Editors:AC. HIGGINS,Chief Paleontologist, Geological Survey of Canada,Calgary; and R. L. AUSTIN,Senior Lecturerin Geology;Universid of Southahpton This is the tirst comorehensivetreatment of the sequenceof conodontsin the British stratiqraphiccolumn, which hasWDe sections s€rvinq as world standards.The book provideddetailed information on ihe occurrenceof ai importantgroup of fossilswithin each system,and is an up-to-datesummary of all known and much new research,of value lo scientistsand teachersin industrvand research.
Editor: RONALDL. AUSTIN,Senior Lecturer in Geology,University of Southampton This comorehensivecoveraqe of investiqativetechniques and aDolicationscomDrises a state-of-the-artreview of coiodont sludies,reflecting ihe increabirdawareness of the importanceof conodontsin micropalaeontology,and of the hazardsto human health associatedwith the extraction8nd concentrationofconodonls. The book representsthe. /"-' work of a team of internationalexoerts.
Editor:M. B.HART, Department of GeologicalSciences, Portsmouth Polytechnic Thiswide-ranging review of carbonateenvironments covers the Silurianto rscenttimes. Fossilgroups include foraminifera, conodonts, dinoflagslletes, acritarchs and r€diolariads. Will be of interestto all scademicand industrial micropalaeontoloqists and Dostoraduate students.Many areas of internationalsignificance in hydrocarbon-exploration ar-s dealt with here,among them Oman, Tunisia, Libya, North Sea, and southern France.
Editors:J. W. NEALE,Profeg6or of Micropalaeontology,University of Hull,and M. D. BRASIER,Lecturer inGeology, University of Hull "a principll merit ofthii bool i! tho vlrletv ol topicail contain! ... wide divol3iw of contributionron which todruw ... rnothor gucces3tulfsatuig of th€ book is tho high dogreeof uniformityof cont butioni. nonaof whlch r]o elthortoo long or too rmbhious,rnd only onc of which ir rgducadrgdu to sxtendodabstract dimgnsionr ... . good rcviqwof cur.ontrsssrrch. Thii collgctionof plpots ghouldssrve s usetulpurpose in stimuhting rlnewed int€ralt in this curentlv most us€tul,bst rometim€r noglcctgdbnnch ol the elrth lnd biologicll lcionce!. h cancsnoinlv be recommendsd for relding by both advancsdundgrgradurt€sundgrgtadurt€s and spocialktr rsseotchrsseorch workgrEworkg.E in the fiold" - B.M. Funnoll,Prof€ssor of EnvironmentalSciences, , Universityof EastAnglia, in y'Vatule
Editor:A. R.LORD, Department of Geology,University of CollegeLondon "ths th€meof thc bookis tha biogtratigr.phic valug ot thecelcrrcous nannofossils ..- makosthe point weff" - GeologicalMagazine "will bo widolyconsultodby microjr.hoontologilts,goologists and ocsanographersboth in the Britbh lslgsand boyond"- B, M. Funn€ll,Professor of EnvironmentalSciences, Universitv of EastAnglia, in t /tls, Book News
Editors:R. H. BATE,Director of StratigraphicServices Limited, London; E. ROBiNSON, S€nior Lecturerin Geology,University College, London; and L. M. SHEPPARD,Diroctor of StratigraphicServices Limitod, London "tho illustrutionga.9 ggDorallysxclllgni .., ths bool holdr lomothing of intgtssttor ov€ry ortracodofogi3t.., r vlfuablg additionto lny librrrl/'- JeanM. Betdanin Palaeogeogrcphy. Palaeoclimatologyand Palaeoecology "this is a rollly good compilrtion ... not gno pot-boil€rrmongst ths 24 pap€rs... extremely vlfuabfo" - J. W. Whitlakerid Journalof Naiuhl Histow
distributedbv published bv HALSTEDPRESS a division of ELLIS HORWOOD LIMITED JOHN WILSY & SONS Publishers. Chichester New York . Chichester. Brisbane.Toronto
EllisHorwood Edition ISBN 0-85312-90S-1 HalstedPress Edition ISBN 0-{70-20788-4 Contents
Contributors 7
Preface 9
1 Conodont palaeobiolog5r:a historical review R. J. Aldridge 11
I Preliminary structural comparisons of protoconodont' paraconodont' and euconodontelements H. Szaniawski 35
3 Re-examination of Silurian conodont clusters frorn northern Indiana R. S. Nicoll and C. B. Rexroad 49
-l The architecture and function of Carboniferous polygnathaceanconodont apparatuses R. J. Aldridge, M. P. Smith, R. D. Norby, and D. E. G. Briggs 63
Form and function of the Pa element in the conodont animal R. S. Nicoll
A conodont anirnal from the lower Silurian of Wisconsin' USA' and the apParatus architecture of panderodontid conodonts M. P. Smith, D. E. G. Briggs, and R. J. Aldridge 9l
Soft tissue matrix of decalcilied pectiniform elements of Hindeodella confluens (Conodonta, Silurian) L. E. Fdhraus and G. E. Fdhreus-vanRee 105
An abrupt changein conodont faunas in the Inwer Ordovician of the Midcontinent Province R. L. Ethington, K. M. Engel, and K. L. Elliott 111 t-- = i_-
:'_. _ rF=:_::r - ;\ - :-.
;- .:-.
:_: -i:
a-.4. c:;',- , ?=-__: : -i
:_- t-: n; - _--
I -- - :
IlUtr I 6LI xepur IErauec 9LI epur J XOUOXEI 99I \ sr'a,{uoJItFu rgg 1"ug"q1 ffi;;S ,, L'I snorarluoqreJ-pruoluBruo^ao -o-.,,,ffio*rfoiJ,l"""llX;1"6r, 6Zr uossddel.1 rJu'e'osno*urous;o saposrda .ro;aruno,n" ,"uoo.o,rf,if,llfrllfi#tdY.;jilXll; , Eualuoc OfFnic -E&|ns 129
747
165
175
779 Contributors
Richard J. Aldridge, versity,St John's,Newfoundland, A1B 3X5, Departmentof Geology,University of Notting- Canada. ham, University Park, Nottingham NG7 2RD, England. GoverdinaE. Fihraeus-vanRee, Thyroid ResearchLaboratory, Health Sciences Derek E. G. Briggs, Centre, Memorial University, St John's,New- Departmentof Geology,University of Bristol, foundland,A1B 3V6, Canada. Wills Memorial Building, OueensRoad, Bris- tol BS8 1RJ,England. Lennart Jeppsson, David L. Clark, Department of Historical Geology and Department of Geology and Geophysics,Uni- Palaeontology,University of Lund, Solvegatan verslty of Wisconsin-Madison,Madison, Wis- 73, 3-22362 Lund, Sweden. consin53706, USA. H. Richard Lane, Kenneth L. Elliott, Research Center, Amoco Production Com- Department of Geology, University of pany,P.O. Box 3385,Tulsa, Oklahoma74102, Missouri-Columbia, Columbia, Missouri USA. 65211,USA. Robert S. Nicoll, Kevin M. Engel, Bureauof Mineral Resources,P.O. Box 378, Department of Geology, University of CanberraCity, A.C.T., Australia2601. Missouri-Columbia, Columbia, Missouri 65211,USA. RodneyD. Norby, Raymond L. Ethington, Illinois State Geological Survey, Natural Department of Geology, University of ResourcesBuilding, 615 E. Peabody Drive, Missouri-Columbia, Columbia, Missouri Champaign.Illinois 61820. USA. 65211,USA. Carl B. Rexroad, Lars E. Fihrreus, Indiana GeologicalSuwey, 611 North Walnut Department of Earth Sciences,Memorial Uni Grove,Bloomington, Indiana 47405, USA. 3::ql Jo II€ ',(lsnol^erd l:!6 I ) uepe^\S 'Pun'I ur ! ?suar^ Jo se4ueJ Isnp art-rliol'116I ur',{u€ur :EJ:l 'Srnqre]^{ uI PIaq FESrlqelse auloceq a\ou ou.-r3 ueedorng eq1 ',{1nr1ceqcreese: s rJauar (]1 puB luelx asaql Jo sesod ir.nuaqcoa8-Einlo^ cIUeeJo Jo -;onEnlE S uoll€Jnl"Iu ficnlo \a'fSolocaoaepd o..:-i aqt ]o anle^ eql nur ilprder eqt ,{q pu" N-UOS JO ssrJa^Osslp (ts -ra{ro^\ luopouoJ eql r :JP sseql SInOAeePUA }c' uortecrpUtleelc e a,rl3 o:o .plarJur€u aql lle Jo p::01 s{ooq eerql esaqJ | :PrJrrroc ol pacnPoJd su];-;rH J Vf,qPalpa F-,.rqd?rSqerls€'prrrll lF:lrpe ueeq sEI{'suoqeJ JlilP;rlse^ur uo'atunlo^ '. 'i;rrl 6Z-92 por]ed aql ) lrs.re^run aql l" pleq Lis ruopouoJ ueadorng a:i r:aded uorJ ,{Ie8rel ? irr euo SI atIInlOA SII{J
eJeJeJd
'lneN ?ruep€{V pr1s1o4'rrSolorqoap dpel1.az ,r{s^\euezs UeqnH 'fuerureg go cqqndeg prapag 'puefug ,OA€ eEpuqrxeJ .taerls 'I urBW rup gggg-q ,97 aEelue8 ZSJ trnplu€rd Surumoq'eSprrqrue3yo flrs-re,rrun,saouarJs -JequeIrues'8:aqualcueg lnlrlsurs8unqcs:og ,qg7 queg Jo tuaru edeq 1e .tou fpuelSug 'ra13er7q14y 4gp ueqEu41otr1 'Imd ,(lrsJe^run .ur?q .,{EoIoeC 'pu?lod -8ur11o;1yo,{lrsrearufl Joluaulredeg .IV 'ttesreid 690-20 '96 frn3r16 I l{rl,t\Z 'ql rs Ined I^
srotnqlJluoJ 8 F
5c:cr:::e:: Snckenber- U ---r--- - =: \lain 1, J -'t-:-r:.
