlamellaeand
nonlaminated,oftenbubblelike,lightercoloredmaterial(whitematter)betweenandcutting
acrossthelamellae;basalplatepresent;elementsoftenornamented;approximately215genera;
CambrianUpperTriassic
XXII.FromStarfishtoFish,LordsoftheWater
SubphylumVertebrata
A.Chordates
1.Characteristics

PALEOLECTURE,PAGE92
possessanotochord,adorsalhollownervecordwithashareddevelopmentalpattern,an
endostyleorgan(equivalenttothethyroidglandofvertebrates),andatailforswimming(atailis
adistinctregiondevelopedbehindtheanus)
2.OriginoftheChordates
WalterGarstang(1928)saidthathigherchordatesevolvedfromseasquirtlarvaethrough
paedomorphosis(adultsretainingjuvenilecharacteristics)butrecentmoleculardatasuggeststhat
seasquirtsareaseparatelineofevolution
modernhypothesesstatethatchordatesarederivedfromhemichordates(bothwithciliatedgill
slitsandgiantnervecellsnotseeninechinoderms)ORfromcalcichordates(basedon
interpretationoffossils)ORthathemichordatesandechinodermsaresistergroups(the
Ambulacraria)andurochordates,cephalochordatesandvertebratesareaclade(with
cephalochordatesclosertovertebatesthanurochordates(basedonmorphologicalandmolecular
data)
B.VertebrateCharacteristics
1.Shape
arebilaterallysymmetricalwithlongaxisofbodyusuallyinahorizontalposition;tendencyof
fishandotherswimmingvertebratestowardsafusiformshape
2.Cephalization
tendencytoconcentratesensoryorgansontheanterior("front")end
3.Notochord
long,rodshapedantitelescopingstructurebelownervetubefoundinsomelowervertebrates;
notpreservedinthefossilrecord
4.Cartilageandbonemakesuptheskeletalsystem
a.Cartilage
soft,translucentmaterialconfinedtodeeperlayersofbody;capableofgrowth;somelower
vertebrateswithcartilageonlybutusuallynotpreservedinthefossilrecord
b.Bone
distinguishedfromcartilagebecausebonehasirregular,branchingcellspacesandismuch
strongerthancartilage;notcapableof"expansional"growth
5.AxialSkeleton
vertebraeconstitutingthebackbone
6.AppendicularSkeleton
consistofbothunpairedandpairedappendagesandtheirsupportinggirdles

PALEOLECTURE,PAGE93
a.Limbgirdlessupportthelimbs
b.Fintypes
Medianfinsunpairedfins
Dorsalfinsonuppersideofbody
Analfinonventral(lower)portionofbodyposteriortotheanus
Caudalfinthetailfin;thereareseveraldifferenttypes:Heterocercal(bodycontaining
backbonetipsupposteriorlyandthefinisdevelopedbelowit;probablygaverisetoallother
caudalfintypes);ReversedHeterocercal(Hypocercal)tail(posteriortipofbodytilteddownand
findevelopedaboveit);Diphycercaltail(symmetricaltailinwhichthevertebraeextend
outwardtothetip);Homocercaltail(derivedfishtailtypewiththetailsymmetricalandwith
abbreviatedvertebrae)
Pairedfinsoccurinpairs(ofcourse);includePectoralfins(foundjustposteriortothegill
region);Pelvicfins(foundusuallyatposteriorendoftrunkinfrontofanalopening);thepurpose
offins(exceptcaudal)isforsteeringandbalancing
7.BranchialArchSystem
cartilaginousorbonybarssituatedbetweengillopeningstosupportthegillmusculatureand
stiffenthegillregion
8.Skull
embryoineveryvertebratehasabraincasethatarticulateswiththevertebralcolumn
nasalcapsulesprotectnostrils;hollowsinsidesreceiveeyes;posteriorportionenclosestheotic
(ear)capsules
9.Denticles
derivedfromthedeeperlayersoftheskinandfoundinmanyfish
usuallyhollow,coneshapedstructures;probablymodifiedtoformfinspinesinsharks,etc.
10.Teeth
specializeddermaldenticlespresentin"all"jawedvertebrates
C.TheRecordoftheearliestVertebrates
1.MiddleEarlyCambrian(525520Ma)
ChengjiangFossilSite,YunnanProvince,southwestChinawithmostlyarthropodsbutalso
possiblebasaldeuterostomes(e.g.,theVetulicoliansandYunnanozoons)andthefirstfishes
(Myllokunmingia,HaikouichthysandZongjianichthys)

PALEOLECTURE,PAGE94
2.LateCambrianVertebrates
conodontsweretheearliestvertebrateswithhardtissues
anothergroupofvertebratesisindicatedbyisolatedpiecesofdermalarmor(Anatolepis)from
WyomingandGreenland(thisdermalarmorismadefromapatite,whichischaracteristicof
vertebrates)
D.Agnathans
jawlessvertebrates;pairedfinsabsentorpoorlydeveloped;primitiveearregion
1.Classification
Thereisconsiderabledebate.Ihaveincludedacommonclassificationherewithsome
synonyms:
PhylumChordata
SubphylumVertebrata(Craniata)
Class"Agnatha"(=Cyclostomata)
SubclassMyxinoidea
SubclassPetromyzontida(=Petromyzonida;Petromyzontiformes)
SubclassConodonta
SubclassPteraspidomorphi(=Diplorhinasensustricto)
OrderAstraspida
OrderArandaspida
OrderHeterostraci(=Pteraspida)
SubclassCephalaspidomorpha(=Monorhini,Monorhina)
OrderOsteostraci(=Cephalaspida)
OrderGaleaspida
OrderPituriaspida
Subclassincertaesedis(i.e.,wedon'tknowtherelationshipsofthefollowing2groups)
OrderAnaspida
OrderThelodonti(=Coelolepida;sometimesplacedwithinthepteraspidomorphs)
2."Cyclostomes"
polyphyleticgroupingofthemodernlampreys(Petromyzontida)andhagfishes(Myxinoidea);
semiparasiticanimalswithnojaws;UpperMississippianPresent
3.Conodonts
seediscussionabove
4.Pteraspidomorphs
oftenincludestheAstraspidaandArandaspida(Ordoviciangroupsrepresentedbypiecesof
dermalarmor),theheterostracans,andsometimesincludesthethelodonts
withpairednasalsacsandopenings;typicallyonepairofgillopenings;skullnotforming

PALEOLECTURE,PAGE95
completeheadshield
CommontypesofPteraspidomorphsinclude:
a.Heterostracans
Range:OrdovicianUpperDevonian(earliestundisputedvertebrates)
Characteristics:mouthusuallyanteriorandventralwiththedevelopmentoflong,narrowOral
Plates(havesmalltinybackwardpointingdenticlesthatwereprobablyusedforcapturingprey);
mayhavegivenrisetootheragnathansbutprobablytoospecializedtogiverisetojawedfishes
5.Cephalaspidomorphs
includetheOsteostraci,theGaleaspidaandthePituriaspida
a.Characteristics
withmassiveheadshieldthatcoveredgillsdorsallyandwithventralplacementofmouthand
gillopenings
withsingle,mediallyplacednasohypophysealopening
b.Osteostracans(Tremataspids+Cephalaspids)
themostcommoncephalaspidomorphs
Range:UpperSiluriantoUpperDevonian,mainlyinwesternLaurasia
Characteristics:usuallysmallfishwithundividedbonyshieldwhichextendsdownthebody;
headdorsoventrallycompressed;dorsaleyesandwithdorsalandlateralfields("electric"or
pressuresensitive"sensory"organs?)ontopofhead;heterocercaltail;believedtobebottom
dwellersand"mudgrubbers"
6.Agnathaincertaesedis(Agnathansofuncertainaffinities)
a.Anaspids
Range:UpperSilurianUpperDevonian
Characteristics:small,streamlined(fusiform),laterallycompressedfishes;eyeslaterallyplaced;
mouthterminalandinformofoval,verticalslitforfilterfeeding(?);hypocercaltail
b.Thelodonts(Coelolepids)
Range:LowerSilurian?;UpperSilurianMiddleDevonian
Characteristics:smallfishwithtinyconelikescales(denticles;usedinclassification);eyesfar
apartandlateral;mouthventral(butnearlyterminal);oftenwithhypocercaltail
somespecimensfromtheDevonianwithadeep,laterallycompressedbody,aforkedtailand
largestomach
E.EvolutionofJawsandFins
1.OriginofJaws

PALEOLECTURE,PAGE96
a.Oldertheoriesstatethatjawsmayhavebeenderivedfromgillarchsupports
butproblemisthatgillsinlampreysdevelopmediallytothesupportingskeleton,whereasin
gnathostomestheydeveloplaterallyandjawsdevelopembryologicallyfromtheneuralcrest,not
fromgills
b.Modernembryologicalstudiesindicatethatoncebonesaroundeyesareformed,aseriesof
connectorgenesmayhavebegunmakingalowerjawcartilage,perhapstostrengthenthe
existingmouthparts
2.TypesofJawArticulations
a.Autostylyarticulationformedbyjawswithoutaidfromgillarchbehindthem;foundin
mostplacoderms;someacanthodians
b.Holostylyfusionofupperjawstobraincase;chimaeras,lungfish
c.Amphistylyfirstgillarchbecomesenlargedandaidsinproppingthejawsonbraincase;
foundinsomeplacoderms,someacanthodians,earlysharks,earlyosteichthyes
d.Hyostylyfirstgillarchbearsmainburdenofjawsupport;foundinmodernsharksand
higheractinopterygians
3.Fins
a.Earlyfinswithspines,flapsorfoldsforstability;latertypeswithtwopairedfins(pectorals
andpelvics)articulatingwithgirdles
b.OriginofPairedFins
FinSpineTheoryprimitivefintypewasskeletonwithlongcentralaxisbearingsidebranches;
thereforeprimitivefinhadanarrowbasedevelopedaroundamovablespine
FinfoldTheoryfinsoriginatedaslateralfoldsfrombodywalls;pectoralandpelvicfins
originatedbysubdivisionofthisfold;thereforeprimitivefinhadbroadbase;mostwidely
acceptedtheorybutfinoriginsmaybecombinationof"finspine"and"finfold"
F.Placoderms
1.Classification
PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
ClassPlacodermi
OrderAntiarchi

PALEOLECTURE,PAGE97
OrderArthrodira
2.Characteristics
typicallydorsoventrallycompressedfisheswithheadandtrunkshields(inadvancedtypes
shieldsconnectedbyballandsocketarticulation)
Range:DevoniantoLowerMississippian;placodermshadnodescendants
Paleoecology:freshwaterormarine;usuallybenthonicandnotverypowerfulswimmers
includethelargecarnivorousArthrodiresandthe"arthropodlike"mudgrubbingAntiarchs
G.Acanthodians
1.Classification
PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
ClassAcanthodii
2.Characteristics
smallfusiformfish;allfinsexceptcaudalwithspinesonanterioredge;heterocercaltail
Range:LateOrdoviciantoLowerPermian
H.Chondrichthyans
sharklikefishes
1.Generalcharacteristics
Range:UpperOrdovician/Silurian?(basedonisolatescalesandteeth)toRecent
cartilaginous(cartilageconsistsofprismaticstructure;firstpreservedprismaticcartilageis
EarlyDevonianage);skincoveredwithdermaldenticlesincludingplacoidscales,teeth,claspers
andfinspines(histologyofteethissometimesusedinclassifications)
2.Classification
Here'sacommonclassificationbutthereareothers:
ClassChondrichthyes
PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
SubclassElasmobranchii

Infraclassesunnamedforfollowinggroups:

OrderCladoselachida

OrderSymmoriida

OrderEugeneodontiformes(=Eugeneodontida)includesthecaseodontsand
edestids

PALEOLECTURE,PAGE98

OrderPetalodontiformes(=Petalodontida)

InfraclassEuselachii

OrderXenacanthiformes(Xenacanthida)

OrderCtenacanthiformes

OrderHybodontiformes

CohortNeoselachii
SubclassSubterbranchialia

OrderIniopterygiformes(Iniopterygia)

SuperorderHolocephali

OrderBradyodontidaincludestheSuborderChimaerina
3.Subterbranchialians
Range:MississippiantoRecent
CharacteristicsofSubterbranchialians:shouldergirdlelocateddirectlybehindheadwiththe
branchialbasket(gillarea)crowdedintorestrictedspaceanteriortoshoulderandmostlybeneath
theskull
almostallwereprobablysluggish,vagilebenthonicanimalsincludingthebizarre
IniopterygiansandtheHolocephalians(includesthemodernratfish)
4.Elasmobranchs
withbranchialbasketexpandedposteriorlyandlyingmostlybehindtheskull;gillpouchesopen
separatelytooutsideandgillarcheswidelyspaced
approximately10orders;themostimportantare:
a.Symmoriids
bestknownPaleozoicsharks
Range:UpperDevoniantoPennsylvanian;NorthAmerica,Europe
Characteristics:cladodontdentition,ashortblunt"snout";somewithwierddorsalfinbrushes
(forsexualdisplay?)
b.Eugeneodonts
Range:UpperDevoniantoTriassic
Characteristicssymphysialteethtendtoformelaboratecuttingdevices(edestid,caseodontid
andhelicopriontypesharks)withwhorlshapedreplacement;lateralteethusuallyformcrushing
surfaces;withhighlyspecializedtailfinsinwhichtheneuralandhaemalarchesmodifiedto
formanumberoflargecartilageplates;pelvicgirdlesandfinsarecompletelyreduced
c.Euselachians
includethexenacanthsandthe"modernshark"groups;unitedinresemblancesintheirfin
structure
c1.Xenacanths(pleuracanths)
Range:UpperDevoniantoUpperTriassic;NorthAmerica,SouthAmerica,Europe,Australia,
India

PALEOLECTURE,PAGE99
Characteristics:freshwater(?)sharkswithfusiformbody;teethwithtwoorthreepointedcusps;
twoanalfins;diphycercalcaudalfin
c2.Ctenacanths,HybodontsandNeoselachians
Range:DevoniantoRecent
Characteristics:primarycharacteristwodorsalfinswithspinescoatedwithanenameloidlike
dentine;skincoveredwithplacoidscales
improvementsfromprimitivetomodernsharksincludeshiftfromamphistylictohyostylicjaw
articulation(withshortenedjawsthatpivotonmodifiedfirstgillsupport;formsprotrusion
mechanism);improvementsinsmell,largebrains,calcifiedvertebralcentra;threebasalelements
inpectoralfins;neoselachiansharksincludemodernsharks,skatesandrays
I.Actinopterygians
bonyfish(Osteichthyes)thatdifferfromsarcopterygiansinthepresenceoffinrays(bony,rod
likefinsupports)
1.Range:(UpperSilurian?)LowerDevonianRecent
foundinbothfreshwaterandmarineenvironmentsfromtheDevoniantoRecent
2.ClassificationandOrigin
possiblyoriginatedfromtheacanthodians
taxonomyofactinopterygiansisamesswithnumerouscontroversies
Psarolepis,fromtheUpperSilurian/LowerDevonianofChinaandVietnam,shows
characteristicsofbothactinopterygiansandsarcopterygians,andisclassifiedaseitherabasal
actinopterygianorsarcopterygianoncladograms
a.Acommonclassificationisasfollows(butthereareothers):
PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
ClassOsteichthyes
SubclassActinopterygii

SuperdivisionChondrostei

SuperdivisionNeopterygii
DivisionGinglymodi
DivisionHalecostomi(Ginglymodi+Halecomorphi="Holosteans")
SubdivisionHalecomorphi

SubdivisionTeleosti
3.EvolutionoftheMajorMorphologicalFeaturesoftheActinopterygians
mostlydealwithfeedingandlocomotorysystems

PALEOLECTURE,PAGE100
IthinkitisagreatexampleofDeBeer'sconceptofMosaicEvolution
a.TheLocomotorSystem
actinpropulsion,steeringandhydrostaticadjustment
a1.CaudalFin
primitiveisheterocercalandtypicallywithahypochordal(lower)lobe(foundin
chondrosteans)
homocercaltail(withasymmetriccaudal)foundinsomeholosteansandinteleosts(probably
accompaniedbygreaterefficiencyofswimbladdersandreductionoffish'sweightbyreducing
scales)
a2.DorsalandAnalFins
Chondrosteanswithstiff,triangularandbroadbaseddorsalandanalfins;finraysnumerous
andcloselysetandwerestrongbutinflexible)
primitiveneopterygians("holosteans")withfinraysbecomingfewer,lighter,andthereforefins
weremoreflexibleandmobile
Teleostsdevelopedstiff,pointedspines(especiallyinthedorsalfins)
a3.Pairedfins
Structuralchangesprimitiveactinopterygianswithtriangular,broadbasedpairedfinswith
numerouscloselyspacedfinrays,withmoderatetolargepectoralfinandwithsmallanalfin;
derivedcharacteristicsincludenarrowfinbases,reductioninnumberandwiderspacingoffin
rays;teleostsdevelopedspines
Positionchangesofpairedfinscommonderivedconditionisforwardmovementofpelvicfins
accompaniedbytendencyforpectoralfintoliehigheronthelateralbodywall(goodforintricate
manuevering)
a4.VertebralColumnandRibs
Chondrosteanswithlarge,unrestrictednotochord;centragenerallyabsent;neuralandhaemal
archeswelldeveloped
Teleostswithvertebraeconsistingofbiconcavecentrumandwithneuralandhaemalarches
fullyossified
a5.Scales
generalscalemorphologyandhistologymaybeveryusefulintheclassificationofthe
actinopterygians
Ganoidscaleslarge,thickrhomboidalscaleswith"pegandsocket"articulationcharacteristicof
primitiveactinopterygians;characterizedbypresenceofathinenameloid(=ganoin)layeron
outersurface
Cosmoidscalesderived(?)scalewhichconsistsofthreeparallellayers;possiblyderivedfrom
ganoidscalesbyreductionofganoin;foundinprimitivecrossopterygiansandlungfish

