VertebrateOrigins

AStudyonFossils,AdvancedFeaturesandVertebrateHistory

By
KarlJ.Stiger

I.VertebralColumnandBraincase

II.Jaws
1.Placodermi
A.Dunkleosteus
B.Eastmanosteus
C.Botheriolepis
D.Stensioella&Gemuendina

2.Chondrichthyes
E.Cladoselache&Symmorium
F.Stethacanthus
G.Hybodus,Spathobatis,Heliobatis
H.Scyliorhinus
3.Actinopterygians
I.Cladocyclus
J.Calamopleurus
K.Eustenopteron
III.4limbs
L.Eryops
M.Buettneria
N.Diplocaulus
O.Pantylus
P.Phlegthontia

IV.EndnotesandConclusions

Introduction:
Vertebrateanimalsareperhapsthelargestandmostthewidelystudiedontheplanetearth.
Theentiretyofthesubphylumidentifies64,000totalspeciesdividedupinto5classes.Each
vertebratevariesinformandfunction.Butthereareadvancedfeaturesthatallunitethem,as
theyareinheritedfromacommonancestor.
VERTEBRALCOLUMN
&BRAINCASE
Thevertebralcolumnisthemostimportantpartofthebody,beingthatitstiffensthebody
andprovidessupportforthemuscles.However,itsoriginslieintheevolutionofasystemof
cartilaginousorbonyrods(whichmakeupthenotochord),andinturndeveloptheskeleton.Early
jawlessfishhadawhollycartilaginousskeletonwithapersistentnotochordthatmay
havebeenanassetintheevasionofpredators,responsibleforquickescapes.Asasubphylum,
vertebratesaresplitintofiveclassesofbackbonedanimals,allhavingabackboneanda
braincase.Inadditiontosupportforthemuscles,thebraincaseorcraniumprotectsthevital
senseorgansincludingthebrain.Theprimitivejawlessancestorsi.e.
Sacabambaspis
,
Drep
anaspis
and
Arandaspis
allhadaheadprotectedbybonyplatescoveringthedorsalandventral
sectionsofthebody.Mostimportantly,theseanimalslackedjawsandprobablysuckedparticles
fromthewater.Diagnosisthevertebralcolumnandbraincaseformsaninternalskeleton
alongsidethebodyaxis.Thevertebralcolumnformsasegmentedanchorformusclesalongthe
bodyaswellas,protectionforthenervechord.Thereisalsoasegmentedregionwithabrain
casethatsurroundsthebrain,sensoryorgansandcranialnerves.

Thesegmentedbodymusclesareanchoredtoadjacentpartsofthevertebralcolumnandthismay
causethemtocontractandrelax.Theostracodermswereextinctjawlessvertebrateswith
hardenedarmorshelllikeplatescalcifiedbybonesintheskin.And,maybethisiswhythese
animalshavebeenclassifiedastheostracoderms,orshellskin.Theonlysurvivingvertebrates
withoutjawsarethelampreyswhichinhabitfreshwaterandthehagfishaquaticenvironments.

JAWS
1.ThePlacodermi
Jawswereapinnacleevolutionaryinnovationfoundinbothinvertebratesandvertebrates,
usedforpreycapture(eitheraspincersorhinges),thatwereencasedinboneorcartilage.These
structuresallowedanimalstobecomepredatorsandotherstobecomeprey.Placodermswere
armoredvertebrateswithjawsfashionedintotoothplatesorshears.SPECIMENS
Dunkleosteus
terrelli
wasoneofthelargestmembersofthearthrodirajointneckedfish,whichlivedinNorth
America360millionyearsagoinpelagicenvironments.Itwasanektonicpredatorwhichranged
thevastpelagicwaterwaysinsearchofpreyandfedonsmallervertebratesincludingprimitive
sharks.Asitsanatomyisconcerned,theskullof
D
.
terrelli
(FF3