Preface
This volume is one of a pair that has arisen European only in terms of their location, and largely from papers presented at the Fourth all have attracted participation from through- European Conodont Symposium(ECOS ry), out the international community. The ECOS held at the University of Nottingham during IV meeting (with two field excursionsto the the period25-29 July. 1985.The companion Welsh Borders and S.W. England) was volume,on investigativetechniques and appli- attendedby more than 100 conodont special- cations,has been edited by R. L. Austin, and a istsfrom 29 countdes,and lecturesand posters third, a stratigraphical index of conodonts were presented by speakersof 19 different edited by A. C. Higgins and R. L. Austin, was nationalities. The meeting catered for all produced to coincide with the Symposium. aspectsof conodontresearch, but two one-day Thesethree books togetherprovide a coverage sessionswere reserved for the two major of all the main fields of conodontresearch, and themes:(1) investigativetechniques and appli- give a clear indicationof the direction of future cations, and (2) conodont palaeobiology. endeavours.These are very exciting times for Apart from the introductory chapter, all the the conodontworker, stimulatedby the recent contributions in this book were formally pre- discoveries of soft-bodied conodont fossils sentedduring the latter thematicsession. and by the rapidly increasingrecognition of The papers which comprisethe chaptersof the value of the group in biostratigraphy, this volume mostlyfall neatlyunder one of two palaeoecology,evolutionary studies, thermal headings: (1) the conodont animal and its maturation evaluation, and the investigation mineralized apparatus, and (2) pattems of of oceanicgeochemistry. It is among the pur- conodont evolution. Chapter 1 provides some posesof these volumes to convey that excite- historical background as an introduction to ment and to reflect the diversity of current these two topics, and endeavoursto highlight researchactivity. where the most vigorous current debatesare The European Conodont Symposia have centred. In the following tive chapters the now becomeestablished events. The first was themes of conodont affinity and of the held in Marburg, Federal Republic of Ger- architecture and function of the conodont many, in 1971, followed by meetings in the apparatusare addressedfrom widely differing dual centresof Vienna and Prague(1980) and viewpoints. Szaniawskidescribes the internal in Lund, Sweden(1982). As hasbeen stressed structure of the earliest euconodontelements previously, all of these gatheringshave been and their putative ancestors,and stressesthe anprrplv'f pnqrrx aqrur ol roud Fl '886I 'IPUo4?UaluIseca-re5 crqder8r1er15 pue pue nlderz epr^lpFoa{uoq"tueurrpas pcrddle E lsedse raqlo ,{u" Jo 'c1dlueg puepr141aql 'OJIUA 'petrutr-Ipoo^\ Jo ss atuq 1eEuurncco suou?rcosse pezlprcads rqp relEerS € peJrdsur ,AUEdtrIOJ -IOH SIIIE EPIISOIJIWIINH 'X 'g pue asre^rplsour eql qllt\ .suolllpuor JrueaJo rxII pels"Ju€ flur€ue3 .{larcog 'c1dlo1ug 'c1druneyorleg qsrlrrg pcr ur se8u€qf,ol spun"J luopouor u?unlrs eua eseql sarlrurJJe Jo Jo -3o1o1uoee1edorcr1a1qsDug eql,lbunoJ rlsrlrrg tuaudole^ap aql sel?leJuossddel .crsser{ €ql Dd eqJ .,{Sololuoepd aqt 'uotdureqlno5 pue ureq8urgop;o sarlrs Jo pua eql o1 qSnorql u?rJnlrsaql luo{ Jaao b pa.raarsueunlzluaru eql dq pelroddns -Jalrufl flsnoraue8 se,t,pue -uJnl Ieun€JluopouoJ Jo suJallzd pJooel sJel r aq o1 slsrSoyoluoepd ,(leroograpue4 aq1 ,{q palu rlrur se,r gg^3g n1 -deqc luenbasqng lueruacelderlecrSoloce ;o FopouoJ ernleu eql .lnoqinoJql Jo pur{ aurosploJer 01 steedd€l€ql p^lelur uerc ' t3qauoa1n4 a ao4 eJu€lsrss?pu? e3r^pe a,re3 poo,rruoll sqlg -r^opJo flJeg ue qEnorql saEueqcpunel eql q '1861se dlluecer sy ,asrgedxe Jo JJBISeql pu€ uoslllrC selx?f pu? Jo lunoJJepelplap e e,rr8(g raldeq3) .121a uo1 ctqde-r8o1:ecpapr,rord uosur116 H)IJJNOOUINI I'I ersol,11aEuy -3urq1g'aldruexa ue sy urelled elqeredruoce u?af ,{g euJoq ure8€s?,rr p€ol lerr€lelJeseqJ s,toqsd:o1srq dreuo[nlola luopouoJ leql a]aq 'saeraJer aqt pu? sJolnqrJluoJeqt Surpnpur peluesardsreded aq1 uro4 luaprle sr tI .stusr 'elqrssoo eIllnlo^ srql epElu oq^\ esoql I[€ ol -ue8-ro;o sdnorErelncrged auos ut pup eloq,ll enp osle .s{u€ql .sJeqlo n a.rars8urreq1e8-poo; eft $lueql eteJurs,{ru E sEproceJ JrssoJ eql ur pezuSocarflSursearcur ,{ueur pue .sualqord 'Jo suoddns s? sluaur 'oser{l Jo 112o; rounu ueeq [lluaJeJ s?q Euela uoqzrper a,l4depe epnlrllnu .uorlnlo^e I s.raqlo q8noqlle 'rusr Jo E pellos pue sraqureru8ur,{ued /uorpunxe ut &rJrlJ,{C luopouoc rfuidse:8 e se snleredde -trroJc€JoJ aruure-r3ord€ pezueElo,uosqv ;o crdol eql ol et€leJsJaldeqc JnoJ eqJ 'uosuDllr/!\ ,lTnJ I?urJ rD leql sasaqlodfq 'eJra ,{trl ersol pu€ uourrs 'uorle8qse,rurd:eurruq ,(puv'uosJeadu?ef ,upao ,qa8uy lE sra{ro,!\ IsJsAes 'gr/r\S {Jery -erd srql ;o esrurordeql sulrJuoc lloa panuq u ol ruees 'ra^aA{or{ ueal i(lercedse 'suoqnquluoJ -roleru aperu -uoJ Jraql leql odoq lsnur e1d .fEolorqoaeled sen8eeloc pue spuer{ reqlo .qlrur5 ![P tsasolr eqJ 'PaPhrP duey41 Jo plarJ eJoq^\ eqt ol lnq ftrurJJ? luopouoc pu? uqsnv pleuoll SurzrueS-ro 'BuaurnSre eql pel?Il In?d 'aaltruturoc ;o uorlsanbaq1 o1 acue,reler;o 1sn[ 1ou se4llrqrs r par-rasard qlft\ suerrJ aql uo sroqueur-oc ,{ur yo suoJJa aqt e8pe -sod Surleurcse; suado suorlerederd rroql o] a^€q,{lrElncrU"d .sluop JO seua^oJsrp lueJe)I l,\{ou)lce I.srql log.sseJcns ur pueleur replac ;o uorlruSoceJJreql € uaeq eAEq o1 sreedde ,uoll?zl Ernala lelalels rreql AI SOCS -ouoJ Jo Eueuale pezrlBreuruteql uI pe{col -ue8ro eql ur pellolur . pua-1eru rme8ro eql Jo sanrugJe lasoycsr auo ueq,n8ur crue8ro aql urcls pue xrJ ol slueu p moro?r,r {q pelleru -1earue eSpnfo1 tlncr;;rp ,{re,rsr 1rq3noq11y -uedxe rragl Jo sllnsereq1 luese.rd (1 rsldeq3) ',{-4gruraqooaS .uoDnqr! lEoJ Jo ,{Jotsq eql tuopouoJJo prlualod apr,naq1 aou u?