PALEOLECTURE,PAGE101

Cycloidscaleslostganoinandcosminelayers,leavingonlytheinnermostbonylayer
(isopedin);thincircularscaleswithgrowthringsfoundinmanyadvancedactinopterygians
Ctenoidscalesverymuchlikecycloidscalesinstructurebutwithctenii(barbs)onthescales;
foundinmanyderivedactinopterygians
a6.BodyForm
primitiveactinopterygianstypicallyfusiform(chondrosteansandholosteanstypically
conservativebutsometypeshadelongateordeepbodies)
teleostswithawidevarietyofbodyshapes
b.FeedingSystems
manyskullandjawmodificationsduetofeedingandrespiratorymodifications
trendtowardkineticheadskeletons(composedofseveralmobileunitsthatcanmoveagainst
eachother)
b1.Chondrosteanstage
skullbonessupportingjaw(suspensorium)stronglyoblique;primaryupperjawbone(maxilla)
isfirmlyjoinedtootherskullbones;branchiostegalrays(elongatebonesonposteroventralpart
ofskull)numerous,closelyspaced,smallandallsimilar;cheekregioncompletelycovered(allof
thesefeatureswereforbracingthejaws)
b2.PrimitiveNeopterygian("Subholostean/Holostean")Stage
suspensoriumofjawbecomesmorenearlyvertical;specializedjawsupport(thesymplectic)
forms;maxillabecomesseparatedfromotherskullbonesbutstillretainsteeth;numberof
branchiostegalraysdecreases(dorsalmostbranchiostegalenlargedtoanchorthemusclewhich
depressedthelowerjaw);allofthesefeaturesgreatlyincreasedfeedingeffectivenessby
increasingthestrengthofthebite
b3.TeleostStage
upperanteriorjawelement(premaxilla)freedandbecomesprimarytoothbearingelementin
theupperjaw(becameprotrusibleinmanyteleosts);maxillafreedsothatitcouldbepulled
downandforwardbythelowerjawasthemouthopened;thesefeaturesweredevelopedfor
"suctionfeeding"
J.SarcopterygianFishes
bonyfish(Osteichthyes)withfleshylobefins,asquamosalboneispresentontheskull,anda
cosminelayerispresentonthescalesandbones
1.Classification
relationshipsoflungfishes,coelacanthsandtheirPaleozoicrelativesiscontroversial,with
severalcompetingcladisticanalyses
Thefollowingisoneclassificationschemewithseveralothersavailable:

PALEOLECTURE,PAGE102

PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
ClassOsteichthyes
SubclassSarcopterygii
OrderDipnoi
InfraclassCrossopterygii
OrderActinistia
InfraclassTetrapodomorpha
SuperorderOsteolepidida
2.Dipnoans
thelungfish
a.Range:LowerDevoniantoRecent
b.Characteristics:
alldipnoanswithholostylicjaw(fusedtothebraincase);theyhavenomarginalteeth;the
externalnasalopeningsareventralinposition;Nochoanae(internalnares)arepresent;allhave
lobatefins
mostwithdiphycercaltailandwithacontinuousdorsal,caudalandanalfin;dentitionusually
consistsofcrushingtoothplates
c.Paleoecology
modernspeciesarefoundinfreshwaterbutextincttypesinhabitedawidevarietyof
environments
essentiallyallwerebottomdwellingomnivores
oldertypesseemtohavereliedonlungs
burrows(foraestivationduringdryseasons,wheretheycansurviveinasemiinanimatestatein
aflaskshaped,mucuslinedpit)arefoundfromtheDevoniantoRecent
3.Coelacanths(Actinistia)
coelacanthsandosteolepiformsarebonyfishwithtwodorsalfins,lobatepairedfins,and
cosmoidstructureofthescalesanddermalbones;mostimportantdistinguishingfeatureis
braincasedivisionintoananterior(ethmosphenoid)andposterior(oticooccipital)partswithan
intracranialjointbetweenthem;smallplatessurroundeye
predominantlypredatoryfishesandmainlyreliedonsenseofsmell(withsmalleyes)
fossilsrangefromMiddleDevoniantoUpperCretaceous(bothmarineandfreshwater);with
one(orpossiblytwo)livingmarinespecies(Latimeriachalumnae)
featuresdifferingfromosteolepiformsinclude:braincasejuncturepositiondifferentfrom
osteolepiforms;nomaxilla;upperjawattachedtobraincase
4.Osteolepiforms("Rhipidistia",inpart)

PALEOLECTURE,PAGE103
importantfishsincethey(oracloselyrelatedgroup)gaverisetotheamphibians
large,voraciousfish
Range:MiddleDevoniantoLowerPermian
Characteristics:mostdifferfromcoelacanthsinthearrangementandnumberofdermalbonesin
thehead(infactthebonesintheheadsofosteolepiformsarelargelyhomologoustothoseof
primitivetetrapods);usuallypossesschoanae(internalnares)
otherimportantfeatureswhichlinktheosteolepidfishestoearlytetrapodsinclude
labyrinthodontteeth(withinfoldedplicidentine),singlebasalelementsinthepectoralandpelvic
fins(homologoustothehumerusandfemur)whicharticulatewiththelimbgirdles;pairsof
radialsarticulatewiththesinglebasals(thesepairsarehomologoustotheradiusandulnaortibia
andfibulaoftetrapods);axialskeletonislikerhachitomoustypefoundinsomeearlytetrapods
(consistsofawedgeshapedintercentrumandsmallpairedpleurocentra)
XXIII.TheGreeningoftheLand
includedhereisadiscussionoftheKingdomsFungiandPlantae(Metaphyta)
A.KingdomFungi
heterotrophsthatsecreteenzymesabletobreakdownexternalfoodsourcesintomolecules
smallenoughtobeabsorbedbycells(extracellulardigestionandabsorption);areamajorgroup
ofdecomposers;Saprobic/saprophytictypesfeedonnonlivingorganicmatter;parasitictypes
feedonlivingorganisms
1.Classification
theFungiisprobablyapolyphyleticgroupandthereforeisinvalid;includestheDivisions
Gymnomycota(slimemolds),Mastigomycota(flagellatefungiorphycomycetes;includingthe
classesOomycetesandChytridomycetes)andtheAmastigomycota(nonflagellatedfungiortrue
fungi;includingtheclassesZygomycetes,AscomycetesandBasidiomycetes)
thereareabout250generaand500speciesoffossilfungi,mostlyfromCretaceousandTertiary
2.EvolutionandtheFossilRecord
theAscomycetesandBasidiomycetesevolvedinCambrianandOrdovicianseasas
heterotrophicfungilivingonalgae
duringtheSilurian,fungimovedtolandenvironmentstoliveassaprophytesandparasiteson
theearlylandplants
allclassesoffungiwerepresentbytheendofthePennsylvanian
fungievolvedandradiatedintonewenvironmentspioneeredbythefloweringplantsduringthe
CretaceousandTertiary;thisadaptiveradiationisprobablyduetocoevolutionoffungiwiththe
evolvingangiosperms
B.KingdomPlantaeClassification
*SubkingdomorDivisionBryophyta(mosses,liverworts,andhornworts)

PALEOLECTURE,PAGE104
*SubkingdomorDivisionTracheophyta(vascularplants)
ClassRhyniopsida(primitivevascularplants)
OrderRhyniales
ClassPsilopsida(whiskferns)
ClassZosterophyllopsida(ancestorsofmicrophyllousplants)
OrderZosterophyllales
OrderAsteroxylales(prelycopods)
ClassLycopsida(clubmossesandtheirrelatives)
OrderDrepanophycales
OrderProtolepidodendrales
OrderLycopodiales
OrderSelaginellales
OrderLepidodendrales
OrderIsoetales
ClassTrimerophytopsida(ancestorsofmegaphyllousplants)
ClassSphenopsida(horsetailsandtheirrelatives)
OrderHyeniales
OrderPseudoborniales
OrderSphenophyllales
OrderEquisetales
ClassFilicopsida(fernsandtheirrelatives)
OrderCladoxylales
OrderStauropteridales
OrderZygopteridales
OrderOphioglossales
OrderMarattiales
OrderFilicales
OrderSalviniales
OrderMarsileales
ClassProgymnospermopsida(ancestorsofgymnosperms)
OrderAneurophytales
OrderArchaeopteridales
OrderProtopityales
ClassGymnospermopsida(plantswithnakedseeds)
OrderPteridospermales
OrderCycadales
OrderCycadeoidales
OrderCaytoniales
OrderGlossopteridales
OrderPentoxylales
OrderCzekanowskiales
OrderGnetales
OrderGinkgoales
OrderCordaitales

PALEOLECTURE,PAGE105
OrderVoltziales
OrderConiferales
OrderTaxales
SubdivisionAngiospermophytina(floweringplants)
ClassMagnoliopsida/Dicotyledonae(dicotyledons)
ClassLiliopsida/Monocotyledonae(monocotyledons)
*Dependinguponthescientist,thetaxonomiccategorywillvary.Somepaleobotanist's
"Subkingdom"isanother's"Division",or"Division"toonemaybea"Class"toanother,etc.(see
classificationsbelowforcomparison)
C.OriginofLandPlants(KingdomPlantae)isMonophyletic
evolvedfromtheChlorophyta(grassgreenalgae)
Synapomorphiccharactersinclude:
1.havechlorophyllaandb
2.storetruestarch
3.havecelluloseincellwalls
4.protostelewithxylemformingcore(xylemformspipelinesforconductingwaterand
dissolvedminerals),phloemonoutside(phloemisthefoodconductingtissue;foundinrootsof
mostvascularplants;stemsofmanypsilopsids,lycopods,sphenopsidsandferns)
5.lifecyclessimilar(alternationofgenerations,etc.)
D.CharacteristicsoftheKingdomPlantae(*preventsdesiccationinterrestrialenvironments)
1.fertilizedeggdevelopsintoanembryowhichisenclosedwithinaprotectivecovering*
2.protectionofsporesandpollengrainsbytoughwallimpregnatedwithsporopollenin*
3.ParenchymaTissuescontinuetoliveaftermaturity;maycombinewithothercellstoform
complextissues*
4.Waxylayersonleavesandbranches*
5.Guardcellsofstomata(openingsintheleaves)*
6.AlternationofGenerations
multicellulargameteproducingorganisms(gametophytes)alternateinthelifecyclewith
multicellularsporeproducingorganisms(sporophytes)

PALEOLECTURE,PAGE106

E.ClassificationandCharacteristicsoftheKingdomPlantae
1.SubkingdomBryophyta
includesmosses,liverwortsandhornworts(maybeapolyphyleticgroupandsomepropose
establishmentofthreeseparateDivisions)
nostiffenedvasculartissueforconductingwaterandnutrients;gametophytesmostimportant
phaseofsexualreproduction
featuresusefulforlanddwellingexistenceincludepresenceofwaterconservingcuticleonthe
abovegroundparts,presenceofprotectivecellularjacketaroundthespermandeggproducing
partsoftheplanttokeepthemfromdryingout,andthesporophytebeginsearlydevelopmentas
anembryoinsidethetissuesofthefemalegametophyte
poorfossilrecord(DevonianRecent)
2.SubkingdomTracheophyta
vascularplants[withconductingcells(xylemandphloem)fortransportingwaterand
nutrients];usuallypossessroots,stemsandleaves
withapproximately11to13Divisions
Themostimportantdivisionsare:
a.Division/ClassRhyniophyta(Rhyniopsida)
MiddleSiluriantoMiddleDevonian
oldestknownvascularplants(MiddleSilurianofIreland);bestfossilsfromtheRhynieChert
(LowerDevonian,Scotland);Examples=Rhynia,Cooksonia
noleavesorroots(thereforewithphotosynthesisoccurringintheoutercellsofthestem);with
dichotomousbranchingofthestems(branchbyproducingtwoequalsegments);stemscappedby
sporebearingcases(sporangia);sporesarehomosporous(withonetypeofspore)andsporeis
trilete(roundortriangular,withthreefoldsonit)
b.Division/ClassPsilopsida
"whiskferns";thelivingwhiskbroomlookinggenusPsilotumisverysimilartothe
Rhyniophytawiththreedimensionaldichotomousbranchingandlowdegreeoforgan
differentiationwithrootsandvascularizedleavesabsent
Butsporangiaareborneterminallyonshortlateralaxes(whereasonRhyniopsidaareterminal
ontheaxial,ormain,portionsoftheplant)
c.Division/ClassZosterophyllopsida
UppermostSilurianthroughMiddleDevonian;Example=Zosterophyllum
ancestorsofmicrophyllousplants
alsoleaflessandbrancheddichotomously,butsporangiaarebornealongthesidesoftheaxis;
sporangiakidneyshapedandattachedtoashortstalk;branchesforkintotwoaxes,onegrows
upwardandonedownward("Hbranched",probablyallowedplanttospreadoutwardfroma
centerandtobeattachedtothesoilasitgrewout)

PALEOLECTURE,PAGE107

d.Division/ClassLycophyta(Lycopodophyta,Lycopsida)
DevonianRecent;includesthemodernclubmossesandquillworts;alsoarborescent(treelike)
lycopodsupto40metershigh(OrderLepidodendrales;DevonianPennsylvanian;dominated
Carboniferouscoalswamps)
evolvedfromtheRhyniopsida
havetrueroots(oratleastrootlikeorgans),stemsandoftenwithsmallleaveswithasingle
strandofvasculartissueandNOleafgap("microphylls");leavesontrunkinpits(leaf
cushions/bolsters)andarrangedinspirals
oftenwithconeshapedclustersofleavesbearingsporesacs(eachclusterisaStrobilus);spores
dispersedfromsporesacsandgerminatetoformsmall,freelivinggametophytes;moderntypes
requirewaterinwhichspermcanswimtotheeggs(thereforerestrictedtowethabitats)
Examples=Lepidodendron,Lepidophloios,Sigillaria(stems/trunks),Stigmaria(rootsor
rhizophores),Lepidophylloides(leaves)
e.DivisionSphenophyta(Sphenopsida)
includethemodernhorsetailsandseveralextinctgroups(DevonianRecent)
withscalelike,smallleavesarrangedinwhorlsaroundanaboveground,bamboolikejointed
(withnodesandinternodes)photosyntheticstemthatishollowinside;wallsofstemcellscontain
silica,sostemsoften"gritty"(hencethename"scouringrushes");sporesforminsidecone
shapedclustersoftinybranchesattheshoottipsandaredispersedbyaircurrents;sporesmust
germinatewithinafewdaystoproducegametophytes(freelivingplants)
aretypicallyfoundinswamps,moistwoodlands,andalonglakeedges
commonfossilsincludeCalamites(aPennsylvanianagearborescentsphenopsid;some
membersupto15metersormoreinheight)andAnnularia(leafwhorls)
f.DivisionFilicinophyta(Filicopsida,Pteridophyta)
includefernsandtheirallies(UpperDevonianRecent)
large,complexleaves[megaphylls;withleafgaps(spacesdevelopedwheretheleafstalk
(petiole)joinsthevascularcylinderofthetwig)];fernleaves(fronds)usuallyfeatherlikewith
bladesfinelydividedintosegments(leafpinnatelycompound,withtheleafletsarrangedalong
thesidesofacommonaxis)anddevelopsrachis,pinnaandpinnules;immaturefrondsunroll
(circinate)inmostmembers;oftenwithclustersofsporecases(Sorus)onleafundersides,with
sporangiumsnappingopenatdispersaltimetocausethesporestocatapultintotheair;
germinatingsporedevelopsintoasmallgametophyte
Treefernswerelarge,arborescentferns(MississippianPermian)foundincoalsswamps(Exs.
=Psaronius,Pecopteris)
arborescentlycopods,sphenopsidsandtreefernsbecameextinctwhentheLatePaleozoiccoal
swampenvironmentdeclined,probablyduetoclimatechangesresultingfromtheformationof
Pangaea
g."Gymnosperms"
probablyapolyphyleticgroup;includesconiferophytes,pteridospermophytes,cycadophytes,
cycadeoidophytes,ginkgoes,andfourplantgroupsofuncertainaffinities