)measures4ft.3in.(1.3m
long),howeveritsbody
lengthmeasured16to20feetanditsjaws,includingbothmaxillaeandmandibullaewerefitted
withserratedtoothplates.Similarly,itsclosestrelative
Eastmanosteus

calliaspis
,wasasmallerplacodermalsobelongingtothearthrodira.Unlike
D
.
terrelli
,the
jawsof
E
.
calliaspis
sportedwedgeshapedtoothplateswithenamellikepoints.Thespecimen
Iexaminedatthehallofvertebrateoriginsmeasured7in.(18.3cm)andthematerialalso
includedaheadshieldandtrunkarmor.HABITATS
Eastmanosteus
livedinwhatisnowthe
GreatBarrierReefinNorthwesternAustraliainalagoonreef.
Eastmanosteus
waspiscivorusand
mayhavefedonsmallfishandarthropods.Placodermsdidnotjustinhabitopenwateror
lagoonseither,infactsomesmallergenera(i.e.
Stensioella
&
Gemuendina
)werebenthic
meaningthatboththeseanimalslivedatorneartheoceanfloor.
Stensioellahientzi
asIsawin
onespecimen,hadfewerbonyplatesonitscephalicregion.However,itsbodythoughwas
coveredintinydenticlescales,similartothoseobservedinsharksandrays.Theentirelengthof
thespecimenmeasured10in.(26.9cm)long.Theanimalsdietmayhaveincludedarthropods
&shellfish,likemanymodernchimaerasandskates(relativesofsharks)anditprobablyhada
similarlifestyle.SPECIMENSAnothergroup,theRhenanids,werealsomarineplacoderms
andlivedmuchliketodaysraysandskates.In
Gemuendina
,(FF4132)boththefinsandbody
wereflattenedandthetailislongandtapered.Butunlike
Stensioella
,thedorsalfinisabsent.
Consequently,allthesefeaturescanbeseeninraysandskates,butthesimilaritiesthough,
representauniqueexampleofconvergencebetweengroupsofunrelatedanimals.

AntiarchswerefreshwaterplacodermsthatlivedduringthelateDevonianperiod360million
yearsagoandareidentifiedbysmallerbodies,armoredheadsandthoracicregions.Features
oftheiranatomyincludeajointbetweenheadandbodyarmor,appendagessituatedbehind
thehead,andflattenedtoothplates.Mostplacodermfossilsconsistgenerallyofbonyplates,
butmanycompletespecimenswerefoundtogether,suggestingthattheylivedinsmallschools.
SPECIMENS
Botheriolepiscanadensis
:materialincludesarmoredheadandthoracicregion.
B.canadensis
dietsuggestsitmayhavebeenascavengertheentirelengthofbodymeasured
10in.(1.24cm).HABITATThedistributionofthisgenusisworldwideandprolificinthe
freshwaterenvironmentsandverycommoninWesternN.America.

2.Chondrichthyes

Sharksproduceasmanyas16to30,000teethinanentirelifetime.Fossilcollectorscanfind
isolatedexamplesofteethbelongingtovariousgeneraandspecieseasily.Sharkteethdifferin
sizeandshapeaccordingtothespeciesage,andsexofindividualanimals.Andtherefore,the
majordifferencesintheteethincludetheirsizeandshapethedegreeofcurvatureinadditionto
thepresenceortheabsenceofcuspsintheserratedmargin.Plusthereisanumberand
arrangementofcuspspresent.Furthermore,thejawsandteethshowhowsharkteethare
arranged.Theteethareattachedtothegumtissuebystrongfibersandthejawsareattachedto
theskullbyligaments.Newteethmoveintopositionbymigrationofthegumtissuetowhich
theyareattached.Thebrokenandoldteethareshedandreplacedasfrequentlyasevery10days,
dependingonthespecies.TheprimitivesharksknownasCladoselachiansorcladodonts,because
oftheshapeoftheirtoothstructure,possessseveraladvancedfeaturespresentinmodern
Chondrichthyesincludingprismaticcalcificationoftheendoskeleton.SPECIMENS

Cladoselachefyleri
(FF239)

wasasmallsharkmeasuring76.2cm(2ft.6in)long.Fossilsofthis
animalshowpresenceofthesmalltoothlikedenticlescalesinadditiontoanotochordand