^-snergyd pu? snerqgg Jo uorleJrpurauos 8ul,rr8,eurnlor uorueduroc -uoc rfieuoqnlo,rarfllenuetod lnq Uoqs p uI IJVUISSY aqt ur papnl3ur pu? .1sed erE AI socg te pelues eql ;o asoql uuql -erd eJa,{ adfl sql ;o suorle8rlsa.lurluecar 8um8rr1urssal ou srfsrerrorluoc lueunJ eql pue seldurexg 'sa^lesuaql ;o slueurela luopouoJ 'eleqepf1e,rq;o crdol e ueeqs,{e.np s?r{ Ieurue .sprourx,{ru eql go srs,{1euepcmraqc q8norql ,{:lsruraqcoeS luopouo3 erll Jo eJnl€u eq1 aq1 a3pup1171y Jel€Ai-Beso1 ,{1pallp slua,la fueuo4nloae qtft\ ,trulJJeu? sraqto pue .saleproqcolzqdec ar1o1 .asrruord ,sqt€uEoleeqr elqrssodaq [Eru lt se ,{gercadsa eql ot sseuesolce IIocrN eql qll,$ ud luopouo3 esuerrJrrrrsploq ?aJ? srql ur serpnlsJo etnlnJ 1uq e Suunoru; r{s^r?u?zs qll^\ .sdqsuorl?Jal eqJ Jrsser{ ete.I aql ul dnorS aqt Jo esrurep luopouoJ ;o ssaqlod,{q lepc4rcd B sprea\ol eqt I€urJ ol eurTJapsqt secerl 1-re13,:aldeqc ua\€rp $ sJoqlne oseql Jo qc?g 'aporrr s^rs IBUrJeql ur pu? '$ped larlJeeeql ur ueql Je,{{ol -sederou reqla8o1pue ur pooy8ur-reqleE pue I s^?^rp sp,r! d:o1srqluopouoc yo sa8els ra1el peJsAOC-enSSIlS" EUaUAIa eql ,{\3rAp"oJxell eql ur flrsra,lrp lnq 'Jrss€uJ pue uerrrrJadaql pue IloJrN alrq,{ 'sprourx,{ur snlereddepn? .seseqd ;o olur rncJo saslnd-reglrg cqsqenperS -uq el4eredo dgerelelq eqt pu? sasnluredde ueql a^rterp€r pe^\olloJ suoDcuqxe luopouoJ uae,tlaq sarSoleue,trerp o1 -relard qlra\ 'snoreJruoqtJ^q flJ€a pue upruolaq eql '1,?a qlrus pae pp a8prlply .sqteu8oleeqe q8norql slue^aJrlc,(c;o sarres e ezru8ocersuel uJaporuJo saurdsEurdser8 eql ol serlrJ"lrurs
aJBJard OI EE- :: -- : a :1::1ISthIOUgh d :-- : :-ious. -., with G: l. e then r. ! nu- :-=:-::,:::\ occur into Il:i=.r: -- :,::sin. in the l,tlr;i; :]:: t:__. *as always t:]r- :E a: ia the final I G:1:i:: :-=: :--: iO the final + r --:. _,!:- f:iassic. The I :l-: :-:: :: l: immenSe Conodont palaeobiology:a historicalreview l.L - : :. :. :!.:!ible tO tie G : ::-*--. :-. Sea-water R. J. Aldridge l!!n :::: ::: .raltsis of the f :li-\- .:. Eramples of rE : : 1: :,.;< $ ere pre_ I r:-: .-: ::::..rded in the ABSTRACT of the soft tissuesof the conodont organism, : rl'._: lr:-: -ndication of hypotheses ranged from the carefully con- The history of conodont studies has been i![ : : :,': ::: :-L-lchemistry. sideredto the bizarre.and taxonomicassign- tiJse marked by vigorous debates concerning the lt ::t:---::: a meet- ment from the algae (Fahlbusch1964) to the affinities of the organisms and the function of m :. t'.:: n the organ- vertebrates(Pander 1856, and many others). their skeletal elements and apparatuses. rc-,€r-: :-. :ta\e been a Miiller (1981, pp. W78-80) listed 46 publi- Recent discoveries of complete fossil speci- lI:L-- r--" :=r e ro acknowl- cationsthat appearedin the period from 1856 mens with presewed soft tissue have concen- i :r-. : ---::erbers on the to 1975, variously suggestingaffinities with paul trated the arguments, but opinions are still ts i : t-: 1..-:tinand plants, conulariids, aschelminthes,gnathos- divided. The closest affinities of the conodonts, Er :-: .:! =:J colleagues tomulids, molluscs,annelids, arthropods, ten- ._ however, seem to lie within the chordates. rrr---,. irpeciallv Jean taculates,chaetognaths, and chordates.These Several workers now favour functional i "'.: :: =:'+.:r. -\ndy Swift, assignmentswere commonly based on mor- -- hypotheses that consider the conodont hsr 'i l:n:r-rn. \ly wife, phologicalcomparisons of structuresin extant apparatus as a grasping and shredding rnechan- I :!: : :-:::13 ior accom_ organismswith conodont elements, although , ism, although others prefer to view the ele- at: it : multitude of severalof the similaritiesare superficial.Some -:--::e ments as supports of ciliated tentacles in a [c .: :e- and many hypothesesrested on associationsof elements - -. food-gathering sieve or lophophore. &r', :=ris are also due with otherfossils, although Hinde (1879)and, ---:: Er- ,.,1'.:ne possible, later, Rhodes(1.952, L954) reported that con- rF-: ti =: the referees. odonts may be found associatedoccasionally r..:. I torne by Jean but not consistentlywith almost every type of 1.1 INTRODUCTION lr: r :::--:: ,:::d canographic contemporaneousmarine organism. The dis- E --. i.: :nd the staff of As recently as 1981, in the Treatiseon Inverte- covery of fossil specimenswith preservedsoft re .::i :: :nd asslstance brate Palmnbbgy, K. J. Miiller wrote that tissue (Briggs etal. L983, Mikulic elal. the nature of conodonts'is consideredby many 1985a,b,Aldridge etal. 1.986)has concen- --i:i !ftrla: : , Pander Society paleontologiststo be one of the most funda- trated the debate,but there are still differences f!:-': rr rhe Univer- mental unanswered questions in systematic in interpretation of the nature of the animals r --:S-.:ihampton, the --: paleontology'.The problem of the zoological and of the function of the conodont elements B ::r: \:--!rpalaeontolog- affinities of these enigmaticextinct fossilshas and apparatuses.Consideration of the com- P::. .:-= rlc. Britoil plc, certainly attractedmore generalattention and position and structure of conodont elements h- - :::.:1. Ellis Hor- inspired a greater diversity of opinions than and of the architecture of conodont appara- Cll ::: \L:l:nd Bank plc, any other aspect of conodont palaeobiology. tuses continues to be of fundamental im- IT:':a: -:::_national. Prior to 1983,in the lack of any direct evidence Dortancein thesediscussions. i::::rd J..{ldridge + PaJnPord(896I) dlsnorcrpnfpue '1ce;redurrse,n dSopue aql terll 'uorssrurrodqlr^{'(9261) uosl3uagrege f sauas elqsde sno pezuSoca:lnq 'sprourxf,ruSulsrxe rlteet aql 'sl[eurelatuopouorna (J pue luopouocerEd(q.luopouocol Jo -ord (e sre,{elcqeqdsoqd eql ,{SolotsrH-I .fiC rr 3ql lnoqe suorsnlt qll,t\ seurJslutrspalou eH .scsn orrrJo sprleuue Jo Jo I FnoqlF 'slql paurnJ o1 paSuolaq ,{aq1 leql uonou eql peloaleJ -ts6l pue I preEurla 'dSoloqdrou pue uorgsoduoc ;o spunorE I 'sdIIIqd '9t6t Sreq aql uo 'pue 'uoqezqrssoJ;oelqzdec rusrue8ro r$enbasqnspuE 'eddl eqt Jo stled ,{po eq1 ara,r slueuale aql leql rus Ptsdlr eql leql paar8e(6491) epulH punoJaq le^e pFo,t\ pru rlFI luanlDsuoJluep -rue luopouoc ar{l Jo stred raqlo 1eq1,{1ar1qun oaoqs (I98I),{elreH s"^{ lr 1?q1peuos€eJ (gggI) Jepu"d ,slueuele E PalJalap relel tnq tuopouoc Jo suoDcalloJsrrl Jo srssqeqt uo rruoJ eJe,t\ feql 13q1 'un8eq ,{1preqpeq et€qapeql .fq"er uI ) rapu?d ,(pnls Jreql '.tsal te las,{1e1e1druocsr uopsanb eql, }Eq1 ffioJaJ se,r slueuJele er€lrep ol 1t.1,rt.es (98€ 'd 'tSg1) uosrqrlnn .rggI .6s8I tx Jrleqdsoqd eql pu? 