PALEOLECTURE,PAGE108
gymnosperms(andangiosperms)arecharacterizedbyseeds;typicallyformedbyfusionofegg
andspermnuclei;thendevelopintoripenedovules(=seeds)
gymnospermshavenoflowersandseedsarenotfullyenclosed(gymnospermmeans"naked
seed")
g1.Division/Class/OrderPteridospermophyta
includethe"SeedFerns";UpperDevonianJurassic
withfernlikecompoundleaves(Fronds)butgymnospermlikeseedsandwood
Exs.=Alethopteris,Neuropteris,Medullosa(trunk)andpossiblyGlossopteris(leaves)
g2.Division/Class/OrderCordaitales
cordaites;PennsylvaniantoMiddlePermian;talltreesabout15to30metershigh
leavesoftenswordorstrapshapedwithdichotomousvenations(withforkingleafveins);
leaveswerebornespirallyinacrownnearthetopoftheplant
petrifiedstemswithpithareacrossedbystrandsofparenchymacells(thetissuethatmakesup
thebulkofthefleshyplantparts)separatedbyairspaces
seedbearingconelikestructuresandpollenbearingconesborneonspecialsmallbranches
g3.Division/Class/OrderConiferales
conifers(Exs.=pines,spruces,firs,hemlocks,junipers,cypresses,redwoods);TriassicRecent
woodytreesandshrubswithneedlelikeorscalelikeleaves;mostareevergreens(shedleaves
throughoutyearbutretainenoughofthemtodistinguishthemfromdeciduoustrees);conifers
haveCones(coneshapedclustersofmodifiedleavesbearingthesporangia);oftentwokindsof
cones(maleconesbearmicrosporesandfemaleconesdevelopmegaspores);seedsdevelopon
theshelflikescalesofthefemalecones
g4.Division/Class/OrderGinkgoales
ginkgos(?Permian;TriassicRecent;onerecentspecies,the"maidenhairtree")
oftenconiferliketreeswithamaintrunkbearingbrancheswithaxillarylongandshortshoots;
leavesareusuallyfanshaped;someleavesaredeeplylobed,othersarenot;venationofthe
leavestypicallyparallel,althougheachveinisdivided
g5.Divisions/Classes/OrdersCycadophyta(cycads)andCycadeoidophyta(cycadeoids)
cycads(PermianRecent)andcycadeoids(TriassicCretaceous)areoftendifficulttodistinguish
asfossils(bothoftenformshrubbyortreelikeplantswithpinnate,straplike,palmlikeleaves
andsimilarwood)
butcycadshavemaleandfemalecellsindistinctlydifferentconelikestructureswhereasin
cycadeoidsthemaleandfemaleconesareverysimilar
XXIV.Amphibians,AncientandModern
A.EcologyandOriginofTetrapods
1.The"Classic"Theory

PALEOLECTURE,PAGE109
landinvasionduringLateDevonianwhenpondsandstreamsdriedoutperiodically
evidenceincludesDevonian"redbeds"(wetdryseasonality);alsoearliestamphibians
(Ichthyostega)werefullyterrestrial(?)
2.Occupancyoflandtookplaceunderwarm,moistclimates
lowoxygencontentinthewater,populationpressures(seekingfood,competitionforspace,
breedingsites,escapefrompredatorsoreggeaters)
evidencefromstudiesofmodernairbreathingfishandamphibiansandcontradictoryevidence
aboutthegeologicalconditionsoftheDevonian
3.Amphibiansoriginatedinthewaterandwere"preadapted"toalifeonland
limbs,hearing,airbreathingandfeedingmodificationsrelatedtolifeinwarm,shallowwater
withlowoxygencontent
terrestrialradiationscamemuchlaterfromthevariouslinesofaquatictetrapods
B."Amphibians"
theearliesttetrapods(fourfootedcreatures)arefromUpperDevonianrocks
somecladisticclassificationsstatethatthe"Amphibia"shouldonlyincludethemodern
lissamphibians
1.OriginsandAdaptations
a.derivedfromtheosteolepiformfishes,efficientairbreatherswhichlivedinwarm,shallow
watersometimeslowinoxygen
b.lungsbecamemoreefficient(butgillbreathingretainedinimmatureandlarvalstagesof
primitivetypes)
c.evolutionofpairedappendages(duetosubstratelocomotion?);asairbreathingbecamemore
efficientthepectoralfinwasusedtolifttheheadoutofwater;finsbecamethetetrapodlimbs
withthelossofthefinrays
2.PrimitiveAmphibianStructure
a.Externalshapeandform
typicallythesmallandprimitivetypestendtoberelativelyhighandroundbodied
bonyscalesreducedbutancienttypesoftenretainedVshapedventralarmor
b.AxialSkeleton
veryimportantintheevolutionoftheearlytetrapods
vertebralcolumnhighlyossified
Vertebraltypesinclude:

PALEOLECTURE,PAGE110
Lepospondylous(husk)vertebrae(foundinmanysmallPaleozoictypes;centrumformsa
singlestructure,oftenspoolshapedandwithholefornotochord)
"Arch"vertebrae[foundinlabyrinthodontsandinallhighervertebrateclasses;consistsoftwo
setsofossifiedarchstructures(intercentraandpleurocentra)]including1)Rhachitomous
vertebrae[mostprimitivetypewithwedgeshapedcrescenticintercentraandpaired
pleurocentra(smallandbetweenneuralarchesandintercentra);foundinrhipidistians,
ichthyostegidsandmost"central"labyrinthodonts(sometemnospondyls)],
2)Stereospondylousvertebrae[pleurocentrareducedorabsentandintercentrumbelowthe
neuralarchandsometimesringshaped;foundinmanyLatePermianandTriassic
temnospondyls]and3)Anthracosaurs[pleurocentraincreasinginsize,fuseandbecomea
completering(=centrumofhighervertebrates);linewhichleadstoreptileshaveintercentra
reducedtosmallventralwedgesbetweenthepleurocentra;secondline(theembolomerous
condition)witheachsegmenthavingtwocompleteringshapedcentralstructures]
Neuralarcheswelldevelopedandwithzygapophysesgivingaddedsupporttothevertebral
column;hastransverseprocessforarticulationwiththeribs
Ribstetrapodswithonlyonesetofribs;foundfromnecktoanteriorpartoftail;reinforcethe
bodywallandprotectsthelungsandviscera;specializedrib(Sacral)connectspelvicgirdleto
vertebralcolumn
c.Skull
primitivetetrapodswithskullcompletelyroofedbydermalbones
tetrapodswithlongersnoutandshorterskulltable
anteriorandposteriorpartsofbraincasebecomesfuse
MajorBonesoftheSkullbonesdownmidlineincludeelongatenasalsandfrontals;shortened
parietalsandpostparietals;premaxillaandmaxillawelldevelopedandtoothed
d.Sensoryorgans
The"Ear"fishhyomandibularnolongerneededtopropthejaws,becomesloosened;land
vertebratesneedtopickupairvibrations(for"hearing");hyomandibularlaidjustbehindthe
spiracle,becomesfreetoformaStapes(probablyfirstfunctionedtosupportbraincaseagainst
thecheek);latertransmitedvibrationsfromtheTympanum(theeardrum;primitivelysituatedin
anoticnotchattherearoftheskullroof)totheinnerear
Laterallinesystemfishhydraulic"sensorysystem"typicallyretainedinwaterliving
Paleozoicamphibiansbutlostinhighervertebrateclasses
TheEyestypicallylargeintetrapodsandofteninprimitivetypessurroundedbysclerotic
plates
e.LimbGirdles
Shouldergirdleskullconnectionlost;humerusfitsintolaterallyplacedconcavity
Pelvicgirdlemuchlargerthanplateseeninfishesandhasjoinedtosacrumabove;consistsof
anilium,ischiumandpubis;femurfitsintolaterallyplacedcavity

PALEOLECTURE,PAGE111

f.Limbs
Pectorallimbshumerus(proximalbone)articulateswiththeradiusandulna;radiusandulna
articulatedistallywiththecarpus(wristelementsformingahingebetweenthelimbsandtoes);
typicallywithfourorfivetoes
Pelviclimbsproximalbone(femur)articulateswiththetibiaandfibula;tibiaandfibula
articulateswiththetarsus;tarsalsarticulatewiththemetatarsals(fivetoes)
g.ReproductiveSystems
probablyinitiallywithexternalfertilizationandlaidlargenumbersofsmalleggsinwater;
manyPaleozoictypeswithgillbearinglarvalstage(referredtoas"branchiosaurs")
3.Classification
PhylumChordata
SubphylumVertebrata(Craniata)
InfraphylumGnathostomata
SuperclassTetrapoda
UnnamedClass

FamilyIchthyostegidae
ClassAmphibia(Batrachomorpha)
Order"Temnospondyli"
FamilyTrimerorhachidae
SuborderCapitosauria
SuborderTrematosauria
InfraclassLissamphibia
OrderUrodela
OrderAnura
ClassUnnamed
SuperorderLepospondyli
Order"Microsauria"(probablyapolyphyleticgroup)
OrderNectridea
OrderAistopoda
SuperorderReptiliomorpha
Order"Anthracosauria"
OrderSeymouriamorpha
OrderDiadectomorpha

4.Ichthyostegids
earliesttetrapodsbutnotancestraltoothergroups;UpperDevonian(?)LowerMississippian
someoftheearlytetrapodshaveasmanyas8toes(Ichthyostegahad7)thatweredeveloped
intopaddlelikeappendages;ichthyostegidswereprobablylargelyaquaticandcouldnotfully
supporttheirweightonland

PALEOLECTURE,PAGE112

5."Temnospondyls"
labyrinthodontamphibiansthatevolvedfromosteolepiformfishes;primitivefeaturesinherited
fromthesefishincludelabyrinthineinfoldingofdentine,palatalfangedteeth,vertebrae
composedofseveralcentraelements
derivedfeaturesincludeformationofspecializedanteriorvertebraeforconnectingtoskull
(atlasaxiscomplex;notfoundinichthyostegids),oticnotchatbackofskull(supportedear
drum?),primitivetypesoftenlarge(over1meter)
MississippianCretaceous;werethemostimportantCarboniferoustetrapods
oftenwithlarge,flatheads;examplesincludetheaquaticeryopoidsandtrimerorhacids;the
terrestrial,armoreddissorophidsandthemetoposaurs(largeskulledaquaticamphibians)]
6."Lissamphibians"
maybeapolyphyleticgroup;mayhavebeenderivedfromdissorophidtemnospondyl
amphibians(orotherssaywerederivedfromlepospondyls)
includesfrogsandtoads(SalientiaorAnura;TriassicRecent),thelongbodiedaquatic
salamanders(UrodelaorCaudata;JurassicRecent)andthewormlikecaecilians(Apodaor
Gymnophiona;JurassicRecent)
withpedicellateteeth(baseandcrownoftoothseparatedbyazoneofweaknessoffibrous
tissue;probablyrelatedtotongueprotrusion);teethbicuspid;spoolshapedvertebrae;frogsand
salamanderswithstrangeearspecializationbywhichanearossicle(theoperculum)hasa
muscularconnectionwiththeshouldergirdle(probablyrelatedtohearingandbalance)
FrogsandToadsgreatlyderived(mostfeaturesrelatedtojumping):only5to9trunkvertebrae;
posteriorofsacrumfused(urostyle);noribs;iliumrodshapedandconnectstolastvertebrae;
longhindlegswithtibiaandfibulafused;radiusandulnafused;proximaltarsalselongate;
shouldergirdlefirmlybraced;skullforms"open"structure;externalfertilization;tadpolestage
(herbivoroussuspensionfeedersoralgaeeaters)knownfromLowerCretaceous;adults
carnivorous;advancedtypesflipbackoftongueoutofmouth
Triadobatrachus(EarlyTriassic,Madagascar)withfroglikeskullbutpostcraniumnotvery
froglike;ancestraltomodernfrogs?
7.Lepospondyls
usuallysmall,MississippiantoPermianageamphibians;mayhaveevolvedfromearly
labyrinthodonts(butwithnolabyrinthineinfolding,nopalatalfangsandpitsandnooticnotch)
characterizedbylepospondylousvertebrae(spoolshapedbonycylindersurroundingthe
notochord)
includesthesnakelikeaistopodsandlysorophids,theeellikenectridians[Diplocaulusand
Diploceraspiswithboomerangheads],andthelizardliketerrestrial"microsaurs"(manybelieve
microsaursareapolyphyleticgroup)
8."Anthracosaurs"andtheirKin
DevoniantoPermian;aparaphyleticgroupsincetheyareancestraltoreptiles(oftenplaced
withintheReptiliomorpha)

PALEOLECTURE,PAGE113
Seymouriamorphswithcombinationofreptileandamphibianfeatures;Amphibianfeaturesof
Seymouriaincludeanthracosaurlikeskullandgillbearinglarvae;Reptilelikefeaturesinclude
skullandcheeksolidlyattached;stapes(earbone)reducedtoanarrowrod;iliumexpandedand
beginincorporatingsecondsacralrib;vertebraewithswollenneuralarchesandlarge
pleurocentrumwithtinyintercentrum
9.Diadectomorphs
PennsylvaniantoEarlyPermianterrestrial"reptiliomorphs",veryclosetotheoriginof
amniotes
herbivorouswithpeglikefrontteethandgrinding"molariform"teeth
XXV.AMyriadofReptilesonLand
A.CharactersofReptiles:
1.Developmentofamnioteegghasalargeyolk,ashell,andextraembryonicmembranes
whichprotecttheegg,supplynourishmentandforgasexchange
Amniotesprobablyamonophyleticgroup;probablyoriginatedintheMississippian
2.lossofintertemporalbone;reductioninsizeofsupratemporal,tabularandpostparietal
3.lossofpalatal"fangs"andlabyrinthineinfoldingoftoothenamel
4.absenceofoticnotch
5.developmentofmoreefficientvertebralcolumn(specializedanteriorvertebrae=atlas/axis)
andreduceintercentra
6.specializedanklejointdeveloped[astragalusandfibulare(=calcaneum)]
7.wheatshapedventralscales;noossifieddorsalscales
B.PaleoecologyofEarlyReptiles
probablyterrestrialsince:
1.earlyreptilesusuallyfoundinterrestrialdeposits
2.limbs,girdlesandvertebralcolumnareossifiedandwelldevelopedforterrestriallife
3.smallbodysize,structureofteethandprobablearrangementofjawmusculatureprobably
indicatelizardlikehabitsforearlyamniotes(probablyatesmallterrestrialarthropods)

PALEOLECTURE,PAGE114
C.ReptileClassificationandRadiation
oftenbasedonpatternsofopeningsofskullroof(temporalopenings)behindtheorbits
1.Anapsidconditionnotemporalopening;Exs.=captorhinids,turtles
2.Synapsidconditionloweropeningwithpostorbitalandsquamosalmeetingabove;Ex.=
mammallikereptiles
3.Diapsidconditiontwotemporalopeningspresent;Exs.=dinosaurs,pterosaursand
ancestralconditionofallmodernreptilesexceptturtles
4.Euryapsid(Parapsid)conditionupperopeningwithpostorbitalandsquamosalmeeting
below;Exs.=plesiosaurs,ichthyosaurs
derivedfromthediapsidconditionthroughlossofthelowertemporalfenestra
D.ClassificationofPrimitiveReptiles(Oneofseveralavailable):
SeriesAmniota
ClassSynapsida

OrderPelycosauria
FamilyCaseidae
FamilyOphiacodontidae
FamilyEdaphosauridae
FamilySphenacodontidae
OrderTherapsida
FamilyDinocephalia
FamilyDicynodontia
FamilyCynodontia
ClassSauropsida
SubclassAnapsida
FamilyProcolophonidae
FamilyPareisauridae
FamilyMesosauridae
FamilyCaptorhinidae
OrderTestudines(Chelonia)
FamilyProganochelyidae
SuborderPleurodira
SuborderCryptodira
E.TheAnapsids
mostprimitiveformswhichareunquestionablyreptilian;consistsof"parareptiles"
(milleretiids,bolosaurs,procolophonids,pareisaurs)andtheturtles
typicallysmall(0.3to0.9meterslong),lacktemporalopeningandoticnotch;massivestapes
propsbraincase