3
acraniamaxillae.HABITATDuringthelateDevonian,
Cladoselache
wasrestrictedtoinferior
seawaysthatcoveredmostofN.Americawereitfedadietofsmallfish.Inonespecimen,I
observedevidenceofinternalorgansbutnoclasperspresent,whichsuggesteditwasafemale.
OtherearlyexamplesofPaleozoicgenerabelongingtotheclassChondrichthyesalsoinclude,
Symmoriumreniforme
(FF19502)and
Stethacanthusthomasi
,mostoftheseacquiredthesame
patternoftraitsincludingcladodontteeth.However,S.thomasiistheonlygenuswhosefeatures
representananomalyinmalesaspinebrushcomplexisobservedanditwastheorizedthatthe
spinebrushservedasasexualfeatureusedasanattractant.Consequently,thefirstspecimenwas
describedin1889byJohnNewberry.AttheAmericanMuseumofNaturalHistoryshallof
vertebrateorigins,thereisonespecimenof
S
.
thomasi
FF565ondisplaythat,showstoothlike
scalessupportedbyamodifiedfin.Inthe1970'sand80'scompleteskeletonshavebeen
discoveredwhichrevealaclosesimilaritywith
Symmorium
andothercladodontsharks.
HABITAT
Symmoriumreniforme
livedinwhatisnowKansasandIndianainshallowseas
duringtheCarboniferousperiod.SPECIMENSAtthetime,sharklikespeciesoftheclass
Chondrichthyeswhichwereabundant,andanatomicallymorediversethantoday.Atthehallof
vertebrateorigins,thespecimenofSreniformeshowsthecranium,finsandgill
arches.Furthermore,inthespecimenFF19502thereisanabsenceoftheelongatedsex
organsespeciallythoseseeninmalesandthepelvicgirdlebegintobisectforminganup
wardsV.Hybodonts,XenacanthsandNeoselachianshaveallbeenclassifiedaselasmobranchs
basedonsharedfeaturesofthegillskeleton.SPECIMENS
Hybodusfraasi
FF7191is
anatomicallydistinctiveespeciallyinthestructureofthebraincase,teethandjaws.1.Thejaws:
highlyamphistylic,meaningthatthemaxilla(upperjaw)ispartlyfreeofthebraincaseandisthen
bracedbythearchofthehyomadibularcartilage.2.Thebraincase:Isformedentirelyofcartilage
withaclevershapedcartilageelementalsopresent,fromwhichthelowerjawattached.3.Teeth:
heterodontconsistingofpointedinfrontforgraspingandbluntinbackforcrushing.Themales
havecephalicspinesinadditiontoheavydorsalspineswhichservedassecondarysexual
features.Thisspecimenof
H.fraasi
FF7191measuresalengthof6ft.7in(2.0m).Furthermore,
anotherdistinctfeaturepresentinhybodontsharkswasawelldevelopedribcagethepresence
ofwhichIcannotsayorhaveareasonfor.Honestly,itisinterestingtometothinkthatan
animalgroupcanevolveanatomythatisbynaturalstandards,outofplace.Howeverinancestral
organismstherearecharactertraitsthatevolveasarepresentationoftheirlineage,butthenis
lostinothers:hybodontsharkshavearibcage.Butintheneoselachians,theribcageislost
completely.HABITAThybodontsharksweremostlycommonduringtheJurassicofEuropeand
N.America,inAquaticandfreshwaterdeposits,butsomegenera(i.e.
Tribodus
),arecommonin
lagoonalenvironmentswherethewaterisshallowsurroundedbyreefs,whichoccurinmixed
sandorgravelcoastlines.Neoselachiansincludemodernsharksandrays.Alloftheseanimals
areunitedbyskeletalfeaturesincludingthestructureofthevertebrae,whicharestrongly
calcifiedtoformbonelikediscsthatconstrictthenotochord.