'suoqrpeTL9I pue eql ol 6fi 'd 't96I sapoqd) secrpuaddy ur ua.u8 e:a,r ua,r6 ,{q sgoder |Irs aleJJes01 su[o,$ ra ng (9€g d 'r9gl uosqcrnr^{) .seprteuuv 6uo a1e:qa1re,re ;o puP s{sn[otr^lpelEu Jo salJquepJo.slaplooq q (areqdsoqdurnrcpc) 'seurdsaq1 q1r,n ,{Soleue tsour eql, lueserd[aq1 ayq,n'ece;:ns reddn eql punoJ€snonuquor aJe r, leJrrrreqceql lsnhtr, t€ql SulpnlJuor s? uaa{O pJ€qcrg papodar u,{loI' ern Jo a€ aIIIel eql ',{poq ps"q ? pu€ 'luegodrur-1e re se,t' Pu? uoqquep qsrJ-al nr ol slueureleluopouoc u,t\om euq",(q e olur lrun aql Jo uorsr^rpB sr 'seleJq ,eSgt) .fl/fl/rs' tfl r\3r^ srquI peJ?duror (Vtt.d uosrqrrntrl aJeqlslueurele'.luopouoc arul, Jo'luopouocna .(1991) ssJnleerc ell(-txro/ir Jo suoqrpera1e1 u1 fape11 ,{q pre,trro; uI (9L6I uoslSuag '1461 lureSop pue .^\ara roueraprsuoc JequnJ lnd se,ln'saurds uzacelsnrc ara,t deql l€ql rallnyq)dp eqt punoJzsnonu4uoJ lou tnq 'eo€J Io sauoeql o.t'.1,{po J€lrurs e pue 'slueruSaselrqolJl Jo spua eql -Jns Ielno Pu? rsuur eql lE peleJcceeeflerrrEl : ;o s,ne[ro qlaal aql aq o1 ,{1e1uerorx era^r daql teql eruq lpq} t? crleqdsoqd llqrqxe slueuale luopouocered , lEql s€a\ uorsnlJuoJ lleJJlasurrl uosrqJin6 qleelqsu aq ot tq8noqt 'lsertuor uI (gSOf 'ZgOt 'Dls^{eru?zs'q€g6I qs puE 'uoDrsoduro3 serpoqalnurur o1 SuuJeJet eruerts Jo ,{urepecy '9161 uosfuag) aseq eql pu? eJEFnsJauur ,xp uo ?aPrs.ur€quecl q3uerCeql ol IggI ur tepwd uauu,t\ la]lel egl l" uo[arcJ? seleJrpurJefel pazq€reuur pepunodord peq ,VSgi .rrunlls ueeq e pelrodar '(ftt,'d uosrqJrntr^q l eqt ur ee elllel ;o lueura8uerre aq1 lsre(e1 are ilecrlrrc pue paql srqut aloulooJ u1 s1sr8o1o1uoee1edueedoJng crue8ro urql [q dleu:a1ur puz fleuralxa " nssn Uos ro; sgoddns IeJeAas,{q pessnJsrppue uaes ueeq ,(lJeelJ papunoq sr ra,(e1 alppru.r crlzqdsoqd 1crq1 |zCr (It6I) ss€HPu€ peq suarurcedss.rapue4 uorlecqqnd slqt aJoJeq E slueurele luopouocolord ep u1 (1 1 3rg) bld aqt pue 'srurot\ Jo t€ql paprorar (ZZ6I) eppnH'gsg;o dnor8 sluopouocna pue'sluopouoo?Jed'sluop romts f,rolepdoc aq1 lJr4xe u€ Jo qteet eql se rueql pepreSerqder8 -ouocolord s" ureql peterluataJJrppue sluop .H ,late^oJ m+ (ttOt) s.u?queq -ouoru9SgI srq ur ogal 'repuEd C -ouoJ eql o1 alqeuSrsseere,r sedfl esaql ;o suo4saSSns eerql Suunp -slp Jreql Jo leql ssn slueurela luopouoJ Jo l?ql perunserd(9461) uoslSuog (qgg61 llsuaqoyg ees)rnole; eJnl€ueql uo passe-rdxeuorurdo lsaqree eq1 uoslSueg) sarnpnr1s pcr8ololsrq ;o fleue,l osse qsrJ Jo prleuue 'sluopouoJne e deldsrpsurro; uerrqrleC pu€ ',fi€punoq ueuq 8ur-req se uraql preSer ol ecuaJaJeJur pesn fll?rurou sr .]ueuala -IUEC-UsrJqrrIererderll ol asolJ pJoJeJIrssoJ ,sla{Ioil\ lsoul Pel sIIIsl luopouoc, rurel pagrpnbun aql pu? .sluau aqt ur reeddz lsrrJ quauele crlzqdsoqd pc rFael aql ol slueuala -ela luopouoJne passessodereueS luopouoc -IuoJ trun aq1yo q1nor8 eql Euunp dllpEnJl4 rP aql 'seurugJe luopo ;o ,{luoferu 1se,raql (qgg61 uosl8uag) sluop -uec pue ,{qecrrluecuocpelarcce ara,n seddl mn aql ge alrdsaq -ouoceredaq1 eq1 q1n sno8oloruoq a,lrlrruudlsour aq1lnq 11?ur qJrq,t\'e€Ieru?l rn '(sss ;o ,dsnc, d aq deru sluaruele tuopouocne eql Jo fpoq -eqdsoqd;o pasoduoc eJ?sluauele luopouoJ eqr epbep ol luarJIJ aql eJ€JJnSJa^\OI eql punoJ?snonuquoc ' I?Seq SJNqIAIU]S INOCONOJ TO NOIICNO.{ 3111, pepnlruoc 4 1?q1 eq lou [Bur ro ,(eru ,{poq Ips?q eqt Jo esoql (INV'SUOJJnUTS 'Norrrsodwot z.I rJ [Z I res I'qJ] 1e lJolslq e :f,iioyorqoeepd luopouoJ ^ra!^ar ZI r [ch. 1 Sec.1.21 Composition,structure' and function of conodontelem€nts t3 hr-- ::-:. : ::]a\ not be concludedthat 'the facts at hand appearinsuf- and regardedthe mineralogyof Devonian ele- b r; r =- .--::::. The basal ficient to decide the question' (Hinde 1879, ments to be of the carbonateapatite, franco- tr!3t i: : .::::,: may be p.3s6). lite. Rhodes (1954) consideredthe composi- E --r- :: ---: DataCOnO- Despite all the uncertaintiesregarding con- tion to be essentiallysimilar to that of the bony --- lB: . :neiority of the closeresemblance of many armour of Devonian fish. This, together with -.- odont affinities, rIn - r-- : -::::t-tont ele- elementsto the teeth of various extant organ- the 'bone-like' natureof the basalbody and the qi;rl:-: :-:l COnOdOnt ismsled mostworkers over the next 75 yearsto tooth-like appearanceof the crown, provided a b l*: : :::erence to regardthem ashaving beenteeth or jaws, with major Iine of evidencefor thosewho favoured annelid or fish associationsparticularly in a fish affinity (e.g. Ulrich and Bassler 1926, I =i:-.:!€: :: Ihe natule favour (seeGlobensky 1970). Other functional Bransonand Mehl 1933,Schmidt 1934, Elli- G L!: -::: t; rheir dis- suggestions during this Period included son 1944). However, Gross (1954, 1957) t r:: ::.1!56mono- Denham's(1944) theory that they represented pointed out that conodont elementsgrew by l.L ,:: ::a_-: ..I an extinct the copulatorystructures of a group or groups the accretion of successivelamellae on the & --l :ctrrded that of worms.and the proposalsby Huddle(.1934) outer surface of the unit, and are thus not n i := . :-crmens had (1941) served as internal homologouswith the enamel teeth of verte- -- and Hass that they m: .:{ -.-.-: br several supportsfor soft tissues.In 1954, Rhodesout- brates. Grosswas still inclined towardsa rela- hrL-,: -: : :--..rnote in his lined and critically analysedall hypothesesthat tionship of conodontswith fish, although he lii- : :l--r. reported a had beenpropounded to that date, and refuted weakened the hypothesis still further by tr- :: : i - :!r rhe French Denham'sidea on the evidenceof morphology, demonstrating that the basal body was not K:-:::t:ninute bodies composition, and size of elements. Rhodes' 'bone-like' as it lacked blood-vesselcanals. d \l -::: ..1nhimself felt conclusionwas that conodont elementswere The idea that conodont elements shared :! r: = ----!. likeh to be the teeth or jaws of an extinct group, and that affinities with worm jaws dates back to 1858 i =.-::--:- =rd a similar only two theories of their affinities deserved (Owen, In: Murchison 1859). The hypothesis q-ll:;,-?:: i:lleS. $'aS put further consideration:they belongedeither to gained further currency with the demonstra- -: li: - '- ::er editions of worm-like creatures or to primitive verte- tion by Zittel and Rohon (1886) that the ele- t:!:: : _--rt compared brates.In his view the compositionof elements ments resembledthe masticatoryapparatuses n :--.