PALEOLECTURE,PAGE115

1."Protorothyeridae"(=Romeriidae,Millerettidae,etc.)
apolyphyleticgroupofPennsylvanianPermian,small,lizardlikereptiles
2.Bolosauridae
small,earlyherbivores
3.TheProcolophonoids
developherbivorousfeeding(jawsshorten,differentiationoftheteeth,enlargeorbitotemporal
fenestraforlargejawmuscles],latetypeswithbonyskullprojections
4.ThePareiasaurs
MiddletoUpperPermian;uptothreemeterslong;skulloftenwithbonyprotruberances;leaf
shapedteeth(herbivores)
5.Captorhinids(Captorhinidae)
withspecializeddentitionwithmultipletoothrows
probablymorekintodiapsidsthanotherearlyamniotegroups
F.Synapsids
haveoftenbeentermed"mammallikereptiles",butsynapsidsarenowtypicallyconsideredto
beagroupdistinctfrom"truereptiles";theSynapsidaoftenincludespelycosaurs,therapsids,and
truemammals
synapsidsweretheearliestcarnivorousamniotes(byPennsylvanianconstituted50%ofknown
amniotegenera;byEarlyPermian=70%)
1.GeneralCharacteristics
withasinglelateraltemporalopening(synapsidcondition;primitivetypeswithpostorbitaland
squamosaljoiningabove;derivedtypeswithuppermarginboundedbyparietal)
2.Pelycosaurs
PennsylvanianPermian
probablyderivedfromprotorothyridcaptorhinomorphs
includesOphiacodonts(carnivores);Sphenacodonts(highlypredaceousformssuchas
Dimetrodon;manywithelongateneuralspinesforming"sails"),Edaphosaurs(herbivores;
usuallywithelongateneuralspineswithcrossbars)andCaseids(herbivores)
3.Therapsids
Tetraceratops(EarlyPermian,Texas)maybetheoldesttherapsid(isintermediateinform
betweenthesphenacodontpelycosaursandtherapsids)
a.GeneralCharacteristics
mandiblewiththin,extensivesheetofbone(thereflectedlamina;forhearing);temporal
fenestra(opening)largerthanpelycosaurs;singlecanine,jawhingeanteriorlyplacedandback

PALEOLECTURE,PAGE116
ofskullisvertical;Skeletonwithimprovedlocomotion
derivedfromadvancedsphenacodonts
b.ClassificationofTherapsids
includetheDinocephalians(verylargePermiancarnivoresandherbivores);Anomodonts
(PermiantoTriassic;herbivorous;mostsuccessfulmammallikereptiles;includethetusked
dicynodonts);Cynodonts(PermiantoJurassic;advancedmammallikereptilesrepresenting
transitionalstagesinthedevelopmentofmammaliancharacteristics)
XXVI.FarewellstoLand
A.Mesosaurs
aquaticparareptilesofPermianagefromAfricaandSouthAmerica;probablyrestrictedtoone
limitedoceanbasinandwasusedasevidenceofcontinentaldrift
uptoonemeterlong,slender;withlong,laterallycompressedtailandneckandpaddlelike
feet;marginalteethlongandslender(forstrainingmicroplankton?)
B.Testudines(Chelonians)
theturtles
probablycloselykintopareisaursandprocolophonids
1.Structure
shellcomposedofhornyscutescoveringbonyplates;withcarapace(dorsalportionofshell)
andplastron(ventralportionofshell)
vertebrae[exceptcervicals(neck)]andribsfusedtoshell;limbsandlimbgirdlesmodifiedfor
sprawlingposture
anapsidskull;teethrudimentaryorabsent
2.Classification
divideintotheProganochelyds(primitiveTriassicageturtles),Pleurodires(sidenecked
turtles),andCryptodires(Sneckedturtles;mostsuccessfulturtlegroup)
C.Subclass(orOrder)Ichthyopterygia(Ichthyosauria)
Dolphin,tunaandsharklikeneodiapsidreptilesoftheMesozoic
bodyshort,laterallycompressedandfusiform
skullhighlymodifiedforaquaticlife(longbeak;nostrilsmigratedfarposteriorly;eyesgreatly
enlargedandsurroundedbybonyplates);withaeuryapsidskullpattern
vertebralcentraamphicoelous(biconcave);tendencythroughtimetodevelopahypocercaltail;
limbsreducedtosteeringpaddles
reproductionprobablytookplaceinwaterandwithlivebirth(somefemaleswithskeletonsof
youngichthyosaursinsidethem)
D.TheSauropterygians(SuperorderSauropterygia)

PALEOLECTURE,PAGE117
lepidosauromorphneodiapsidsthatincludethenothosaurs,pachypleurosaurs,plesiosaurs,and
possiblytheplacodonts;aquaticreptileswitheuryapsidtemporalopening
Nothosaurs(limbsrelativelynormal;Triassic)andPlesiosaurs(developpaddlesbyaddingtoe
joints;JurassicCretaceous)withnostilsmigratedfarbackonskull;ventralribsformbasketlike
structure;ventralportionofpelvicgirdleexpanded
E.Placodonts
wierdTriassic,aquaticmollusceatingneodiapsids(butwitheuryapsidtemporalopening);
mostwith"pavementteeth";kintothe"Sauroptergyia"andnowoftenplacedwithinthatgroup
XXVII.ScaleBearersandLizardHippedDinosaurs
A.Diapsids
diapsidshavetwotemporalopeningsseparatedbythepostorbitalandsquamosal
includesallmodernreptilegroupsexceptturtles;alsoincludesdinosaurs,pterosaurs,
plesiosaursandseveralotherancientgroups
1.ClassificationoftheDiapsids(oneofmanyavailable):
SeriesAmniota
ClassSauropsida
SubclassDiapsida
InfraclassIchthyosauria
InfraclassLepidosauromorpha
SuperorderSauropterygia
OrderPlacodonta
OrderNothosauroidea
OrderPlesiosauria
SuperorderLepidosauria
OrderSphenodontida
OrderSquamata
SuborderLacertilia
SuborderSerpentes(Ophidia)
InfraclassArchosauromorpha
FamilyTrilophosauridae
FamilyRhynchosauridae
FamilyProlacertiformes
DivisionArchosauria
FamilyProterosuchidae
FamilyErythrosuchidae
FamilyEuparkderiidae
SubdivisionCrurotarsi
FamilyPhytosauridae(=OrderPhytosauria)

PALEOLECTURE,PAGE118
FamilyOrnithosuchidae
FamilyStagonolepididae(=OrderAetosauria)
FamilyPrestosuchidae
FamilyPoposauridae(=OrderRauisuchia)
OrderCrocodylia
SubdivisionAvemetatarsalia(Seebelow)
MostimportantGroupsAre:
B.InfraclassLepidosauromorpha
includessphenodontids,lizards,snakes,andtheextinctaquaticplacodonts,nothosaurs,and
plesiosaurs
differentiatedfromarchosauromorphsbyretentionofsprawlingposture;lateralundulationof
bodyduringmovement;earlylepidosauromorphswithlargesternum("breastplate"forgreater
flexionandstride)
OrderSphenodontida(FamilySphenodontidae)lizardlike,smallreptilesfromTriassic
Recent
OrderSquamataincludesSubordersSauria,AmphisbaeniaandSerpentes
Sauria(lizards)withtendencytowardsstreptostyly(withlossoflowertemporalbarandloose
connectionofposteriorskullbonesforgreaterbitingforce);teethprimitivelysubpleurodontor
pleurodont(attachedtoinsideofjaw);maysecondarilybecomeacrodont(teethfusedtotopof
jaw)orsubthecodont(teethinshallowpits);includesmanytaxa;Cretaceousmarinelizards
includedolichosaurs,aigialosaursandmosasaurs
Serpentes(Ophidia)=snakes(UpperCretaceousRecent);temporalarchesabsentandupper
andlowerjawsverylooselyattached(forconsuminglargeprey);vertebraeverynumerous;
pectoralgirdleandanteriorlimbabsent(somesnakeswithvestigesofpelvicgirdleandhind
limb);implantationofteethsubacrodont
C.PrimitiveArchosauromorphs(InfraclassArchosauromopha)
mostimportantstructureunitingarchosauromorphsisankle(tarsus)andfootstructure(related
touprightposture)
MostImportantGroupsare:
1.FamilyTrilophophosauridae
smalltomediumsized,Triassicage,lizardlike"herbivorous"reptiles(teethtypically
tricuspid);postcranialskeletonlikeprimitivearchosaurs;Ex.=Trilophosaurus
2.FamilyRhynchosauridae
heavilybuiltTriassicherbivorouslepidosaurians;advancedtypeswithupperjawwithbroad
crushingtoothplatesandparrotlikeedentulous(toothless)beak

PALEOLECTURE,PAGE119
D.DivisionArchosauria
the"rulingreptiles"includingthedinosaurs,crocodiles,pterosaursandmanyprimitivegroups
(thethecodonts)
1.Characteristics
Skullwithdiapsidcondition(twotemporalopenings);openingsdonotloseanyoftheirarcades
("arches");alsowithanantorbitalopening(rarelymorethanone)betweentheorbitandthe
snout;thecodontdentition(teethplacedinsockets)
Postcranialskeletonwithhindlimbmuchbetterdevelopedthantheforelimb;tendencytowards
bipedalposeinvolveschangeinhipandfemurstructure
2.SubdivisionCrurotarsi
withimprovedanklejoint(tarsus)thatallowsrotationbetweentheastragalusandcalcaneum
tarsusmaybeusedtodivideMiddleandUpperTriassicarchosaursintotwogroups;
CrocodileNormalPatternwithprocessonthelateralsurfaceoftheastragalusfittingintoarecess
onthemedialsurfaceofthecalcaneum[foundincrocodiles,phytosaurs,aetosaursand
rauisuchids);CrocodileReversePatternwithprocessonthecalcaneumfittingintoarecessin
theastragalus(foundinlagosuchids,ornithosuchidsandEuparkeria)
a.FamilyPoposauridae(=Rauisuchia)
large,fierceMiddleandUpperTriassicthecodonts(upto6meters)
b.FamilyStagonolepididae(=Aetosauria)
relativelylargeherbivorousquadrupedsofLateTriassicage;bodywithsolidarmorplate
c.FamilyPhytosauridae(=Phytosauria)
veryabundant,crocodilelike,UpperTriassicthecodonts
E.Crocodilians(OrderCrocodilia)
Crurotarsithatincludethecrocodiles,alligatorsandtheirrelatives
1.Morphology
Skullelongate,flattened,massive;evolutionarytrendinposteriorextensionofthepalatalbones
(premaxillae,maxillae,palatinesandpterygoids)toformasecondarypalate(foraquaticmodeof
lifeortosupporttheelongatesnout?);elaboratepneumaticducts(forhearingairbornesounds?)
Bodyelongatewithlong,flattail;gastralia(ventralribs)present;amphicoelous(biconcave)or
procoelousvertebrae(concavityinfront,withconvexposteriorsurface);dermalarmor;witha
semiimprovedgait[hindlegslongerthanfrontlegs;pelvistriradiate;improvedtarsaljoint]
2.Classification
includestheProtosuchiaandtheMesoeucrocodylia[largestassemblageofcrocodiles;include
manyMesozoicmarinetypes;posteriorextensionofpalatinebonestoformsecondarypalate;
MesoeucrocodiliaincludestheEusuchia(includesgavials,alligatorsandcrocodiles;secondary
palatefullydeveloped)]

PALEOLECTURE,PAGE120

F.Dinosaurs
Over800speciesofdinosaursareknown
1.Posture
limbsbroughtunderthebodyandmovedinaforeandaftdirection[femurinturned;pelvis
"socket"(acetabulum)open(perforate);improvedtarsaljoint(withformationofamesotarsal);
digitsformmainsurfacethatcontactsground(digitigradeposture)]
2.Ancestry
probablyderivedfromOrnithosuchians(closetoLagosuchus)
dinosaursbecamethedominantreptilegroupafterthe"endCarnianExtinctionEvent"during
theUpperTriassic,whichclearedecospaceforthedinosaurstotakeover
3.ClassificationofPterosaursandDinosaurs
Thefollowingisoneofseveralavailable:
SeriesAmniota
ClassSauropsida
SubclassDiapsida
InfraclassArchosauromorpha
DivisionArchosauria
SubdivisionAvemetatarsalia

InfraorderOrnithodira
OrderPterosauria
SuborderRhamphorhynchoidea
SuborderPterodactyloidea
SuperorderDinosauria
OrderSaurischia
FamilyHerrerasauridae
SuborderTheropoda
InfraorderCoelophysoideaFamilyCoelophysidae(Procompsognathidae?)
InfraorderCeratosauriaFamiliesCeratosauridae,Abelisauridae
InfraorderTetanurae
DivisionCarnosauria
SubdivisionSpinosauroideaFamiliesMegalosauridae,Spinosauridae
SubdivisionAllosauroideaFamiliesAllosauridae,Carcharodontosauridae
DivisionCoelurosauria
FamilyCoeluridae
SubdivisionManiraptoriformes
FamiliesTyrannosauridae,Ornithomimidae
InfradivisionManiraptora

PALEOLECTURE,PAGE121
FamiliesAlvarezsauridae,Therizinosauridae
CohortDeinonychosauria
FamiliesDromaeosauridae,Troodontidae
SuborderSauropodomorpha
FamiliesPlateosauridae,Massospondylidae
InfraorderSauropoda
FamiliesVulcanodontidae,Euhelopodidae,Omeisauridae
DivisionNeosauropoda
FamiliesCetiosauridae,Diplodocoidea
SubdivisionMacronaria
FamilyCamarasauridae
InfradivisionTitanosauriformes
FamiliesBrachiosauridae,Titanosauridae
OrderOrnithischia
FamiliesPisanosauridae,Fabrosauridae
SuborderThyreophora
FamilyScelidosauridae
InfraorderStegosauria
InfraorderAnkylosauria
FamiliesNodosauridae,Ankylosauridae
SuborderCerapoda
InfraorderPachycephalosauria
InfraorderCeratopsia
FamiliesPsittacosauridae,Protoceratopsidae,Ceratopsidae
InfraorderOrnithopoda
FamiliesHeterodontosuridae,Hypsilophodontidae,Iguanodontidae,
Hadrosauridae
4.Saurischians
withprimitivepelvisstructurewithiliumattop,pubispointingforwardandischiumbackward;
frontlimbshorterthanhind;digitsofmanus(hand)andpes(foot)reduced;teethoccupiedthe
rimsofthejaws;largeopeningsreducedtheweightoftheskull
saurischiansdominatedduringtheearlyMesozoicbutwereoutnumberedbytheornithischians
duringtheupperMesozoic
ThemostImportantSaurischiansare:
a.Theropods
includeallofthebipedalcarnivorousdinosaurs
neckisgenerallyshorterthanthetrunk;thetibiaislongerthanthefemur;handsbearsharp
clawsandtherearetwoorthreefingersonly;feetwiththreeclawedtoes(thefifthisalways
reducedandthefirstorbigtoeisshortenedandturnedbackwards);abdominalribspresent
Range:LateTriassicCretaceous
includesthecoelophysoids(small,slendertheropoddinosaurs),ceratosaurs(manywithhorns

PALEOLECTURE,PAGE122
andcrests)andthetetanurans[withanopeninginthemaxilla(maxillaryfenestra),anddorsal
vertebraehavecavitiesintheirsides(pleurocoelousvertebrate)andtheascendingprocessofthe
astragaluscoversaportionofthetibia;includestheCarnosaurs(spinosaursandallosaurs)and
theCoelurosaurs(mostimportantcoelurosaurtypesaretheManiraptoriformes(reductionto
threelongfingers;withahalfMoonshapedwristbone)includingornithomimids("ostrich
dinosaurs"),tyrannosaurids,anddeinonychosaurs("raptors")]
b.Sauropodomorphs
typicallyheavilybuiltquadrupedaldinosaurswithsmallheadsandlongnecks;mostwith
pegliketeeth;UpperTriassicUpperCretaceous
"Prosauropods"smalltolargesized;possibleancestorsofsauropods;carnivorous,
herbivorousandperhapsomnivorousforms
SauropodswerehugeJurassic/Cretaceousherbivoreswithquadrupedalpose,powerfullimbs,
longtail,longneckandsmallhead;jawsshortandweakwithsmallpeglikeorspoonshaped
teeth;frontlegsshorterthanhindlegs;metapodials(proximalfootbones)andphalanges(toes)
short,stoutandspreading;includeCetiosaurids,Brachiosaurids,Camarasaurids,Titanosaurids
andDiplodocids
XXVIII.BirdHippedDinosaurs
A.CharacteristicsofOrnithischians
pubispointsbackward(birdhipped);withsinglemedianboneatthetipofthelowerjaw(the
predentary);jawwithbeak,posteriortowhichisagrindingdention;mostwithconcavecheek
region(thereforemostwithmuscularcheeks);tendencyforinternalnostrilstobedisplaced
posteriorly
B.ClassificationofOrnithischians
1.Cerapoda
withagapbetweentheteethofthepremaxillaandmaxillaandwithfiveorfewerpremaxillary
teeth;withathicklayerofenamelontheinsideoftheteeth
includesornithopods,pachycephalosaursandceratopsians
a.Ornithopods
hadbirdlikefeetwithbluntclawsorhooves;includeFabrosaurids,Heterodontosaurids,
Hypsilophodontids,Iguanodontids(Cretaceous)andHadrosaurids("duckbilled"dinosaurs)
b.Pachycephalosaurs
smallgroupofLateCretaceous"boneheaded"dinosaurs(withunusuallythickskullroofs,
probablyusedfor"buttingcontests"betweenmales)
c.Ceratopsians
smalltolargedinosaurswithskullsrangingfromrelativelylargetogigantic,oftenwithhorns