4
Thevertebralcalcificationpatternhadevolvedabout200millionyearsago,andmayhave
provided
astrongerbackboneforswimming.Someprimitivefamiliesofsharksandraysareknownfrom
theJurassic,whilemorespecializedgenera(i.e.BullsharksandEaglerays)haveamuchshorter
fossilrecordextendingbacktotheearlyTertiary.SPECIMENSEarlyrelativesofthemodern
angelshark,includetheJurassicsqualean
Pseudorhinaalfera
.Likerays,someof
P.aliferas
synapomorphiesinclude1.AnExpandedregionofthevertebraethatconnectstoskull2.An
Absentanalfin.The
angelsharkisarelativeofrays,however,itlacksmostoftheotherfeatureswhichdescriberays.
Neoselachiansynapomorphiesincludemainlyfeaturesofthejawstructureinsharksandrays,
the
jawsareconnectedwiththecraniumbythehyomandibulararch.Thismodeofsuspensionallows
theseanimalstoprotrudetheirjawsforward,inordertosnatchpreywithasinglemovement.
Hence,thejawsaredescribedasfreefloatinginsidetheskull.SPECIMENSTheearliestrays
resemblerhinobatoids(banjorays),becausetheyallasaresult,retainmanysharklikefeatures
includinglongbodyandheterocercaltail.Thelengthofthe
S.bugesiacus(FF7494)
specimen
measures5ft.,3in.(1.6m)anditssynapomorphicfeaturesinclude1.Frontfinexpandstomeet
snout2.Pairedclaspers.HABITATS
S
.
bugesiacus
livedamongspongesandcoralreefswhere
itfedonmollusksandsmallfish.SPECIMENS
Heliobatisradians
isafossilstingrayfrom
depositsinWyomingandUtah.HABITATSItlivedinfreshwaterlakesduringtheEocene
epoch55millionyearsago.ThespecimenIobservedwasafemale(asnotedonlybyabsenceof
claspersinherpelvicregion),shewassurroundedbyseveralofherpups.Thisindividualseems
tohaveundergoneastressinducedbirth,shortlybeforeandafter,thepupsdied.SPECIMENS
DogfishsharksfirstappearedduringtheJurassicperiodabout200millionyearsago.Initially,
thecommonnomenclatureforthisanimalhasledtoconfusionregardingitsgenus.
Consequently,thenameoftheAMNHspecimen(labeledcatalogue13983)measuresalength
of9in(22.9cm)andisfrom85millionyearlatecretaceousdepositsinLebanon
Scyliorhinus
elongatus
means:thesharkwithanelongatednose.TheSynapomorphictraitsin
S.elongatus
aswellasothermodernsharksandraysinclude1.Calcifiedvertebrae2.Spacebetweenhyoid
andgillarch3.Pelvicgirdleformingasolidtransversebar.Tomyknowledge,theonlyshark
knownbythatgenusisthecatshark.HABITATSasalivinganimal,
S
.
elongatus
hada
piscivorusdietanditfedonsmallfishandinvertebrates,whichimpliesthatitprobably
inhabitedshallowseas.

Chimaerasarerelativesofsharks,sometimescalledrabbitfish,liveindeepwaterandfeed
onmollusks.Thereare35speciesdividedinto3ordersand3families.SPECIMENSThe
toothplatesin
Edaphodonmirificus(
representedbyFF2223andFF2227)aremore
specializedthanthoseofrelatedgenera(i.e.
Coccliodus
).Traitsinclude:1.teethwithhard
areascalledtritorsthatareheldtogetherbydentinethatarealignedacrossthe

anim
als
jaw.2.Theautostylicjawsuspensionincludesthecraniamaxilla.SPECIMENS
Ischyodusativus
Thisspecimen:(FF7189andFF7190)representstheonlyholotype
speciesandwerethefirstcompletefossilchimaeroid,althoughitsbodyimpressionsare
faint,thespecimenIobservedatAMNHmeasures2ft.9in(83.8cm)long.InFF7189,
theheadisequippedwithsensorycanalsinadditiontodorsalspinesandapairmodified
claspers(foundonthemalesonly).Furthermore,theheadandjawsarepresentalongwith
thetoothplatesaswell.Itsdietmayhaveincludedsmallfishandmollusks.

Actinopterygians=Teleostei

Halecomorphs:
Calamopleuruscylindricus
SPECIMENSIn
Amia
(thebowfin),themandible
articulateswiththemaxillaandconnectswithasmallbonecalledthesymplecticinthenormal
jaw
joint.InFF11816,thespecimenIonceobservedinarecreationofafoodpyramidshowing
interrelationshipsamongpredatorsandpreyofthemidcretaceous,thestrongjawsarepresentin
additiontothepositionofvertebralcolumnsituatedbeneaththeanimalsscales.Furthermore,
C
.
cylindricus
measures3ftlong(91.5cm).