-:-: i: denrition and was all-important, and he posed the question: of annelids,especially the echiuroids and the )*:-: :,:ncluding that of conodonts sipunculoids.Compositionally, though, annelid -. 'Must the chemicalcomposition E ::- :1 rrth the spines, (calciumphosphate) be interpretedas evidence jaws are formed of chitin, and although some = ; -.r.: \Iollusks and of a vertebrate origin, or is it possible for worms produce phosphatic tubes, none sec- a!i: - r-- :. -r-:6). Fuller worms to secretesuch a substanceinternally?' retes calcium phosphate internally, which rE: :-r ::: rn _{ppendices (Rhodes1954, p. 449). would be necessaryto produce the external ri- -.-: editions, and The phosphatic composition of conodont appositionof lamellae.A furtherargument is '- J l: ir to declare elementswas recognized early in the history of that annelid jaws are built by the addition of s ::=: -;:::.r let at lest'. their study. Pander (1856) believed initially material to the inside of the base (Lindstrom e r-:: :--Jr t'egun. that they were formed of calcium carbonate, 1964) and cannot grow in size like conodont tr ::r i,-:lt! of conodont but later detected calcium phosPhate,which elements. LindstrOm (1964, p. 120) con- t---': =---:ed rhat it was Harley (1861) showedto be the more abun- cluded: 'The situationis evidentlythe samefor nr--. -: --:: conodont ani- dant constituent.Ellison (1944) demonstlated the annelid hypothesis as for the fish m': :-: :.:: r1S-9) agreed that the crystal structure was of the apatite hypothesis. It is false if applied to known, Er= --:- t:-'. pans of the type, and subsequentanalyses (Hass and Lind- existing structures.Otherwise, it must be so d:{! -:: !rn- and. on the berg 7946, Philips, In: Rhodes 1954, Rhodes vague as to be rneaningless.' Er'r- r:J :norphology, and Wingard 7957, Pietznet et al. 1968) con- The evident weaknessesin the hypotheses L ---: :.:i itelonged to firmed this, although producing differing con- that conodont elementsrepresented the teeth, b: : -::: .:itllariries with clusionsabout the membersof the isomorPh- jaws, or radulae of vertebrates,annelids, or I-r-.: t :i. ::i recognized ous apatite series involved. Pietzner etal. molluscsled Lindstrom (1964) to developthe Er--,:. :ri judiciously (1968) produced the most detailed forrnula, proposalpreviously argued by Hass(1941) that idr aru€u aql pau8lsse pue goddns-e1ce1ue1 'lepour aql roJ arue uorl"uruexe ruorg 6ueq13ue-4sfoql prp ue8:o ntuolaq a,req lsnu 11e -re;erd penuquoc e pesseJord(9961) srrroyg Jo puDItEq,1\ ueql .sloddns ptele{s ere,t\.(eql r't\ENUI AFqS ,ttlotflaT I IIOA fe,ruo3 uorlecylqndleur8rro eqt sE 'palse33nsp€q sseH se ,JI .aJIISuunp pesodxa aJeFns aurppeq I Jo 'purnof arues eql ut suonnqrJluo3elrsJnJsrp re^a aJea\deql leql eurnss?ol uoseeJou sea{ I)QPTJOSSE ul pAAIAS uoqs Jo tas e o1 pa1raded 6461 s,uossddel aregl teqr (nt'd 'VgAt) uo4spur-I pepBns mJ're^aaoq'(6L8I) 'sera0loqdJotu -rad ql,{or8 naqt lnoq8norql snss4,{q perenoc . |tr perlnrco aA€q II? IxroJruoo pue urloJnue.rfueur 1msplnolir qcrq^r aq ol sluarxeTaluopouoJ ro; fllssaceu aq; puB 'qtoours .adeqs I 'pelaol 'Suqeeur tnoqlr,t\ ds?J8 d€ur [aq1 ro 'aellauuletrtede lsouuelno rIIr lr perePrsuoJaq alerrdordde u" el€q sluauele rxJoJrurpo{ aql Jo uon€nuquoc e 1o flar4ue palsrsuool?ql plno^{ ,l?aqs P aq IPnprarpur -r€zo qcrq/( toJ ,{euI ,{aql Jo.slueuela uo4eue,rnfere ueql teleaa eq plnoa oJn]c€{ s",ti ,tal^ (998I) ,,{l[eclu€qJotx tql ruro;1e1df,ueru ro1 peEesr,rueaq ,(?ru r{crq,,{\ pepaq e eJurs snoa8Etueapeeq brur ue u1 peuoqrunJ 'apnlcro ,{€ul qleel sngJ suoqJunJeleredsrp pFo,r\ lleJ aq 'srql .paqrosarro dpoq aqt uro-ry Idz ye1e1ar1sluaureye rraql fq paureldxe aq f1-relurs p1nocsluerrlele palladxe uaeq e^Eq plnor slred ue:1orq1eq1 u:6 uA\ool .suSrsep -{aql lzql luopouoJ ur secuare;ypssor8 leql pue Suruoseerfq srql paralunoc (y961) ruo4spurl das se ureql peleerl tuaraJJrp Jo qla6t ,{q peqsrldruocce araarr 'pasnJ-eraq ppom suorgod peJnlJe4 o/ru aql UOnf,UnJ .atrl ||IOpOuOsJO elrq ;o sedrfitueJeJJlp leql petou os[? eH f€ql ,{Jalrl aJour$ lr enssnur peppaque eJa/i\ qr yo qcnur qSnoqtyy -€de 'leueleru eues aq1Sursn suorlJunJ Jelruns lueulale eql;r ilso1 s,{e,np lsoutp sr uorlrod urro;red It'ddv JNo(oNoJ o1 setnlJnJls lelurrs Jo sdnor8 o,n1 ue{orq eql ssecordsrqt ur leql pazrseqdruoeq sq1,{q .drqsuorl EIJHCUV SHJ, €'I lueurdole,repeql pa}Jegar1nq 'sluauala ualorq Jo uoqereuaSaraq1 o1 pte8er -elal Jluouoxel due ,{ldurr prp lou serlrJeJnlrs qlr |I?rJel€ru snouDlqJ.lauos e ]o per[JoJ esaqt ler{t sserts ot sured ge searr,(646y) ere daql q8noq1 ua,r.e.uortu e oJeJlns Jo uossddal 'qleel aletqauel pu€ slueuele su8rs,noqs ,{1erer slusruale luopouoJ se eleJls 'ecueldeoce luopouooueel\leq serSopueesolJ Jo JeqrxnuE erxeseql uro{ (stuopocaloJs)s,rnef alaeqc,{1od 'L86I psorxau pelerlsuouap ,6461) .Jee,!\ f,mrN osp og,t\,(096I uossddal JISSOJleql palou osl? eH luecryuSrs puu .peFaJJe DIII e (L861 IDta ,{q pecro;ure: are,lr qurod esaql 8ultroqs lnoqlu( qloel se uonJunJol slueurele uoqcunJ qtool e eq pFo^{ sdp aJcrlueppue {t') dsnJlsol pu? eql lrtolJeol luercqJnssl alrledu Jo sseupJeq h.sr-gt ur 'ecueH uolle ua{oJq^lrpear uoqeJolsar,eJorrrtagung 'papedxa Jo aqt leql lno pelurodpue'aJqrns lero eql uo al slB(uruB luopouoc aq lou plno^\ eJ€JJnsJelno eql uo Jee,!\Jo uot1rr11elueredde Suuroqs sluaurele acua pu? ;o afl ue^e 'uorlel etuaprle puE 'sporradpuorpunl uaal{teqA{ol3 -lsue aql pauoder 'ra^e,roq .(7961) sapoqg flslr-\auraJe araql lnq ot onurtuoJplnoc tuauala eyoq,raq1 .1sarEur 'esn ol peqrrcs€oq Aluo plnoc p lEql u1{oqs,{Peerl€ -rnp papeJlar pue uortrunJur uerllr{lalcod aqt uourpuoJ Jreql ,{e,r e qcns ur uro,r sddal s-re1-ro,nluop txo4 PepnJtxaoq ueql plnor u,lrrorceql .,(poq l€ql suaurrJads spJoJeJou etea{ aJaql pue ce 3uraq ,{Soyoqdroru les?q eqt o1 dpo 1ua-rsqpeenssrl Su4arces Jo u8rsou ,noqs , aql ur palFseJ ,l'eJ eql qlr,'{rsta{rod ur pe;1n8ued1e1e1d 'ree,r ;o ,{luourruocsluarualg (g) leqaqlrda 'uoq g1) uosl?uag;o arer,r. -rrroJeJea{ sluaurele luopouocna 1eq1?ur1e1n lfurnp pereloc-anssq -eJcJerelletrrp[ luanbasqns Suunp perolser -1soddq uoqcun; l€uraue tnq q1.l.roJ8Ieurelur uaql pue de,tre ualorq area\ eJnpnrls rla luopouoJ reqtaq.