PALEOLECTURE,PAGE123
andlargeshieldsofbone;snoutbeaklike;almostexclusivelyquadrupedal;oneoflastevolved
(Cretaceous)andmostabundantgroupsofdinosaurs
2.Thyreophorans
withkeeledscutes(bonyarmor)alongthesidesoftheirbody
includestegosaursandankylosaurs
a.Stegosaurs
MidJurassictoLateCretaceousarmoredquadrupedalornithischians
relativelylarge;withsmallskull,frontlegsshort;backarchedhighoverlonghindlimbs;with
seriesofplatesandspinesarrangedinarowdowntheneck,trunkandtail
b.Ankylosaurs
stockydinosaurswithshort,broadfeet;withextensivedevelopmentofbony,armoredcarapace,
oftenwithtailclub;LowerJurassicUpperCretaceous
C.BiologyandExtinctionofDinosaurs
1.WereDinosaursWarmBlooded?
Evidencecitedthatdinosaurswereendothermsincludesthefollowing:
a.Erectposturelimbsheldvertically(withmetabolismlikebirdsandmammals)
b.Bonestructuredinosaurshavehaversiancanalsystemsintheirboneslikethoseof
mammals(indicatesmorerapidmetabolicprocesses;buttheseseemtobepresentinlarge
animalsingeneralandareabsentinsmallanimals)
butdinosaursdidnothavedeterminantgrowthandcontinuedtoincreaseinsizethroughoutlife
(unlikebirdsandmammals)
c.PopulationStudies/CommunityStructurecarnivorousdinosaurnumbers(versus
herbivores)aremorelikethatofmammalsthanreptiles
d.Longneckeddinosaurswouldhavetohaveamoreefficientheartinordertopumpbloodup
totheirbrains
e.Afewdinosaurswereatleastasintelligentasbirds
f.Dinosaursshowsocialbehavior(suchasherdingand"nurseries")thatisunknownamong
otherreptiles
g.DinosaurshavebeendiscoveredinMesozoic"polarregions"
h.Somedinosaurfossilshavefeathers

PALEOLECTURE,PAGE124

i.GrowthRatesreptilesgrowslowly,dinosaursgrewquicklylikebirdsandmammals
j.OxygenIsotopesratiosareinfluencedbytemperature;indicatesmoresimilaritytomodern
endothermsthanectotherms
However,dinosaurswouldprobablymaintainarelativelyconstantinternaltemperaturedueto
theirsmallsurfaceareaversusvolume(HomeothermsorGigantotherms).Also,ifdinosaurswere
suchgreatendothermswhyalloftheplates,frills,spikesandnasalcavitiesthatprobablyserved
asheatexchangers,helpingtowarmandcooltheirbodies?
2.ExtinctionoftheDinosaurs
dinosaursoriginatedintheMiddleTriassicandbecameextinctattheendoftheCretaceous
(Cretaceous/Tertiary,K/T,orMaastrichtian/DanianBoundary;approximately65Ma)[somesay
afewsurvivedintotheearlyCenozoic]
a.CatastrophicDinosaurExtinction
a1.ExtraterrestrialCauseslargeasteroid(10to20kilometersacross)hittheearth,creatinga
cloudofdustandsomethingsimilarto"nuclearwinter";decreaseinphotosynthesis,increasein
carbondioxide,increaseinacidityofoceansandashortterm"greenhouseeffect"?
evidenceincludestheiridiumlayerattheCretaceous/Tertiaryboundaryfoundatabout50
localitiesthroughouttheWorld(probablydepositedoveraperiodnomorethanafewthousand
years),thepresenceofglassyspherules(tektites)and"tsunamibeds"attheK/Tboundary
a2.Vulcanologymodelsgeochemicaldatainboundaryrocksindicatemajorvolcanic
eruptions(e.g.,TheDeccanTrapsofIndia)attheendoftheCretaceous
wouldproducegreenhousegasesthatwouldtriggerrapidclimatechange
b.HypothesisofGradualChange
endofCretaceousmarkedbymajorregressionanddryingupofepicontinentalseas
tectonicactivity,mountainbuildingledtomajorchangeinclimateandseasonality
WesternNorthAmericamayhaveseengradualdecreaseintemperaturebetweenlate
CretaceousandPaleoceneof10C;evidenceincludesgradualextinctionandreplacementof
dinosaursandothergroups(includingplesiosaurs,pterosaurs,ostracods,bryozoans,ammonites
andbivalves,allwithlowdiversityattheendoftheCretaceous)
XXIX.FlyingandGlidingReptiles
PterosaursactiveflyingreptilesfromtheUpperTriassicthroughUpperCretaceous;mostfrom
shallowmarineenvironments;mostgenerashortlivedandfromsmallgeographicareas
A.Classification

PALEOLECTURE,PAGE125

SeriesAmniota
ClassSauropsida
SubclassDiapsida
InfraclassArchosauromorpha
DivisionArchosauria
SubdivisionAvemetatarsalia
OrderPterosauria
SuborderRhamphorhynchoidea
SuborderPterodactyloidea
B.Morphology
1.Skull
typicallylarge;bonestendtofuseinskull;largebrains;largeorbits;quadrateslantedanteriorly
andstreptostylic,forincreasedjawmobility;nostrilsmigratedposteriorly;withlongbeakand
long,sharpteeth
2.Postcranialskeleton
neckelongate;trunkveryshortwithlongsacralregion;rhamphorynchoidswithlongtail
ActiveFlightindicatedby:hollowbonesandwithbirdlikepneumaticforamina(for
respiration;withhighmetabolicrate?);keeledsternum;pterodactyloidswithwelldeveloped
shouldergirdle[scapula(shoulderblade)articulatedwithfusedanteriortrunkvertebrae
(notarium)];humerusformspulleylikestructure(forattachmentoflargeflightmuscles);carpal
("wrist")bonemodifiedtoformsplintlikepteroid(servesasanchorforanarrowmembranethat
extendstothebaseoftheneck);firstthreefingersshort,fourthfingergreatlyelongatetosupport
thewingmembrane(attachedtoposteriorportionoftrunk),fifthfingerabsent
rhamphorynchoidshadlongtails;pterodactyloidshadshorttails
XXX.Birds,InventorsoftheFeather
ClassAves(Birds)verydifficultvertebratefossilstoworkwith;generallypoorpreservation
andconservativemorphology(beneaththefeathers)
mostdiversificationintheCenozoicbutmostmodernfamiliespresentbyendofEocene
taxonomicinterrelationshipsaredebated,evenamongmanymoderngroups
A.Characteristics
highestmetabolicrateofanyvertebrate
manymodificationsinconnectionwithhabits,socialorganizationandflight
1.Skeleton
bonesarepneumatic(withextensiveairsacsystemforrespiration)
compactskeletonwithwingandlegbonesarereducedinnumberandmanyelementsfused

PALEOLECTURE,PAGE126
[includinghand,foot,sacralvertebrae(fusedtoformasynsacrum),tailvertebrae(fusedtoform
apygostyle)]
ribswithuncinateprocesses(smallbonystrutsthatbindtheribcagetogether)
claviclesfusedtoformthefurcula(the"wishbone");stabilizestheshoulderjointandprevents
collapseoftheshoulderduringflight
sternum("breastbone")withlargekeel(carina)thatprovidesabroadbasefortheflight
muscles
typicalavianfootwiththreetoesinfrontandonebehind(anisodactyly);someunrelatedgroups
withtwotoesinfrontandtwobehind(theyoketoedorzygodactylcondition;includes
woodpeckers,cuckoosandparrots)
2.Skull
withlargeorbits(incompletelysurroundedbybone),infrontofwhichisanantorbitalopening;
singletemporalopening(butderivedfromthediapsidcondition)
largebraincase;skullbonestypicallyfusedandsuturesobliterated
modernbirdstoothlesswithbeakcoveredwithahornybill
B.Originofflight
theoriescanbedividedintotwogroups,the"arboreal"theoriesand"cursorial"theories
1.Arborealtheoryhasbeenproposedbymostworkers;fourfooted,grounddwellingreptile
becamebipedal,thenclimbing,thenbeganleapingfromtreetotree.Lateritbeganparachuting,
glidingandfinallyincludedactive,poweredflight
2.Cursorialtheoryfeathersdevelopedasthermoregulatorydevicesforinsulation;thenused
fortrappinginsects;thenprovidedliftduringrunningandleaping;thenflight
C.ClassificationofBirds(Oneofmanyavailable):
ClassAves
SubclassPygostylia

FamilyArchaeopterygidae

FamilyConfuciusornithidae
OrderOviraptosauria
InfraclassOrnithothoraces

OrderEnantiornithes
SupercohortOrnithomorpha
CohortOrnithurae
OrderHesperornithiformes
SubcohortCarinatae
OrderIchthyornithiformes

PALEOLECTURE,PAGE127
SuperdivisionNeornithes
DivisionPalaeognathae
OrdersLithornithiformes,Ratites
DivisionNeognathae
SubdivisionGalloanserae
OrdersAnseriformes,Galliformes
SubdivisionNeoaves
Superorderunnamed['waterbirdassemblage']
Infraorderunnamed
OrdersGruiformes,Ralliformes
Infraorderunnamed
OrdersPelecaniformes,Ciconiiformes
InfraorderUnnamed
OrdersCharadriiformes,Phoenicopteriformes,Podicepidiformes,
Falconiformes,Procellariformes,Gaviiformes,Sphenisciformes
OrderStrigiformes
Superorderunnamed
OrdersApodiformes,Caprimulgiformes
OrderMusophagiformes,Columbiiformes,Psitaciformes,
Cuculiformes
Superorderunnamed['derivedlandbirds']
OrdersPiciformes,Coliiformes,Trogoniformes,Bucerotiformes,
Coraciformes,Passeriformes
Therearemanygroupsofbirds.Iamonlygoingtocoverthoseofpaleontologicinterest:
1.Archaeopterygids
includesArchaeopteryx,theearliestknownbird(pigeonsize);Jurassic
nouniquefeaturesinthebonyskeletontodifferentiatethemfromdinosaurs(dinosaurfossils
fromChinaindicateatleastsomedinosaurshadfeathers)
Skullbirdlikewithanexpandedbraincaseandlargeeyes;suturesmostlyclosed;but
Archaeopteryxhadthecodontteeth
Skeletonwithvertebralcolumnprimitivewithamphicoelous(biconcave)vertebrae;tail
dinosaurlikewithtwolateralrowsoffeathers;hindlegsandpelvissimilartosaurischian
dinosaurs;claviclesjoinedtoformabirdlikefurculabutnokeeledsternum
2.ToothedBirdsandDivers
includesHesperornithiformes(loonlike,flightless,toothedCretaceousbirds),Grebes
(Podicipediformes),loons(Gaviiformes),penguins(Sphenisciformes;wingbonesformaswim
fin),pelicans(Pelecaniformes;2meterlongplotopteridswerelargestswimmingbirds),
albatrosses/shearwaters(Procellariiformes)andIchthyornithiformes(ternliketoothed
Cretaceousbirds)
3.WaterBirds

PALEOLECTURE,PAGE128
includeOrdersCharadriiformes(shorebirds,includingplovers,snipes,gulls,terns,auks,and
sandpipers),Anseriformes(ducks,geeseandswans)andCiconiiformes(wadingbirdssuchas
storks,herons,bitterns,ibises,spoonbills)
CharadriiformsmaybeancestraltoallotherlargewaterbirdsexcepttheCiconiiformes;most
LateCretaceousfossilbirdsthatarenothesperornithiformsbelongtotheancientshorebirds
(example=Telmatornis)
4.FlightlessBirds
Ifthereisnocontinualselectionforthemaintenanceofflightapparatusbirdstendtobecome
flightless
majorchangesincludelossoftheflightmuscles;reductioninthewingandbonesofthe
pectoralgirdle;lossofthekeeledsternum;obtuseangleofthescapulocoracoidarticulation;
withbroadunossifiedregionbetweentheiliumandischium(theilioischiaticfenestra);some
ratiteswithskullsuturesapparent;tendencytobecomelarge(notethatmostoftheseareneotenic
featuresi.e.theyretainthe"youthful"condition)
"Ratites"includegiganticflightlessbirdssupposedlydifferentiatedbytheirpalaeognathous
palate;isprobablyapolyphyleticgroupthatincludesmoas(NewZealand;someoverthree
meterstall),elephantbirds(upto500kilograms),ostriches,rheas,cassowaries,emus,tinomous
andkiwis
NeoavianbirdgroupsthatdevelopedflightlessmembersincludetheordersGruiformes(cranes,
rails,andthegiantEarlyTertiarycarnivorousphorusrhacids),Diatrymiformes(overtwometer
tall,probablycarnivorous,flightlessbirdsfromtheEarlyTertiary)
5.BirdsofPreyincludethehawksandowls
a.Falconiformesincludesfalcons,hawks,eagles,vultures,ospreysandsecretarybirds;with
raptorialadaptations(sharplyhookedbeak;powerfulfeetwithlongclawsandanopposabletoe;
strongfliers)
b.Owls(Strigiformes)nocturnalbirdsofprey;featuressuchastheshorthookedbeakand
powerfultalonsareconvergentwiththefalconiforms;PaleoceneRecent
6.LandBirds
verydifficulttofigureoutphylogenies;arebeginningtobeworkedoutbyusingdiagnostic
bonessuchasthestapes
mostimportantgrouparethePasserines(OrderPasseriformes);songbirds;withover5000
modernspecies(threefifthsofalllivingbirds);placedinfrom50to70familiesofSuboscines
andOscines;withperchingfootconsistingofaverylargefirsttoedirectedstraightbackand
opposedtotheotherthree;differentiatedbymorphologyofthepalatalbones(the
aegithognathouscondition),bythespermstructure,bythevoiceboxandstapesstructure
XXXI.HairyReptileswithComplexEars:theEarlyMammals

PALEOLECTURE,PAGE129
A.ClassificationofMammalsthisisoneofseveralavailable:
ClassMammalia
SubclassMammaliaformes

FamilySinocondontidae
FamilyMorganucodontidae

InfraclassHolotheria

FamilyKuehneotheriidae
OrderDocodonta
SuperdivisionAustralosphenida
DivisionMonotremata
SuperdivisionTheriimorpha
OrderTriconodonta

DivisionTheriiformes
OrderMultituberculata
SuperlegionTrechnotheria
OrderSymmetrodonta
LegionCladotheria
SuperfamilyDryolestoidea
SublegionBoreosphenida
OrderDeltathroida
InfralegionTheria
CohortMarsupialia
MagnorderAmeridelphia
OrderDidelphimorphia
FamilyDidelphidae
OrderPaucituberculata
FamiliesCaenolestidae,Argyrolagidae,
Caroloameghinidae
OrderSparassodonta
FamiliesBorhyaenidae,Thylacosmilidae
MagnorderAustralidelphia
OrderMicrobiotheria
OrderDasyuromorphia
OrderPeramelemorphia
OrderNotoryctemorphia
OrderDiprotodontia
CohortPlacentalia(Eutheria)
MagnorderAfrotheria
OrderTubulidentata
OrderAfrosoricidaFamiliesTenrecidae,Chrysochloridae
OrderMacroscelidea
GrandorderPaenugulata

PALEOLECTURE,PAGE130
OrderHyracoidea
MirorderTethytheria
OrderSirenia
OrderProboscidea
FamiliesMoeritheriidae,Deinotheriidae
SuborderElephantiformesFamiliesMammutidae,
Gomphotheriidae,Stegodontidae,Elephantidae
MagnorderXenarthra
OrderLoricata(Cingulata)FamiliesDasypodidae,
Glyptodontidae
OrderPilosaFamiliesMyrmecophagidae,Bradypodidae,
Megalonychidae,Megatheriidae,Mylodontidae
MagnorderBoreoeutheria
OrderLeptictida
OrderAnagalida
OrderApatemyida
OrderTaeniodonta
OrderTillodontia
OrderPantodonta
OrderPantolesta
OrderDinocerata
GrandorderLaurasiatheria
OrderLipotyphlaSubordersErinaceomorpha,
Soricomorpha
OrderChiropteraOrdersMegachiroptera,Microchiroptera
MirorderFerungulata
SuperorderCetartiodactyla
OrderArctocyonia
OrderMesonychidae
OrderArtiodactyla
FamilyDichobunidae
SuborderSuiformes(Bunodontia)Families
Entelodontidae,Suidae,Anthracotheriidae,
Hippopotamidae
SuborderSelenodontia
InfraorderTylopodaFamilies
Merycoidodontidae,Camelidae
InfraorderRuminantiaFamilies
Hypertragulidae,Tragulidae,Antilocapridae,
Giraffidae,Cervidae,Mochidae,Bovidae
OrderCetaceaSubordersArchaeoceti,Odontoceti,
Mysticeti
OrderPerissodactyla
SuborderHippomorphaFamiliesEquidae,