Ichthyodectids
:
HABITATS
Cladocyclus
representsan
extinctgroupthatoncewereverysuccessful.Thisgenus,differsfromothermembersofthe
groupbecauseofitsabilitytoliveinbothfreshwaterandsalineenvironmentsduringthemid
cretaceousofCeara,Brazil.Inonespecimen(FF2983),Inotedasimilaritybetweenboth
Calamopleurus
and
Cladocyclus
respectively.Andinbothspecimensstrongjawslinedwith
sharpneedleliketeetharepresentwhichfurtherindicatesthattheywerepredators,identifiedby
thesmaller
Rhacolepisbrama
insideitsstomach.

ConsequentlyIchthyodectidssuchas
Xiphactinusaudax
undoubtedlyhadastrictlycarnivorousdiet,duetoevidenceofitslastmeal
whichturnedouttobeashrimpinitsstomachcontents.

Wesometimesobtainpatternsor
glimpsesofbehaviorinfossils,butrarelydowehavetheopportunitytoexamineextinctanimals
indetail.Theremainsofstomachcontentshavebennfoundinmanyfossilfish,butby
identifyingtheremainsthemselves,itspossibletocarefullyreconstructanintactfossil
communityinitspurestform.
Fig.1basiccladogramshowingrelationshipsandfeaturesamongvertebrates

Eustenopteronfoordi
,Coelacanths&Lungfish
Evolutionaryrelationships
Thereareprofounddifferencesintheratesofevolutionarychangebetweencoelacanthsandearly
tetrapods,thatarefoundpresumablyinthetracesoffossilevidenceleftbyancestralkin,that
wererelatedtolobefinnedfish,namelytheosteolepiformes.
Eustenopteronfoordi
forexample,
sharesfeaturesincommmanyonwithtetrapodsincludingthearrangementofthenasal
passagesthatopenedtowardtheroofofthemouthandthereforeitisconsideredanancestorof
limbedvertebratesincludingus.
Eustenopteron
sfossilspecimenalonemeasures1ft.7in(48.3
cm)long.
Scaumenacia
,wasalateDevonianlungfishwhichhadjawslinedwith,continuously
growingtoothplatesadaptedforcrushingshellfish.Itlivedinbrackishwatersandsharedits
territorywithotherfish.Likeotherlungfish(i.e.Neoceratodus)ithasapairofanal(belly)finsin

additiontoahumeruswhichattachesitselftotheshouldergirdlethesearebonesthatsupport
appendages(armsandhands).SPECIMENSFF3338
Sauripterustaylori,
isanotherextinct
relativeoftetrapodssuchasfrogsandevenhumanswhichisidentifiedbyscalesinamatrix.I
madeanoteoftheextensivebasalbonesin
S.taylori
,asapomorphiccharacters,because
althoughithastiestotetrapods,itisstillarepresentativeofbonyfish.
Dipterus
wasa
coelacanththatlivedinlakesandfreshwaterstreamssurroundingtheriftthatlayneartheeastern
marginofNorthAmericaduringtheJurassicperiod.Likeothercoelacanths,itsharesclose
relationshipswithothercoelacanthsinAfricaandAustraliadueto,apatternwithaunique
presenceofahumerusthatfitsintotheroundedheadoftheshouldergirdle.

4LIMBS
Oneofthegreatesteventsinthehistoryofvertebrateswastheevolutionoflimbs.The4
limbsoftetrapodsallhaveincommon,aninternalbodystructure,whichisdividedinto
threesections:I.asinglebonethatarticulatesthebodyandarmscreatinganaxialbilateral
foundationofthelimbs.II.Next,therearetwobones(tibia+fibulaorradius+ulna)that
arelighterandcombinesupportwithagreaterdegreeofmaneuverability.III.Thesection
farfromthebodycontainsmanysmallerbonesincludingfingers,toesandanklesthatseem
toprovideflexibilityforlocomotionandfoodgathering.

Diplocaulusmagnicornis

Skull:10in(25.4cm)
Diagnosis:wideskullwithlargebonyprojections.SPECIMENS:
Diplocaulusmagnicornis
(FF
4466)anectridean,plusallofthenectrideanshave,asetofhornsorprojectionsatthebaseof
the
skull.In
D.magnicornis
,thebonyprojectionsformaboomerangshapeskullalthoughits
functionisunknown,itmayberelatedtothewaytheanimalswam.HABITATSthisanimal
livedmostlyinpondsandstreams.InmostofthespecimensIobserved,Inoticedvariationsin
thegrowthstages,asdemonstratedbydifferencesintheirsizes:Verysmallskeletonswith
unfusedskullsareindicativeofthejuvenileswhilethefusedskullsareindicativeofmature
adults.