{\ ;o xopered lueredde eq1 eruec:eao (9461) s1-red a\oqs ql,tror8 su aqt uo luaruaerSe uoslSuag 'arp 01 pesnJeJseq slrun Surdser8 -rraq11o lpql se8els stueurals snoreurnN (Z) lleql se^rlr '8uqt auo ro qlael s3 peuo4JunJ Eueruale luopouoc II€ Jo luopouoJ 'urnrpaurSurlerces ErJO s/a?lc slqBpe4eJ t€ql eepr aqt'(t96I) rxortspur-Ipue (It6I) aql uror; peldnra peq pue u.tror8. e-re,r,r ;sra.ra;o f,llpqrssodurr sse11,{q pelq8qq8q saqpcrJJrpeqt elrdseq 1ela1druoc {eql rage peuuoFad e,req ,{po plnoc pue .qlnor8 aba plnoc ,suralqord .Jelel passnssrpeq ?urrnp enssrl dq pepunoJrnseraar,{eq; (y) f4a'(086I) uoslSuag ru^\ ,(lrugJeluopouoJ ro; suorlec4durrs1r pue dq aperu .snleredde 1161) locrp sseqlod,{q srrrroJtspur-JJo s1?leq :spre e,u1se8urse peuoq qord acedse 3urraq1u8-poo1 lueserd e urqth{ seloeluol pauoddns -cunJe^eq lou plnor Euauele eqt pe^erTeqeq o aldrurs ueAe rrro{ ,t.tAt e,req lq8rnr teqt pa1se38ns(V t1t ,(q.trsuosear eerqt tno palurod dlnlecro; sse11 htoad aqt 1eq1penEre urorlspurl^eql .safoloqdJour '7961) luerrJeleJo 'slroddns 1etela{slBurelur se peuoqcun; deql 'res [€'I Lqcl ,rra!^al lBrrrolsru e :,{liololqoaepd luopouoJ nt r [ch. 1 Sec.1.31 The archit€cture of conodont apparatuses t) b'rra-- Lindstrom (1964, arguedthat the peelingback of secretorytissue whole association.Hinde did not figure the -:.:: E: rhev might have from even simple conodont elements would assemblage,but useddiscrete specimens from E Tr: 3 tbod-gathering present a spaceproblem, a point previously various localitiesand horizonsto illustrate the r { -:,:Sirdm's hypothesis made by Nicoll (1977). This was counteredby different forms incorporatedin his conceptof f i:E r:::,rlont affinity will Bengtson(1980), who suggestedthat 'space the species.Huddle (1972) re-studiedHinde's problems' could equally be used to prove the material and published photographs of two Ahrt-5 --:alishred by Hass impossibilityof eversibleproboscides or of the conodont-covered shale surfaces from the |cE ---t. rhe idea that retractableclaws of cats. If this debateproved deposited collection. Huddle thought that I :Lrlu-::::d as teeth or one thing, it wasthat there was still no general these were two separateassemblages, but it l rF-L
'CZ 8!f'SZ ld'tn6l) srcg nO {q peterlsnllr ,{llpu€uo srouIII ,//€9_x .ou ueunad5 ygx ,s1lsd Jo 'srourlll ',{lunoJ eltes E-I ,{elrcgJo snorelruoqreJ6q} uorJ eaelquressv^lrsre^run iueja aurppecv11.B!J
l1
a
[g I ras I'qJ] FJJrolslq e :,{Solorqoeepd luopouoJ ^re!^or 9I [ch. 1 Sec.1.31 The architectureof conodontapparatuses t7
a I
r: ::
L.:: -:l,--. L,rnors. 6 -r : _::::1). Pb
: :: O -: -arkodina, L ;: l;r.rnj21 6f 1le E6:r .: -r _l shows a lErE:-_:_: _:CkS the Sd Ia ':t:: =\-mblages are erL- \ :-r1l 19?7. Puch- Po fu-.:-,.. Fers.comm.) E iir,:: - 9S0). but the FtrJ-a--:::e lrom car- Fig. 1.3-Diagrammatic representationof the multielementcomposition of a Carboniferouspolygnatha- rEE€:: ::: . ilered to the ceanassemblage, modified from Norby (1976, p. 103). :fr--::-: l\ a le$ of the mrr:; ::::pratations of atusesof single individuals that have become 1980, Nowlan 1979, Pollock 1969, Lange E -L:: :-:a::Lrn har e relied fused in their original alignments.Although 1968,Nicoll 1982,Austin and Rhodes1969, I -a:iti i:: l:t:. most clustersgive information on the elemen- Behnken1975. Ramovs 1978). F :: :::---ELrntelements tal compositionsof apparatuses,they are gen- It is important to note that conodontassem- i:r: : :---.i:r:. in which erally lessvaluable than bedding plane assem- blages,both on beddingplanes and as clusters, Etfl:: :- I : :: IOe Sediment blages in helping us to reconstructthe three- are rare. Only a few hundredof eachof the two =::. . ,-:' le I 'cas Lqtl lBqJolsq e :,{ilolorqoaeled ^ra!^ar luopouoJ 8I The architecture of conodont apparatuses 19 ' [ch. 1 Jec. r.Jl _:I-.:]om\. EE;: :. : where essentialpoints of his reconstructionare: direction from the remainder.Scott's diagram ir : -::: :'. :ence of the also shows the denticulated surfacesof pec- (a) All elements are longitudinally aligned, c : and fot tiniform and ramiform elementsfrom the same ';<:.::- with the apparatusbilaterally symmetrical G :rE _r :- iie baSisfor side of the apparatus facing in the same about a vertical, longitudinal plane. lE-:;--:: ::: Of (inward) direction. In his restudy of North -rleration (b) The arrangementis linear and elongate Et- American assemblages,Rhodes (1952) revised with the paired platform elementsat one itst--:: :. --.*on (1934) Scott's plan and produced a diagrammatic end and the ramiform group at the other; : Ir.L-::-: ::: :nent orien- arrangementin which the elements form an the intermediate angulate pectiniform lf 1]r -:-- =: :hat lime tO elongate,linear series,as in Schmidt'smodel. elementsare placedclose to the platforms. d r": .-- However, Rhodes' arrangement (Fig. 1.5C) --= "i;hitecture. (c) The distal ends of the ramiform elements m==:l; :: :Ol\ gnatha- shows all the elementsin the reverserelative -:; (conventionally posterior) point in the *r ;. :: : nLrle regular orientation from that shown by Schmidt, and same direction as the cuspsof the Pb ele- bn --::-= :: ihe construc- he additionally illustrated the dolabrate ele- -': ments and the platform ends of the Pa diagram of ments as the closestrarniforms to the axis of elements(also conventionallypostedor). Fn--.---r:=--:--:.i ::;1.-<.{). The symmetryrather than the furthest from it. This (d) The dolabrate (M) elements are interpretation of the apparatus pattem was positioned on the flanks of the arched probably strongly influencedby a single well- group of ramiform elements preservedassemblage collected from the Car- (e) The denticulated surfaces of the pec- boniferousof Illinois by Du Bois (1943, pl.25, face in the opposite tiniform elements fig. 14; refiguredby Rhodes 1952, pl. 126, direction to the denticulated surfacesof fig. 11).This specimen is illustratedin Fig.1.6, the rarniform elements. where the general pattern put forward by In the absenceof soft tissueassociated with Rhodes is apparent, although the ramiform the assemblages,there was,of course,no direct element group is somewhatdisrupted. Rhodes evidence for the antero-posterior or dorso- did not directly attribute an anterior and a ventral orientation of the skeleton. Schmidt posterior to the apparatus,but the inference (1934), however,regarded the elementsto may be taken from his usageof conventional representthe mandibles,hyoid arch elements, orientation in the descriptionsof component and gill rakersof primitive placodermfish, with elementsthat