PALEOLECTURE,PAGE131
Brontotheriidae

SuborderAncylopodaFamilyChalicotheriidae
SuborderCeratomorphaSuperfamilies
Tapiroidea,Rhinoceratoidea
?SuperorderBulbulodentataFamilyHyopsodontidae
SuperorderMeridiungulataFamiliesLitopterna,
Notoungulata,?Astrapotheria,?Pyrotheria
SuperorderUnnamed
OrderCreodonta
OrderCarnivora
FamilyMiacidae
SuborderFeliformia
FamilyNimravidae
InfraorderAeluroideaFamiliesViverridae,
Herpestidae,Hyaenidae,Felidae
SuborderCaniformia
FamiliesCanidae,Ursidae,
Amphicyonidae,Mustelidae,Procyonidae
InfraorderPinnipediaFamilies
Enalilarctidae,Otariidae,
Odobenidae,Desmatophocidae,
Phocidae
OrderPholidota
GrandorderEuarchontoglires
SuperorderArchonta

SuborderPlesiadapiformes
OrderPrimates
SuborderStrepsirrhini
InfraorderAdapiformes
InfraorderLemuriformesFamiliesLemuridae,
Indriidae,Daubentoniidae,Lorisidae,
Galagidae
SuborderHaplorhiniFamiliesOmomyidae,
Tarsiidae
SuborderAnthropoidea
InfraorderPlatyrrhiniFamiliesCebidae,Atelidae
InfraorderCatarrhini
FamiliesOligopithecidae,Parapithecidae,
Propliopithecidae
SuperfamilyCercopithecoideaFamily
Cercopithecidae
SuperfamilyHominoideaFamilies
Proconsulidae,Hylobatidae,Hominidae
OrderScandentia

PALEOLECTURE,PAGE132
OrderDermopteraFamiliesParomomyidae,
Galeopithecidae
SuperorderGlires
FamilyZalambdalestidae
OrderRodentia
SuborderSciurognathi

SuperfamilyIschyromyoidea
InfraorderSciuromorpha
InfraorderMyomorpha
SuborderHystricognathi
InfraordersHystricomorpha,Phiomorpha,
Caviomorpha
OrderLagomorpha
B.CharacteristicsofMammals
1.SoftAnatomy
havehairandspecializedmammaryglandsforsucklingtheiryoung
platypusandechidnasinegglayingOrnithodelphia(Prototheria);marsupials(Metatheria)have
amarsupium(apouchinwhichmostembryonicdevelopmenttakesplace);placentals(Eutheria)
havedevelopmenttakingplaceintheuterusandtheembryoisnourishedbytheplacenta(the
tissuesshedfollowingabirth)
mammalsareintelligentwithcomplexbehavioralpatterns
mammalsareendothermicandusuallyhavehighmetabolicrates
2.Osteologicalfeatures
a.Skeleton
determinategrowth(growrapidlybutachievedefinitivebodysize);oncegrownepiphyseson
longbonesfusewiththeshaft;cynodontsandearlymammalswithatlas/axisvertebraeforming
ringinlivingmammalsforrotationofheadonvertebrae);mammalswithdorsoventralflexion
betweendoubleoccipitalcondyle(posteroventral"knobs"onskull)andatlas;trunkvertebrae
andribsintherapsidstoearlymammalsrestrictlateralflexionandenabletrunktobeflexedina
sagittalplane;appendicularskeletonmodifiedforuprightposture
b.Skullfeatures
usualdefinitionofamammalosteologicallyispresenceofadentarycondyleonthejaw
articulatingwithaglenoidinthesquamosal;incisors,caninesandpremolarsreplacedonlyonce;
molarsnotreplaced,withtworootsandspecificpatternofocclusion;withsinglebonynasal
openingintheskull
MiddleEarOriginsreptilesandbirdswithsingleearossicle(stapes);mammalearwithchainof
threebones(malleus,incusandstapes)whichconductvibrationsfromthetypanumtotheinner
ear;formationofdentarysquamosalarticulationfreesarticular(mandiblebone)tobecomethe

PALEOLECTURE,PAGE133
malleusandthequadrate(skullbone)becomestheincus
c.Teeth
veryimportantinmammalianpaleontology
mammalsarediphyodont(withtwotoothgenerations;deciduousandpermanent)
molarshavetwoormorerootsandcomplexcrownmorphology
livingmonotremeslackfunctionalteeth;othermammalswithfourclassesofteethineachjaw;
anteriorteeth(incisors)simpleinform;caninesingle,largeandconicalandisanteriormost
toothofthemaxilla;teethimmediatelybehindupperandlowercaninesarepremolars[usually
becomeincreasinglymolariforminstructureposteriorly;incisors,caninesandpremolarsusually
withdeciduousprecursorsshedearlyinlife];molarswithnodeciduousprecursorsanddonot
eruptuntilanimalbeginstoreachitsdefinitivebodysize
uppermolarsusuallywithsquarecrownswiththreerootsandlowermolarswithnarrower
rectangularcrownsandtworoots
numbersofteethineachclass(incisors,canines,premolarsandmolars)summarizedinadental
formula(commonopposumDidelphiswithformulaI5/4,C1/1,P3/3,M4/4andgeneralized
placentalswithdentalformulaI3/3,C1/1,P4/4andM3/3)
marsupialsandplacentalsderivedfromCretaceoustherianmammalswithtritubercularor
tribospheniccheekteeth(bothupperandlowermolarswiththreeprominenttuberclesorcusps)
uppermolarswithcuspsformingatrigon;consistsofamedialprotocone,anteriorparacone,
andposteriormetacone
lowermolarswithcuspsformingatrigonid;consistsofalateralprotoconid,anteriorparaconid
andposteriormetaconid
fourthcusp,ifpresent,isalmostalwaysdirectlybehindtheprotoconeorprotoconidandis
calledthehypocone(upper)orhypoconid(lowermolar)
fifthcusp,theentoconid,isoftenpresentonlowermolars;theentoconidandhypoconid
encloseaheel(=talonid)posteriortothetrigonid
trigonsandhypoconesonsuccessiveuppermolarsformaseriesoftrianglesformedbythe
trigonid,hypoconidandentoconidonlowermolars;upperandlowercheekteethoccludein
differentwaysandcanbedeterminedthroughstudiesofcuspspatternsandwearfacets
molarswithcusps(pointsforpuncturing),crests(linesforshearing)andbroadbasinedareas
(planarareasforcrushingorgrinding)
C.PrimitiveMesozoicMammalsandMonotremes
nearlyallearlymammalswereverysmall(probablyupto10cmand2030grams)
1.Monotremes(EarlyCretaceous?Recent)
havemilkglands,hairandonelowerjawelementbutlayeggsandhavemanyreptilian
characteristicsintheirskeletonsandsoftanatomy
includestheduckbillplatypusandspinyanteatersorechidnas
2.Triconodonts
LateTriassictoLateCretaceous
basicdentalstructurewithtricuspidalignmentonthemolars

PALEOLECTURE,PAGE134
includesthemorganucondontids,amphilestidsandtriconodontids
3.Symmetrodonts
shrewsizedmammalsknownfromjawfragmentsanddentition;UpperTriassicUpper
Cretaceous
mandibleslenderandlong;lowermolarstriangularwithasymmetricalcusparrangement
dividedintothekuehneotheriids,AmphidontidsandSpalacotheriids
4.Docodonts
MiddletoLateJurassic;wereprobablysizeofsmallmouseandwithelongatesnouts
hasarticularquadratearticulationbutprimaryhingeisthedentarysquamosal
developedadvanced"squarecusped"molarpatterns

5.Multituberculates
mostdiverseandnumerousMesozoicmammals
rodentlike;LateJurassicEarlyOligocene;replacedtherodentliketherapsids(tritylodonts)in
themidJurassicandwerelaterreplacedinthePaleocene(bycompetitionwithcondylarths,
primatesandrodents?)
skulllowandbroadwitheyesfacinglaterally;withpairofenlargedprocumbentlowerincisors;
withlowmanycuspedmolariformteeth
6."TheriansofMetatherianEutherianGrade"
awastebasketcategorytoincludetherianswithtribosphenicdentitionbutnotsufficiently
knowntoplacewiththemarsupialorplacentalgroups
includespoorlyknownCretaceoustherianssuchasthe"TrinityTherians"(Kermackia,
PappotheriumandHoloclemensiafromthePaluxy/AntlersFormationofTexas)
theoldestundisputedeutherianisawellpreservedshrewsizedplacentalfromtheEarly
CretaceousofMongolia,whichprovesthatthereweretrueplacentalmammalsbythistime
D.Marsupials
youngborninaveryimmature(altricial)developmentalstateandundergosubsequentgrowth
attachedtotheteatsofthefemale(usuallyinanabdominalpouch)
nowoccuronlyinNorthAmerica,SouthAmericaandAustralia;fossilsfromEarlyMiddle
CenozoicofEurope,earlyCenozoicofnorthAfricaandearlyCenozoicfromAntarctica
1.Marsupial(Metatheria)versusPlacental(Eutheria)Characteristics
MARSUPIALS

PLACENTALS

1.Postorbitalbarusuallylacking

1.postorbitalbarpresent

2.Braincaserelativelysmall

2.braincaserelativelylarge

3.Posteriorpalatalvacuitiesusuallypresent 3.posteriorpalatalvacuitiesrare

PALEOLECTURE,PAGE135

4.apoorlydevelopedauditorybulla

4.auditorybullacommonlywelldevelopedandof
variousoriginsderivedfromalisphenoid

5.Angleofjawusuallyinflectedmedially 5.lowerjawusuallynotinflected
6.Dentalformuladerivedfrom
I5/4C1/1P3/3M4/4

6.DentalformuladerivedfromI3/3C1/1P4/4M3/3
(6premolarsinprimitivespecies)

7.essentiallymonophyodontteeth;only 7.diphyodontteeth;replacementofmostantemolar
thirdpremolarisreplaced
teeth

8.relativelybroadstylarshelfonupper
8.relativelynarrowstylarshelfinmostforms
molarsinmostpolyprotodontforms

9.hypoconulidandentoconidaretwinned 9.notwinning
andseparatedfromthehypoconid
10.Epipubicbone;bothsexesofmostforms 10.noepipubicbones
11.Nonepossessbaculumorosclitoridis 11.baculumorosclitoridiscommon
12.presumedtohaveretainedtheoriginal 12.withderived,placentalmodeofreproduction
therianreproductivemode

13.withaltricialyoung

13.precociousyoungcommon

13.retainsomewhatlowermetabolicrate 14.typicallydevelopanelevatedbasicmetabolic
rate

2.InterrelationshipsoftheMetatheria
classificationshavelargelybeenbasedonincisormorphologyandmodificationofbonesinthe
hindfoot
a.Incisormorphology
Polyprotodonttypicallymanyincisors(upto5aboveand4below);incisorsapproximately
equalsize;includesallknownNorthAmericanandlivingSouthAmericantaxa
Diprotodontupperincisorscommonlyreducedto3pairsorlessandlowerincisorsconsistof
onepairofprocumbentteeth;oneofeachincisorisenlargedrelativetoothers;canineslacking
andwithlongdiastemaseparatingincisorsfromcheekdentition;cheekdentitionstrongly
bilophodont;includesmost,butnotall,Australianforms
b.Footmodifications

PALEOLECTURE,PAGE136
SyndactylymodificationofbonesofhindfootfoundinmanyAustralianforms;digitsIIand
IIIofhindfootreducedandincorporatedinasingledermalsheath(probablyforgrooming)
Didactylynofootmodification
2.ClassificationoftheMetatheria
ThisisaclassificationschemethatwasmodifiedfromWoodburne(1984)fortheMarsupials(I
haveonlyincludedthemostcommontaxa):
InfraclassMetatheria
a.CohortAmeridelphia
includesAmericanMarsupials
a1.OrderDidelphimorphia
polyprotodontanddidactylous;LateCretaceousRecent;includestheFamiliesDidelphidae
(mostprimitivemarsupialsandprobablyancestraltoothertypes;includestheAmerican
oppossum),Necrolestidae(molelike),Argyrolagidae(kangarooratlike)
a2.OrderPaucituberculata
pseudodiprotodontanddidactyl;EarlyEocenetoRecentofSouthAmericaandLowerEocene
ofAntarctica;includesseveralgroupsofrodentlikemarsupials
a3.OrderSparassodonta
includestheBorhyaenidae(verysuccessfulgroupofmediumtolargesizeddoglikemarsupials
fromtheLatePaleocenePlioceneofSouthAmerica)andtheThylacosmilidae(lateMiocene
PlioceneofSouthAmerica;"sabretoothed"marsupials)
b.CohortAustralidelphia
includesAustralianMarsupials
b1.OrderDasyurida/Dasyuromorpha
includessuperfamiliesDasyuroidea[smalltomediumsizeddidelphidlikeinsectivores,
carnivoresandomnivores;dentalformula4/31/123/234/4;includesFamiliesDasyuridae
(shrewtowolfsizedcarnivorousmarsupials;"nativecats",Tasmaniandevil),Myrmecobiidae
(numbats)andThylacinidae("pouchedwolf")
b2.OrderNotoryctemorphia
includestheSuperfamilyNotoryctoidea(FamilyNotoryctidae;"marsupialmoles")
b3.OrderPeramelemorphia
polyprotodontandsyndactyl;includestheSuperfamilyPerameloidearectangulartooth
crowns[includingFamiliesPeramelidaeandThylacomyidae(bandicoots,rabbitlike)]

PALEOLECTURE,PAGE137
b4.OrderDiprotodonta
diprotodontandsyndactyl
Thisorderincludesthefollowingtaxa:
SuborderVombatiformes
SuperfamilyVombatoidea[includesFamiliesThylacoleonidae("marsupiallions";Miocene
Pleistocene),Vombatidae(wombats;burrowers),Palorchestidae(MiocenePleistocene;
oftengroundslothlike),Diprotodontidae[sheeptohippopotamussizedquadrupedal
marsupialsfromAustralia(Miocenerecentlyextinct)andNewGuinea(Pliocene
Pleistocene)]andWynyardiidae
SuperfamilyPhascolarctoidea[includesFamilyPhascolarctidae(koalas)]
SuborderPhalangeriformes/Phalangerida
SuperfamilyPhalangeroideamaybemostprimitivediprotodonts;includesFamilies
Macropodidae["ratkangaroos",wallabiesandtruekangaroosfromAustralia(Miocene
Recent)andNewGuinea(PlioceneRecent);dentalformula3/301/02/24/4]
Truekangaroos(macropodines)typicallylophodont(looplikeenamelpatternonteeth)and
withhighcrownedteeth;mostwithonlyoneortwomolarsfunctionalatanyonetime;most
aremediumtolargesizewithbipedalleapinggait(tibiagreatlyelongate,metatarsus
elongate;lossoftoes;long,strongtail)
E.PrimitiveEutherianMammals
classificationdifficultduetoincompletefossilrecordandverysimilardentitions;classification
maybeassistedbymorphologyoftheauditorybulla(bonessurroundingtheearregion)
"Insectivores"("OrderInsectivora"ofolderclassifications)
tremendousproblemsinclassification;postcranialskeletonprimitive[pentadactyl(fivetoed)
limbsandtypicallyplantigradelocomotion];typicallysmall,lackanossifiedauditorybulla,
relativelycompletedentitionwithsharpcuspedteethandcaninesoftenreduced
withapproximately60modernand150fossilgenera
includestheLiptotyphlans(hedgehogswithsquaremolarsduetodevelopmentofhypocone;
shrewsoftenwithpigmentedteethandwithparaconesandmetaconesformingaWshapedouter
wallontheuppermolars;moleswithpowerfulforelimbsfordiggingandparaconesand
metaconesformaWshape)
trueinsectivores(Liptotyphla)areasistergrouptothebats
XXXII.TheWorldBlossoms
DivisionAnthophyta(Angiospermophyta,Angiospermophytina,Angiospermae)flowering
plants(CretaceousRecent);includethemajorityofrecentplants
A.CharacteristicsofAngiosperms