Phlegthontialinearis
Body:ft(45.7cm)
Diagnosis:Asharplypointedskullwithsmallteeth.SPECIMENS:Phlegthontiaisthebest
knownmemberoftheaistopodsagroupknownfortheirlimblessandsnakelikebodies.They
werecompletelylimblessrepresentativeswithintheleposponspondyls(agroupthatalsoincludes
nectrideans)andwereprobablyrelatedtothem,asaresult.Oneoftheaspectsofearlytetrapod
evolutionisthatsomanyoftetrapodsreducedtheirnewlyacquiredlimbscompletely.Butwhy
theydid,isobjectivelyandhotlydebatedamongpaleontologists.
Pantyluscordatus

Skull:3in(7.6cm)
Diagnosis:Asolidlyroofedskullwithopenings.SPECIMENS:Themicrosaursarethemost
diversegroupwithinthelepospondyls.Whiletheirskullsaresimilarincompositionandform,
theirbodieshowevervaryfromelongatedandlimbless(asinP.linearis)toshortandlimbed(as
inP.cordatus).Pantylusisanexampleoftheshortbodiedextremehavingalmostnotailanda
verylargeskull.Microsaurskullshavesimilaritiestothoseofmostamniotes,
andithasbeensuggestedthatthemicrosaursmayberelatedtoamniotes.
Eryopsmegacephalus
Diagnosis:AlargeheadedtemnospondylSPECIMENS:(FF4178).Oneofthebestknownland
dwellers,Eryopshaswelldevelopedlimbsandappearstobemostterrestrialofthetetrapods.
Dissorophus
Dissorophusisanarmoredlissamphibian,andlikesometemnospondylsitmayhavebeen
terrestrial,asdemonstratedbythearmoredplatesonitsbodywhichmayhavebeenusedas
defenseagainstpredators.

Fig.2

Skullof
Buettneria
showing1.Palatal2.Orbits3.Nares

Buettneriaperfecta
Theskullsofstereospondylsarelargeandflat,withmostofthemlackingmobilityduetothe
solidfusionoftheirbones,asinstereospondylswithmoreprimitiveskulls.In
Buettneria
,the
skullhasgroovesonitssurfacethatcontainedasystemofsensoryorganswhichitusedtodetect
themotionofpreyinthewater,asdosharksandcrocodiles.

Fig.3Skullof
Mastodonosaurusgiganteus
showing

1orbits2.palatalopenings3occipital4nares

Mastodonosaurusgiganteus
Like
Buettneria
,
Mastodonosaurus
mayhaveresembledacrocodileinitsfeedinghabits.The
skullhastuskliketeeth,butlikemostmembersofitsgroup,islargeandflatandthetusks
protrudedoutoftheskullwhichwereonlyseeninmaleanimals
,
buttheseadornmentsare
generallyabsentinthefemale.HABITATSStereospondylswereabundantinaquatic
environmentsworldwideduringthePaleozoic,buttheyperishedduringtheCretaceousand
Jurassicperiods.

EndNotesandConclusions

1.Allvertebrateshaveavertebralcolumnandbraincase,whichisobservedinthelarvaestages
ofthelivingriverandsealamprey.

2.TheskeletalanatomyinChondrichthyescontainsaframeworkofnumeroustinyprismsof
calciumphosphate,whicharetiedtogetherbycollagenfibersmakingtheskeletonflexibleand
hard.Consequently,theearliestgnathostomeancestorhadabonyorcalcifiedskeleton.

3.Temnospondylsincludelivingfrogs,salamanders,andtoadsthereforetheyarelinkedwith
extinctrelatives,allhavingadvancedfeaturesoftheearandteeth.Furthermore,intheextinct
animals,thereisasquamosalflangepresentintheearwhichisabonyplatethatformsan
attachmentwiththetympanicregionandmayhaveimprovedthehearingabilitiesofairborne
soundsintetrapods.

4.ThediskshapedvertebraeinStereospondyls(i.e.
Mastodonosaurus
)wasmoresolidthanin
othermembersofthegroup,andmayhaveaidedinstrongertailpropulsionforswimming.