the platform elementsare at the the platform elements, therefore, at the anterior. In a later contribution, Rhodes anterior of the apparatus.A consequenceof (1962) commentedon the symmetryof cono- this view is that all elements were actually dont elementsin assemblages,and conclud€d oriented in the reversedirection to that con- that it suggestedlateral oppositionof elements ventionally applied in descriptivework. It is (as left and right forms) rather than an also implicit from Schmidt's diagram that he upper-lower jaw-like structure. considered the denticulated surfaces of the Although Rhodes(1952) stressedthat the pectiniform elementsto face dorsally. relative positions of components in his The discovery of additional assemblages apparatus illustrations were purely diagram- from Montana led Scott (7942) ro propose a matic, his putative arrangementhas certainly very different model. He felt that there was influenced several authors' interpretations of insufficientevidence to draw a wholly accurate how the conodont skeleton might have oper- picture, but produced the schematicarrange- ated.This may partly be a result of the refigur- ment shown in Fig. 1.5B. The apparatusis ing of his diagramsin widely consultedpub- again interpreted as bilaterally symmetrical, lications,including both editionsof the Treatise but Scott placed the pectiniform elements on InvertebrqtePaleontology (Rhodes 1962' directly alongside the ramiform group and Rhodes and Austin 1981). Pdor to Rhodes' oriented the platform elementsin the opposing work, however,Du Bois (1943, p. 158), had god 'slueuele- pazlyssoye stueser uuoJlrllpJ aql Jo ror:a1sod eql o1 peuorlrsod ry snoaceuoqftcuaoJq ara.u,{aql l"rJelptu srg ur parnrJo Fepuq ? Jo uoq?rJoss€ slueurala ru:o31e1daraq,r l?ql palou .{p{ 6I) slos n( pue 'sepqs -Jess?^\[exegl uro+ sJelsnlJueruoaaq raddq pue seqcled ,(9961) mrsse-rdurr Jo^requnue Surqucsapur a8uel ruetn passe aueld-Surppaqsno -aiueJJ? pelonJlsuoJeJ.sepo{ll ,{cetnrJ? ,JeAeA{oqJo qord a,req teql speq eql agl lnoq" lqnop euros .s?^\elaqJ IXr qth\ peleltoss?anss[ 'qs$ pup sluopolaqlJo salJrluaplpurep aql ol ol aoeld snol^qo euo gll^r 'Adoloruotlp sdeq.radpue.i(BoJeuB up pJsru -doJer 'ls?rluoJ ul .(gL6l) :lrzQ .sluopouoJ ]VI IINY aql JoJ laporrJe,rrleruduroc aq1 pepraord ;.iO TUNJVN flHJ, lsaq sql?uaoleeqc,eql s€^i ''I lErlt uorsnlcuocs.laqcs -laD{ sql?uSoleegc;o sesnlereddeEurdser8 aqt .selaBqJ{lod pue Jo s,ref aql .spodollspa 'sJ"af,lueceJ ul rc 1o ged luelrodul ue .rT qf,JPesE pu€ 'slsuJlu? ds aralduroc Jo ,fta^oJslp r pelset,{lsnolJes eq pFot I q{q,$ uI ,(e.t ,(Iuo eqa FlduoJ aql Jo eJueplae Suarnalapel"losr Jo uoll? up peq leqr sasaqloddq E eqJ uo[cunJ pu? rb ;o suorlelerdrelur 3u1 raoala luopouoc;o se8elq Tnts 'enssD$os pa^res I q 'reqr luel?dds sl u 'dnorE urro;ruer E tJSsluauale elsrqslop relrurts,{lpeorq -aSueuzIorN'qsrJ pltsluoeBled qosl 'snlo1n8unmpo1ng I uI ',{ll?Jtuecu?ql Jeqtel i s 'sntereddzaql Jo apts rEr$ stuatxele hI eql leql : sa3elqurasseuro{ eJuap , (60I d '116I) uossddel gteld eq1pue rorrelue eq1 ,urols,{s rrJluer eql lEql peldeJJe -pur.I e{rJun sql u1 .ueEro,{:olzrrdser Ha1 ,{q pagoddns se,t e pue derl pooJ ? s€ pa^Jes e^"q plnoc rld Jo l€ql qlla u"ql uoD IIcIqr{ enss4 polslllJ Illl.,lirpera,roc ara,r dnOrE 5 qll.t lUeUrearEeur eroru trrJopruet eql Jo sreq peleyncrluepele8uole eql [ept^a eql pue 'slueuale slueualJuJloJrurer aql.;o sdsnr aql ,{qlno u pur5uo aq11o uorlecrpur pJuJ?Jbur^Jrs fuepuoJas qll^1 .sluau,lJla(qd) {aqr'otlgordoc,{lqeqord epplq peqJteaql ,{q peuroJlad se^\ uorlJunJ + reql tlal eq q8noqrly tur^els{teuud V puelorJJlue aqt ?ursop pue 'oes 'qc] [t'r I lBJrJoNrq E :,{€olotqoaplBd ^rar^ar luopouoC 0z F** 27 [Ch. 1 Sec.1.41 The nature of the conodont animal a:r:-:r end. A prirnary sieving Although he felt that these clusterswere all some worm-like creature'. The structuresthat :n,r-_:d by the arched blade probably coprolitic, they should retain some Du Bois interpretedas parapodiaor cirri are' (Du rtr:: i=_-Jndary sieving carried indicationof the odginal relative positioningof however, not clear from his illustration rc :e :rriJorm elements. The elements,and the evidencethey provided was Bois 1943, pI.25, fig. 16), nor are similar de!*: ---s of the ramiform more in agreementwith schmidt's reconstruc- impressions associatedwith complete, well- t=: r -_: Jliated tissue which tion than with that of Rhodes.This conclusion preserved assemblages.Those assemblages film are o . iood trap and a was supported by Jeppsson (1971) who which are surroundedby a brownish to tr- -,: :- :r.:em. unlike Lind- acceptedthat the ramiform elementswere at commonlydisorganized, and are most likely t :=r----::r posterior. coprolitic or similar origin. The asphaltic - elements were tbe anteriorand the Platformsat the be of hcf,ls]-L: by Scott (1969) from the :<- '*'hile the func_ Jeppsson(7971, p.109) also noted that evi- 'blebs' described J r:e Pi Pt. ald. perhaps, the dencefrom assemblagesand clustersindicated Heath Shaleof Montana are also almost cer- E after development. that the M elementswere located on the out- tainly faecal in nature. Scott initially believed -q\..i por- u-- =-,;rhors supported the side of the apparatus,as proposedby Schmidt, that they representedthe cartilaginous but the tft- J.:odont apparatus as a rather than centrally. In his reconstructionof tions of the headsof conodontanimals' t:. Halstead 1969, Nicoll Oulodus angulatus,isolated from the gut of a elementsthey containare chaoticallyarranged, !@=-:: ro prefer a mastic_ palaeoniscid fish, Nicoll (1977) adopted a commonlybroken, and do not constltutecom- !. !-i: :-rr;-r_s function. priddle broadly similar arrangement,but with a pair of plete apparatuses.There is no evidenceof soft ry5 3 r.