PALEOLECTURE,PAGE138
withpollenproducingflowers(flowers=modifiedleaves);pollenwindcarriedorinsect
borne;pollenlandsonstigma(endportionoffemaleelement);pollentubegrowstotheovules
forthetransportofsperm;oneportionofthespermfertilizestheeggandanotherportionunites
withasecondportionoftheovule(whichgenerallyformsastructurewhichprovidesnutrients
forthegrowingembryo)="doublefertilization";seedthatdevelopsistotallyencasedinsidea
fruit
angiospermsareverysuccessfulbecausethediploidsporophytedominatesthelifecycle(asin
otherseedplants);thesporophyteoflanddwellingtypeshasrootandshootsystems,aswellas
otherfeaturestoallowittotakeupandconservewateranddissolvedminerals;thesporophyte
retainsandnourishesthegametophyteandtheembryosarenourishedbyauniquetissue(the
endosperm)withintheseed;theseedarepackagedinfruits,whichalsohelptoprotectand
dispersethem;evolutionofflowergreatlyledtothediversityoftheangiosperms
B.ClassificationofAngiosperms
1.ClassMonocotyledonae/Liliopsida
includegrasses,lilies,sedges,palms,pineapplesandorchids;?Triassic,CretaceousRecent
usuallywithfloralpartsingroupsofthree(e.g.,threestamensthepollenbearingportionsof
theflowers);leavesusuallyparallelveined;usuallywithonlyonecotyledon(the"seedleaf"of
theembryo);stemsareusuallyherbaceousandrarelyhavesecondarygrowth
2.ClassDicotyledonae/Magnoliopsida
includeherbsandwoodyplants,cacti,andwaterlilies;?Jurassic,CretaceousRecent
floralpartsoccuringroupsoffourorfive;vascularbundlesarearrangedinacirclearoundthe
pithofthestem;oftenwithsecondarygrowth(thickensthevascularbundlesandmakesthem
stronger);usuallyleavesarenetveinedandtheirembryoshavetwocotyledons("seedleaves")
thedicotsareprobablyparaphyletic;most"dicots"havetricolpatepollen(thesearetermed
"eudicots"or"tricolpates";otherdicotsandthemonocotyledonshavemonosulcatepollen(it
thereforeappearsthatthemonocotswerederivedfrommonosulcate"dicots")
C.TertiaryClimateandVegetation(NorthAmericanModel)
1.Broadleavedevergreen"gymnosperms"becameextinctattheCretaceous/Tertiaryboundary,
tobereplacedbydeciduousdicots
2.PaleocenetoEarlyEoceneglobalclimatebecamewarmerandprecipitationincreased,with
expansionoftropicalforeststoabout50to60Northlatitude
3.UpperEocenewithdeclineintemperatureanddrierclimates,andwithspreadofbroadleaf
deciduousforests
4.DuringMiocenewithdrierclimatesanddevelopmentofwidespreadgrasslands(grasseshave
continuouslygrowingleavesandcanwithstandheavygrazing),andwithadaptiveradiationof
"weeds"(theCompositae,mostlyannualorperennialherbscapableofrapiddevelopmentand

PALEOLECTURE,PAGE139
colonizingdisturbedhabitats)
XXXIII.TheGreatPlacentalRadiation
A.Bats(OrderChiroptera)
onlyorderofmammalsspecializedfortrueflight
innumberofspeciesarethesecondlargestorderofmammals
mayhavedescendedfromsoricomorphinsectivoresandarethesistergrouptothe"true
insectivores"(liptotyphlans)
1.Structure
Postcranialwingssupportedbyfourfingers;thumbisfreedandclawed;legsandpelvis
developedfor"hangingaround"
Skullorbitusuallyopenbehind;auditoryregiongreatlyenlarged;dentitionvaried;oftenwith
tribosphenicmolars(uppermolarstriangular)orsquaredwithaWshapedectoloph,alarge
protoconeandsmallerhypocone)
2.Classification
a.MegachiropteransorMegabats(SuborderMegachiroptera)
includethefruiteatingbatsor"flyingfoxes"oftheOldWorldtropics;OligoceneRecent;
welldevelopedeyes,largeolfactorylobes(goodsenseofsmell;usuallydonotecholocate);
molarsoftenspecialized
b.MicrochiropteransorMicrobats(SuborderMicrochiroptera)
includemostspeciesofbats;almostworldwideindistribution;LowerEoceneRecent;small,
nocturnal(activeatnight)andusuallywithsmalleyes;mostareecholocators;considerable
diversityindiet;uppermolarswithwshapedshearingcusps
B.Plesiadapiforms
previouslyplacedwithinthePrimates(andstillconsideredtorepresentasistergrouptothem)
lateCretaceousEocene;Plesiadapidshadarodentlikedentitionwithalongdiastema("gap")
betweentheprocumbentincisorsandgrindingmolarteeth(probablyherbivorousdiet)
C.Primates(OrderPrimates)
possiblethatprimatesandrodentsshareacommonancestorinthelateCretaceous
usuallyscansorial("scurrying"),smalltomediumsizedforestdwellingherbivoresor
omnivores
1.GeneralCharacteristics
Skeletonretainaprimitive,generalizedskeleton,withfivefingersandtoesonthehandsand
feet;trendtowardincreasingthemobilityofthethumbandbigtoe;orthograde(upright)posture;
typicallyclingorsitverticallywhenresting;locomotiongenerallyquadrupedal
Skullfacialpartofskullisreducedinmoreadvancedprimates;nasalapparatusgenerally

PALEOLECTURE,PAGE140
reduced;eyesfaceforwardonskull;brainrelativelylarge;oftenwithbroadlybasinedupperand
lowermolars;hypoconeusuallyaddedinuppermolars
2.TheStrepsirhines
withpostorbitalbarpresentontheskull;graspingthumbandbigtoe(higherdegreeofarboreal
adaptation);auditorycapsule(earregion)specialized
includeAdapids,Lemurs(OldWorldtropics;small,arboreal,nocturnal,furry,withfoxlike
face)andLorises
3.TheHaplorhinines(Tarsiiformes)
ectotympanicboneformsatubeleadingoutwardfromtheauditorybullaintheseandhigher
primates
includesOmomyids(EoceneMiocene)andTarsiids(LowerOligoceneRecent)
4.Anthropoids
monkeys,apesandman(lateEoceneRecent)
LateEoceneandearlyOligocenewithsharpincreaseinseasonalityandreducedtemperatures
innorthtemperatezones(numberoffossilprimatesdrop);LaterCenozoicwithfossilprimates
almosttotallylimitedtosouthernAsia,AfricaandSouthAmericaandconsistalmostentirelyof
anthropoidprimates
probablyomomyidancestry
withderivedfeaturesofskull;nevermorethanthreebicuspidpremolarteeth;uppermolars
usuallywithquadratepattern(cuspsin"square"pattern);braincaseexpandedandforamen
magnum(wherethebackboneconnectstotheskull)tendstobeundertheskull(thereforefaceis
turnedforwardalmostatarightangletothebackbone);twohalvesofthejawarefused
a.NewWorldPlatyrrhines(Ceboids)marmosetsandcebidmonkeys;mostprimitive
anthropoids;flatnoseswithpairedbutwellseparatedoutwardlydirectednasalopenings;dental
formulaI2/2C1/1P3/3M3/3;OligocenetoRecent
b.OldWorldCatarrhinesCercopithecidMonkeysandtheApes;protrudingmuzzle;nostrils
closertogetherandopenforwardanddownward;dentalformulaI2/2C1/1P2/2M3/3;upper
molarsquadrangular,highcrowned,bilophodont
substantialchangesinglobalclimatesduringMioceneduetonorthwardmovementofthe
Africanplate(createdAntarcticCircumpolarCurrentwithsavannahbecomingdominant);
catarrhinesdidwell,otherprimatesdidn't
c.Hominoids
earlytypesincludesmallbodied,longfacedPropliopithecidae(mostprimitivegroup),
Proconsulidae(AfricanDryomorphs)andDryopithecidae(EuropeanDryomorphs)
Ramamorphs(ancestraltoorangutan),Pongids(orangutan,chimpanzee,gorilla),hylobatids
(gibbons)oftenarboreal;pongidswithcaninesenlarged,dentalarcadeUshapedwithmolarsin
twoparallelrows(toothrowofmanishyperbolic)
basedprimarilyonDNAevidence,ithasbeentheorizedthatatapproximately56Magorillas,

PALEOLECTURE,PAGE141
chimpsandhominids(man'sfamily)divergedwhenclimatebecamecooler,drierandmore
seasonal(termedtheMessinianClimateCrisis)
However,recoveryofa67MayearoldskullwithahumanlikefacefromnorthernChad,
Africa(Sahelanthropustchadensis)mayindicatethatthedivergenceofhumansfromotherapes
occurredat8to10Ma
c1.TheHominidae
therearemanycompetingclassificationsofthisfamilybutthisisoneinterpretation:
c1a.Australopithecines
thefirst"humans"
Ardipithecusspeciesweretheearliestknownaustralopithecines,includingArdipithecus
kadabba(ca.5.7Ma?)andArdipithecusramidus(ca.4.54.3Ma)fromEthiopia,Africa;consist
ofgracile(lightlybuilt)australopithecineswithchimpsizedbrainsthatinhabitedwoodlands
Ardipithecuswasbipedal(asindicatedbythepelvisandlegstructure)butthebigtoeonthe
footwasdivergent(thefootcouldbeusedforgrasping),suggestingArdipithecusmayhave
nestedandfedintrees
Australopithecusspecies(ca.42Ma)weregracile(lightlybuilt)hominids;fullybipedal
(determinedbyhips,thighbonesandfossilfootprintsatLaetoli);veryapelikeinmostof
skeletonwithlongarmsandfingers;thebrainischimpsize(400500milliliters);moderateto
markedsexualdimorphism(maleslargerthanfemales);heightfrom1.0to1.5meters(3'3"to4'
11")andweightfrom30to70kilograms(66to154pounds);a"gracile"australopithecine
probablyleadtoHomo
Paranthropusspecies(2.61.2Ma)wererobust(heavilybuilt)australopithecines;relatively
longarms;height1.1to1.4meters(3'7"to4'7")andweight40to80kilograms(88to176
pounds);markedsexualdimorphism;prominentcrestsontopandbackofskull;verylong,
broad,flattishface;strongfacialbuttressing;verythickjaws;smallincisorsandcanines;large,
molarlikepremolars;verylargemolars;brainsize410to530milliliters

c1b.Homo
thegenuscontainingmodernman
c1b1.EarlyHomoSpecies
Homorudolfensis(ca.2.51.9Ma)andHomohabilis(ca.2.11.5Ma)weresimilarto
Australopithecusbutbrainsizeincreasedtoabout650750ml
TheOldowanCulture
firsttoolculture;althoughtheOldowanCulturehasconsideredtobea"pebbletool"culture,
theirprimaryuseappearstohavebeenaschoppers,scrapersandpounders;theOldowanCulture
wasprobablyduetoAustralopithecusgahri,Homorudolfensis,H.habilisandearlyH.ergaster
datesat2.51.5Ma;earlyhumansusedthesetoolsfor"expandingtheirniche"cutting,

PALEOLECTURE,PAGE142
crushing,digging,projectilesandcarrying;itappearsthatthehominidshadnopreconceived
shapeofthetoolduringmanufacture(i.e.,no"mentaltemplate")
earlyHomospeciesmayhavelivedinmultimaleandmultifemalegroups;malescompetedfor
accesstofemales
noevidenceofintentionalburials,gravegoods,art,etc.;noclearevidenceofarchitectural
features
c1b2.Homoergaster
approximately1.9to1.5MaineasternAfrica
maybeancestraltoallsubsequentHomospecies
theslenderbodied,longlegged"TurkanaBoy"skeletonisessentiallymodernandwitha
highlyefficientstridingstructure;adultsprobably1.8meterstall(6')ormore;brainsize800
1050ml
oldestH.ergastermadeOldowantools;atapproximately1.65MadevelopedAcheulian
industry(withlargehandheldstoneaxes);mayhavebeenfirsttousefireat1.7Ma(fire
provideswarmth,usedinhunting,protectionagainstpredators,removetoxinsfromfood)
c1b3.Homoerectus
Asiaticform[ca.1.5Mato225Ka]witharelativelylargebrain(8501150ml),flatskull,
largebrowridges,slopedforehead,nuchalcrestonbackofskull,almostnochin;probablydid
notgiverisetolaterHomospecies
c1b4.OriginofHomosapiens
probablyevolvedfromH.ergasterlikespecies
by500200KawithformsintermediatebetweenH.ergaster/"erectus"andH.sapiens
OriginTheoriesforHomosapiensinclude:
MultiRegionalHypothesisevolutionfromseveral"stocks"ofmigratedHomoergaster/
"erectus"(especiallyAfricaandeasternAsia)
OutofAfricaHypothesisevolutionfromasinglestockofH.ergaster/"erectus"thatlater
migrated(mostpopulartheory)andreplacedoldergroups
Homofloresiensis,atiny(adults42incheshigh!)islandspeciesfromIndonesia,issimilarto
Homoergaster;itmayhavelivedaslateas18,000yearsago(iftrue,thisgreatlychangesour
ideasofthediversityanddistributionofancienthominids)
c1b5.Homoneanderthalensis
earlypreNeandertalsat400Ka;Homoneanderthalensisat150Kato27Ka;mostlylivedin
EuropeandwesternAsia
oftenmassivebrowridges;largecheekbones;protrudingface;nochin;"bun"shapedskull;
largecranialcapacity(oftengreaterthanmodernman);short(1.5meters;5ft.)butverypowerful
probablynotancestraltoHomosapiens(withdistinctDNA)
handaxesdecline,flaketraditionbecomesdominant

PALEOLECTURE,PAGE143

MousterianTradition
usuallyattributedtoHomoneanderthalensis
strikeflakefromundersideofaprepared"tortoiseshell"coretocreatemanytooltypes;many
ofthesewereCompositeTools(artifactsmadefrommorethanonecomponent)
c1b6.Homosapienssapiens
HomosapienssapiensevolvedfromarchaicH.sapiensinAfricaandthenreplaced
neanderthalsinEurasia?
theremayhavebeenanearlydispersalofanatomicallymodernlookingHomosapiensfrom
Africaatabout100Ka;theremayhavebeenasubstantialbottleneckofpopulationafterthat,
withnumbersdroppingtoaslowas10,000individuals
HomosapienssapiensdevelopedtheUpperPaleolithictooltechnology(35to9Ka);often
typifiedby"punchstruck"bladeindustries(abladeisalongflake);thesewere"specialized"
huntergatherers(concentrateonafewresources)thatoftenhuntedherdanimals
ReligionBurialswithceremonialburialsandgravegoods
UpperPaleolithicArtfirstwidespreadproductionoftrueartwasbymodernHomosapiens(Ex.
=cavepaintings),probablywithareligioussignificance

D.OrderRodentia
rodentsandrabbitsaresistergroups;oftenplacedintheSuperoderGlires
rodentsincludesquirrels,rats,miceandguineapigs
approximately40%ofallknownmodernmammalianspecies(over2,000livingspecies);
approximately50familiesevolvedintheCenozoic(approximately1/4oftheseareextinct)
probablyevolvedfromanagalids(asdidthelagomorphs)
1.EcologyandDistribution
foundonallcontinentsexceptAntarcticaandinnearlyallhabitats;mostlyterrestrial,small
scamperingquadrupedswithclaws,longtailsandwhiskers
2.Osteology
a.Postcranialskeleton
hasnotchangedmuchfromtheprimitivecondition
b.Skullandteeth
aremuchmodifiedforgnawing;onepairofincisorteethinupperjawandonepairinlower
jaw;teethcurvedandcontinuallygrowing;incisorswithtwolayersofhardenamelpresentonly
onfrontface(edgebehindwearsawaymorerapidlyandtoothstayssharp);diastemaposteriorto
incisorteeth

PALEOLECTURE,PAGE144
usuallyfourgrindingteeth(P4M3)posteriortodiastema;manytoothpatternsdeveloped
massetermuscle(onethatclosesthejaw)enlargedandinsertsontoflangeofmandiblebelow
andbehindteeth;masseterattachmentpatternsareimportantinrodentclassification
E.OrderLagomorpha
includepikas,rabbitsandhares;latePaleocenetoRecent;probablyderivedfromanagalids
herbivores;feedongrassesintheplainsandonshrubsinrockytundraanddesertterrains
twopairsofpersistentlygrowingincisors(onepairbehindtheother)inupperjawandenamel
completelysurroundsthetooth;onlyonelayerofenamelonanteriorsurfaceofincisors(two
layersinrodents)
F.CarnivorousMammals
1."Creodonts"("OrderCreodonta")
probablypolyphyletic,butsomememberswereancestralofthe"truecarnivores"
dominantTertiarycarnivores;onallcontinentsexceptAustraliaandSouthAmerica;with
approximately50generaknownfromtwofamilies;carnassialsusuallyinvolvingM1+2/M2+3
(moderncarnivoreswithcarnassialsatP4/M1);shorterlimbs,unfusedwristbones,terminal
phalangeswithfissuredclaws,usuallynolossoftoesversustruecarnivores
includesmustelidandfelidlikeOxyaenidsandsabretoothcat,dogorhyaenalike
Hyaenodontids
2.CarnivoraVeraor"TrueCarnivores"(OrderCarnivora)
includestheorderofmoderncarnivores
wasrelativelyminorpartoffaunasinPaleoceneandearlyEocene;lateEocenewithcanid
(dog),viverrid(civet)andpossiblymustelid(weasel,stoat,mink,marten,skunk,badgerand
otter)lines
a.GeneralFeatures
usuallysimpledigestivesystemandconservativedentition(usually3/31/14/42/3)withupper
canineenlargedandlastupperpremolarandfirstlowermolarformcarnassials;oftenreduceor
loseclavicle("collarbone")andwristbonesfused[relatedtocursorial("running")locomotion]
typeofossificationofauditorybullaisimportantinclassification(includingelementswhich
makeupbullaandpresenceorabsenceofabonyseptumdividingthebulla)
b.Classification
havebeendividedintotheFissipedia(landdwellers)andPinnipedia(sealsandwalruses)but
doesnotreflectthetruephylogeneticrelationshipsoftheCarnivora
MostimportantgroupsoftheCarnivorainclude:
b1.Arctoidea(Canoids;Caniformia)
includetheCanidae,Amphicyonidae,Procyonidae,Ursidae,MustelidaeandtheMarine

PALEOLECTURE,PAGE145
Carnivores;internalcarotidremainsanimportantarteryandtheauditorybullaisnotclearly
dividedintotwochambers
believedtohavebeenderivedfromtheprimitivemiacinecarnivores
Canids(FamilyCanidae)includesthedogs,foxes,wolves,andjackals;usuallysavannah
cursorialcarnivores;goodcarnassialdentitionandretaincrushingteethbehindthese;longlimbs,
digitigradefeet(walkontoeswiththeheelsnottouchingtheground);mostlysocialanimals;
EoceneRecent
AmphicyonidsorBeardogs(FamilyAmphicyonidae)successfulbearlikecanoidstock;
EocenePliocene
Ursids(Bears)probablyevolvedfromcanidsduringtheMiocene;posteriorteethform
crushingsurfaceforomnivorousdiet;plantigradefeet(walkonthesolesofthefeet)
Procyonidsincludestheraccoons,kinkajou,coatisandtheirkin;small,arborealomnivorous
types;carnassialsloseshearingfunction;plantigradefeet
Mustelidsincludesskunks,weasels,badgers,wolverinesandotters;usuallysmallnorthern
temperateformswithshortstockylimbsandnolossoftoes;noseptumintheauditorybulla;
goodcarnassials;sometimesplacedwithinaseparatelineage(Musteloids)versusdogs,raccoons
andbears;OligoceneRecent

b2.AquaticFamilies(the"Pinnipedia")
enaliarctids,desmatophocids,otariids,odobenidsandphocidsareusuallygroupedinthe
SuborderPinnipediabutstudyofearregionindicatesthatphocoids(seals)andotaroids(the
otherfourfamilies)haddiphyletic(separate)originsfromterrestrialcarnivores
Odobenidswalrusandkin;simplifiedcheekteethwithonlyasingleroot;derivedtypeswith
tusks(forpryingclams);MioceneRecent
Otariidssealions,earedsealsandfurseals;hindlegscapableofpositioninganteriorlyfor
locomotiononland;cheekteethwithsinglecusps;MioceneRecent
Phocidsincludethe"earless"seals;hindlegsalwaysposteriorlypositioned;cheekteeth
doublerootedandwithaccessorycusps;fossilrecordpoor,butmaybemorekinto"musteloids"
than"canoids";MioceneRecent

b3.Aeluroids(Feloids)
includesviverrids,felidsandhyaenids;mainbranchofinternalcarotidarteryreducedorlost;
externalcarotidwithcountercurrentexchangerinthevicinityoftheorbitforcoolingblood
enteringthebrain
Viverroidsincludeviverrids(civets),herpestids(mongooses),hyaenids(hyaenas)andfelids
(cats);haveaseptumintheauditorybulla
Felidscats;mostaresmalltomediumsizedandliveonadietofrodents;rangeworldwide
exceptforAustralia;digitigrade(digitsbearweight)withaflexibleskeleton,sharpretractile
claws;carnassialsextremelywelldeveloped;include"truecats"and"sabretooths"(arenot
monophyleticgroups)

PALEOLECTURE,PAGE146
G.EarlyRootersandBrowsers
earliestherbivoresweremostlyPaleoceneandEocene;fromrabbittoelephantsized;probably
mostwererootersorfeedersontubers(withclawedfeet,largecaninesandbroad,lowcrowned
cheekteeth;typicallycompletedentitionwithnodiastema)
includesthepiglikeTaeniodonts,semiaquaticPantodonts,massiveherbivorousDinocerata
(the"Uintatheres"),thelargeclawedfootedTillodonts,therhinosizedEmbrithopods,andthe
"Condylarths"(ancestraltoallotherherbivorousgroups;includesthe
arctocyonids,hyopsodontids,andphenacodontids)
H.SouthAmericanherbivores
1.GeologicandPaleontologicHistoryofSouthAmerica
fromPaleoceneuntilPliocenewasan"islandcontinent"
SouthAmericanmammalianevolutioncanbedividedintothreephases:
a.LateEocenewithmarsupials,edentatesandcondylarths
b.EarlyOligocenethroughlateMiocenewithmarsupialsandedentatesradiatingintomany
adaptivenichesandcondylarthsdiversifyintosixordersandabout25families;alsowith
primatesandrodentsreachingSouthAmericaduringtheearlyOligoceneby"rafting"fromNorth
AmericaorWestAfrica
c.GreatFaunalInterchangeduringPlioPleistocenewithfaunasmigratingnorthandsouth
acrossCentralAmerica(SouthAmericareceivedmastodonts,horses,tapirs,peccaries,camels,
deer,shrews,hares,squirrels,mice,dogs,bears,raccoons,ottersandcats;alloftheSouth
Americantaxabecameextinctandthesearethecrittersdiscussedbelow!)
SouthAmericanherbivoresprobablyderivedfromdidolodontcondylarths
2.OrderLitopterna
primitivelywithcompletedentalseries;nodiastema;uppermolarswithlowcusps;trigonidand
talonidonlowermolarsnearlythesameheight
includestherabbitlikeAdianthids,thehorselikePrototheriids,andtheweirdcamellike,
trunkbearingMacraucheniids
3.OrderNotoungulata
largestorderofSouthAmericanherbivores(4suborders,1314familiesandover100genera);
latePaleocenetoPleistocene;alsoknownfromlatePaleoceneearlyEoceneofWyoming,China
andMongolia
notoungulateswithshortskulls,flattenedabovewithabroadbraincase;earregiondevelops
twochambers;earlytypeswith44lowcrownedteeth;latertypesformdiastemawithhigh
crownedmolarswithpishapedlophs;feetaremesaxonic(thethirddigitformstheaxis)with
reductionofthelateraldigits;mostfeetwerehoofed
mostimportantgroupsaretherodentlikeTypotheres;thehorse,chalicothereorrhinolike
"bowtoothed"Toxodonts;therhinolikesemiaphibious(?)Astrapotheres;andtheshortlimbed
pigortapirlikePyrotheres
I.The"Edentates"

PALEOLECTURE,PAGE147
Apolyphyleticgroupincludingthepangolins,armadillos,anteaters,slothsandtheextinct
palaeanodonts(butseebelowforprobablephylogeneticrelationships)
reductionorlossofdentition(ifretainedteethlackenamel);mostwithlargeclaws(especially
onforelimbs)usedfordigging;dietoftenconsistsofants,termitesandsmallinsects;living
memberswithmuscular,gizzardlikeportionofstomachforcrushingprey
1.Xenarthrans(OrderXenarthra)
includetreesloths,anteatersandarmadillosandtheextinctglyptodontsandgroundsloths
withaccessoryareaofattachmentonpelvicgirdle;mostwithextravertebralarticulations(=
xenarthra;probablyinitiallyfordigging;latersupportedcarapace/shellinsomegroups),scapula
(shoulderblade)withtwoparallelspines;feetusuallybearlargeclaws,somegenerawith
ossicles(bonyplates)intheskinandotherswithabonyshieldoverthebody;modern
xenarthranswithvariablebodytemperaturewhichislowerthaninotherplacentals
a.InfraorderLoricata(Cingulata)
includestheDasypodidae(armadillos;probablyancestraltootherxenarthrans;Paleocene
Recent)andtheGlyptodontidae(MiddleEocene(?)MiocenePleistoceneage;oftenhuge,
armadillolike)
b.InfraorderPilosa
includestheBradypodidae(treesloths)andtheGroundSloths(OligocenePleistocene;from
catsizedtooversixmeterslong;includestheMylodontoideaandMegalonychoidea)andthe
Myrmecophagidae[trueanteaters;Eocene(fromGermany!)toRecent]
2.Other"Edentates"
includePholidota(pangolins;with"pineconelike"scales;EoceneRecent;moleculardata
indicatesthattheseareactuallyasistergroupofcarnivores!);Tubulidentata(aardvarks;Miocene
Recent;moleculardataindicateskinshiptotenrecs,goldenmolesandelephantshrews)and
Palaeanodonta(includeLowerTertiarymolelikeandslothlikeforms)
J.OrderProboscidea
includetheelephants;EoceneRecent
1.RelationshipsoftheAfrotheria(Afrotheres)
recentcladisticstudiessuggesttheexistenceoftwocladesofafrotheres;onethatincludesthe
aardvarks(Tubulidentata),tenrecs(Tenrecoidea)andelephantshrews(Macroscelidea);and
anotherthatincludesthe"Tethytheres"[elephants(Proboscidea),hyraxes(Hyracoidea)and
dugongs/manatees(Sirenia)]
2.GeneralCharacteristicsoftheProboscidea
overallevolutionarytrendsincludeincreaseinsize;cheekteethtrendtowardscyclicsuccession
andenlargedcrownvolume;enlargementofsecondincisorteethastusks(firstforfoodgathering
andlaterfordisplay);developmentofproboscis(nasalopeningisposterior)withincreased
shorteningoftheneck

PALEOLECTURE,PAGE148

3.EvolutionaryHistoryandClassificationofElephants
maybederivedfromphenacodontidcondylarths;GeologicRange=EoceneRecent
mostimportantgroupsaretheDeinotheroidea(withdownturnedandbackwardlycurvedtusks)
andtheEuelephantoidea[includetheMammutidae(mastodonts),Gomphotheridae("shovel
tuskers")andtheElephantidae(elephantsandmammoths)]
K.OrderSirenia
manateesanddugongs
totallyaquatic,huge(upto8meterslong)shapelessanimalswithalargetail(the"swimfin"),
smallfrontflipper,bluntmouthwiththickoverhanginglips;skeletaladaptationsforaquaticlife
includepelvicgirdlereducedtoavestigeandthick,denseribs;EoceneRecent
L.OrderDesmostylia
ponysizedaquaticanimalswithshort,stoutlimbsandshortshovelliketuskedmouth;each
cheektoothmadeofclusterofstoutdentinetubules;UpperOligoceneMiocene
M.OrderAcreodi(Mesonychids)
difficulttoclassify(oftenplacedclosetothecetaceansandmayhavebeenancestraltowhales)
descendedfromarctocyonid"condylarths";mayhavebeenhyaenidlikebonecrushing
scavengers;includelargestknownlandcarnivores/scavengers
N.OrderCetacea
includethewhales
mesonychids,whales,artiodactyls(cattle,antelopes,pigs,etc.)andperissodactyls(horses,
rhinosandtapirs)arerelatedandoftenplacedwithinthe"SuperorderCetartiodactyla"
1.GeneralCharacteristicsofCetaceans
specializedforaquaticlifewithstreamlinedbody;tailformshorizontalflukeforpropulsion;
hindlimbabsent;forelimbformsshortflipperforsteering;thickfatlayer;nosenseofsmelland
visionispoorbutsenseoftouchandhearingverywelldeveloped;brainslargeandcomplex;
primarilycarnivores(feedonsquid,fishorplankton)
bonesfilledwithoil(forflotationandenergyreserve);snoutelongate,restofskullis
telescoped(maxillaemaytouchbonesatbackofbraincase);nosacrum,typicallynohindlimb
andonlyremnantofpelvicbone;pectoralgirdlelacksaclavicle,humerusshortandpaddlehas
fourorfivedigits,eachwithmanyphalanges
2.ClassificationofWhales
includesthreesuborderstheArcheoceti(earliestwhales;derivedfrommesonychidsor
artiodactyls;EoceneOligocene,Miocene(?);includesthelongsnouted,toothed"zeuglodonts");
Odontoceti(toothedwhalesincludingdolphins,porpoises,spermandkillerwhales);Mysticeti
(includeplanktonstrainingbaleenwhales;largestanimalsthateverlived)
O.OrderPerissodactyla

PALEOLECTURE,PAGE149
"oddtoed"ungulatesincludingtapirs,rhinoceroses,horses,brontotheresandchalicotheres
derivedfromphenacodontidcondylarths
firstappearintheEocene(alsopeakedintheEocene);goodfossilrecordinNorthAmericaand
EurasiaandlatermembersfoundinAfricaandSouthAmerica
axisofweightbearingpassesthroughthemiddleorthirddigit(mesaxonic);mostmembersare
threetoedbutlaterhorseseliminatedthelateraldigitstobecomeonetoed
astragalus(anklebone)withasingle"pulley";molarteethwithalophodontpattern(with
enamelloops)
1.SuborderCeratomorpha
includestapirsandrhinoceroses(developcrosslophsontheteeth);includelargestland
mammalsknown(Inthricotherineswererhinosupto5.4matshoulder)
2.SuborderAncylopoda
a.SuperfamilyChalicotheroidea
chalicotheres;EocenePleistoceneofNorthAmerica,EurasiaandAfrica;Moropus(Miocene,
NorthAmerica)washorsesizedclawedbipedalbrowser
3.SuborderHippomorphaincludesthehorsesandbrontotheres
a.SuperfamilyEquoidea
horses;EoceneRecent
evolutionarytrendsincludeincreaseinsizeandheight,increasedcomplexityofenamelpattern
oncheekteeth,elongationoflegs,reductionoftoestoone
b.SuperfamilyBrontotheroidea
includethetitanotheres(brontotheres);mediumtoverylargesizedherbivoresoftheearly
TertiaryofNorthAmericaandeasternAsia;withwshapedenamelridges(ectoloph)onthe
uppermolars
P.OrderArtiodactyla
"eventoed"ungulates;includespigs,camels,giraffes,deer,antelope,goats,sheep,cattleand
otherextinctandmoderngroups;with79livinggenera;27familiesfromCenozoic(10survive);
derivedfromthearctocyonidcondylarths
1.Morphology
footaxisbetweenthethirdandfourthdigits(paraxonic);digitsequalinsize;metapodialsoften
fusetoformcannonbone;astragalus(anklebone)formsdoublepulleystructure
primitivestocks(Ex.=pigs)withcompletedentitionandoftenwithenlargedcaninetusks;later
stockswithupperincisorsreducedorlost,withadiastema,premolarsbecomepartially
molarized(i.e.molarlike)
earlystocks(palaeodontsandsuiformes)withbrachydont(lowcrowned)andbunodont(with
low,roundedcusps)molars;allotherswithselenodontdentition(cuspscrescentorhalfmoon

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shaped)
earlystockswithswinelike,shortlimbswithfusionofthebonesandlittlereducedlateraltoes
complexstomachsfordigestingvegetation(pigsandhipposwith2to3chambers;camelsand
llamaswith3chambersandruminate;cattleanddeerwithfourchamberedstomachsand
ruminate)
withorigininthePaleocene;firstradiationinearlyEocenegaverisetomanypiglikestocks
(forestandwoodlandrootersandbrowsers);secondradiationinlateEoceneandearlyOligocene
gaverisetoearlyruminants;inMioceneruminantsdiversifiedtoexploitthesavannahsand
grasslandsandremainthedominantherbivorestheretoday
2.Classification
includethe"Palaeodonts"(Eoceneprimitiveartiodactyls;probablypolyphyletic),Suina
[Suiformes;includethepiglikeentelodonts,peccaries(javelinas),truepigs,andhippos],
Tylopoda[cainotheres,xiphodonts,merycoidodonts(=oreodonts;themostsuccessfulextinct
tylopodgroup;pigorsheeplike),agriochoerids,camelids(camelsfirstevolvedinNorth
America;includemanydiverseextincttypes)andprotoceratids(deerlike;oftenwithweird
"nose"horns)],andtheRuminantia[includetheTraguloids(chevrotains,muskdeer)andthe
Pecora(includesthehigherruminants;cattle,deer,giraffesandantelope;withdevelopmentof
variouspaired"horns";withossiconesingiraffes,antlersindeerandhornsincattle,antelopes,
goats,sheep,etc.;oftenwithfusedlimbbones)]