-.rpred a sugqestion dolabrateelements set anterior to the elongate tissue surrounding well-preserved,organized lc;rrr,,-- ;;; \fnUer 196;) that ramiform grouP, assemblagesin any of the Carboniferousblack f ' r..roe horny cusps It is apparent that, in the lack of any pre- shales. ltr::-. - --:_);.erplaining their served soft tissue, studies of natural assem- It was in Carboniferousstrata of Montana, - rrr-{ internally sec_ blagesofconodont elements led to widelydiffer- though, that the first indisputableassociations a=- n-log Priddle's proposal, ing interpretations of apparatus architecture of soft-bodied organisms and conodont ele- fca :x array of such horny and function. The conflicting functional mentswere discovered.During a routine check Erstle lin-sual structure simi_ hypothesesthat had developedfrom examin- in 1969 of a fossil fish locality in the Bear re: rl the extant myxinoids. ation of isolatedelements were not resolvedby Gulch Limestone, W. G. Melton Jr and J. J. mr::::ional arrangement of evidence of the complete skeletal structure. Horner found carbonizedimpressions of ani- m :\< flecessitatedby such a The only way in which the various proposals mals that proved on laboratory examinationto I r*i--r be accommodated in could be seriouslytested was likely to be by the contain conodont elements (Melton 1971' t-:,-: analogies of apparatus discoveryof complete specimensof conodont 7972).'lhese specimenswere shown to a rr,L-i ed by Rietschel (1973), animals,and a searchfor suchfossils has been number of conodont specialistsat the North ! &-:rngements proposed bv an important part of conodont palaeontology American Paleontological Convention held I R:.rles with thi radulae of in recent years. that year in Chicago.Among thosepresent was rr5 !.f polychaetes, and the Dr Harold Scott, who had describedthe first ces of chaetognaths. Riet_ North American bedding-planeassemblages in r r:_r that the chaetognaths 1934. Scott was deeply impressed by the 1.4 TIIE NATURE OF THE CONODONT I conparative model for the specimens,and announcedto the Convention ANIMAL ( 19-6). in contrast, recog_ that the conodontanimal had beenfound. Mel d perbaps a homoloey. wiih One obvious place to look for traces of soft ton and Scott(1973) went on to collaborateon hs of rhelodontsand-fish. tissueassociated with conodont elementsis in the publication of a full descriptionof the new Erer- some doubt about the the beds that have produced the most numer- fossils,which includedfour completeanimals. ts' reconstructed arranqe_ ous bedding-planeassemblages. Carbonaceous The Bear Gulch animalsare exquisitelypre- l). rn describing a numbe; f patches and impressionsdo occur in these served on bedding surfaces in fine-grained frsrers from the Kellwasser_ shales,and Du Bois (1943, p. 156) noted an limestones. They are elongate, around shere platform elements associationof a hindeodellid element with a 50-70 mm long by 12-15 mm high, with an terial they were positioned brown carbonaceousfilm that 'apparentlyrep- anterioropening and a Posteriorfin ( Fig.I .7). d rhe ramiform elements. resents a fossilized portion of the cuticle of In the original specimens,conodont elements .slueuale rxD pu? s33 g '€g6I eql JI 'slroddns eroqdoqdol se e^{ se uoqrel paqrrsepun u? uI €d rnoJlseel le 3ur l sqt ot petelerdlesolc sl -ord a,rrssedsz Sur,rressnql 'sp-rz,n1noSuqurod -pnlJur sluauele perellpf,s,(lapm flq8re re,ro 4lun sreaddeA\ou lI sereJJnspelelnc[uap rlegl qlliR qlnoul eql peqquepr eq qcq,r q '0909 hln eldruexeroJ 'e^Isnlcuo3s€ pelJJrcuastueruela aql 1eq1pasodord rsqpn; 'suerurcadsaruos ur sluauele;o reqrunu eErel rsueJ Jr 'Suuserelulsr IroJlspurT'EuarueleJo f8oloqdrour leuorlcun; ,{leleurprour aql palou osp ag (1919 141n f, i€-r\uoC pue s.uroJls aqt pue surallzd aSelquasseeueld-Surppaq '90EZL-9 LINJ) sraqlo1o sln8 eq1ur sursrue8ro ssepr Jo $uaEra^uos Jo uoqeraprsuocqSnorql',padeqs-rrIJor'r pue Jaqlo pu€ qsrJJo sureual aql 3o aruasardaql soPouoc erIl Jo asoql Euol ueql raqler pedzqslerrzq ua,rero 8uolqo pue suaurrcadseuros ur slueureleluopouor Io I pII€ uorlrsoduroJJreql ueeq e^?q o1 ,{1e)II1aroru, s?^{I?urue eql leql pq eq1Eursse-qs 'enbDuc aarsuoqeJdruoc f,pe1 ,srql luesaJd oJe slueuala btt'd'Vrct urorlspurl) paraprsuocaq -ncqred e pephord puz suaurrcadsu,rronl eql qr fluo sV 'Ieuuels JOA\3ra UI 'SlueurelerrrJoJrrrreJ eql Jo s]?q eql Jo lsolu peuruexe seq (5g6I) surol{ de,{'uoC ,s?1{r lltIII'ra^aa\oq'eJuapr^a se 3uo1se sour[ xls ro e^rJ,lsourle lr lpgl '(SAOt'SfOt) suro141de,nruo3 pue (9161)des I -iq pasodorduoDcnrts ecnpap ot lrrortspurT pel etspJoqcoluopouot -rrr?d pu? s?urqc1r11'sraqlo Suorue.fq paJ?qs I qtlll luaueaJSPelqs e ;o 1nB aqt olur ss?d o1 q8noue lpus s?rtl uorurdo ue 'sJelee luopouoc se sal?pJoqco qdol a1oq,neql ',{Ipuo 1l leql lrBJ eql lnq'(€L6I '7961 uo4spurl) -tuopouoJaqt pep:e8are11 .sn1zredde eleldruoc pouor uerrqure3aldrurs rarlr€a pel?o[ ueeq p€q eleroqdoqdol e se,ll ,(yreeu e;o surerxeJaql sureluoc uerurcads .aroqdoqdol o ur aruelqurasar3uo4s puru? or{t legl uo4saSSnsaq1 euo fpo l?qt pue 'pereprosrp ,(llsour ere g aqr go slroddns,{4ds eleF eluet € loJ ern1cnr1sSurgoddns eq1 se SIeIIJIUPaql ulqllrl\ sluauleleluopouoc eql leql .purrue eouof, € ;o eldruexe snlerzdde oq] po8esrlruaeq qrlqlr ur 1no pelurod (VtAt 't.t11) uortspur-J .sre{ro,!\ €uruoseel ;o sauq pd luopouoJ pcqaqtoddq e;o u8rsep aql ureJls luopouoc [q uorldecarpexrur p papJocreetaa Ilns osle all el,{qdater -uoc o seSeqdoluopouocaql posn pu€ .sql deqt lnq '1serelurpcrSololuoaepd alqereprs .,{8o1oca qdoqdol aqt qll^\ elq" pezuSocor(7461) rrrortspur.I pue i(3o -uo3Jo {lpelqnopun eJa,{ sarJe^oJsrpeseql I qlnour ? PapunoJJns lorqoeepd luopouoc lnoqe soD€uuoJulauos '?lPpJoqcotuopouoJeql t JBlnsslual 3 S€ aJnl qlrn fllcerlpur sn apur.o-rdop faql'sluopouoc 'unl. qdqnsaeu ol s1errlru?,reu eql peu?rsse " E |m aH selcetueteseql lou aJa^iselppror.lJoluopouoJ :qr qBnoqtly llocs pue uollew puE 'J€alJunelea seleploqJ lItreJ eql eJea paaarleq 'se8eqdoluopouocere^r slprrru? raqlo o1sdlqsuo4?leJ aql etepJoqJEuruurlns s sz-07 Jo spFotx esaql leql salecrpurflEurwleq^iJelo af,ueprle -ea{ e s? paprcSer faql qr1q,n 'leruru" ssel F 'selsPlual pel'ellel peurquoJ eql puB 'se8plquasseoueld-Surppeq -,r\efsrql urqtr,rr\uollueydol{qd lallrJ ol pelJas iq pelerd:a1ur era,r ruorl sasnle-redde luopouoo 3o Surpuels snlereddetuopouor eql 1eq1pe,ra11eq (9461) r-iy-rz1n8ar'uedralunoo -Jepunrno qlrl\ eJu?ue^le l(lJ€elJsr srql sezrs llocs pue uolletr{ 'luesq? ere ,{aq1 (6699 Ee dool aq1;o semteel snorJEAJo slueuraleed gI lseall€ pelunoceAeq I{n:€t6I ltors aas'999€8I WNSn) ret€l rror pu" lred qloq fq I r{Jrr{a\ur slueuela 1o seqcledo,trl surcluoc 1n3 palJelloc suarurcedso^{l lseel 1e ur qEnoqlp '(g'f '8rd) pue tu-o eqr eql 'uosrpBw 'ursuoJsrl\ Jo ,{lrsJe^ru0eql l? '(€t6I Uo"S pue uo11eytr) .uoteluoellep, p:doo1 e qtr^\ 'Ieurue plaq fpuarrnc prreVy6Z,L perequmu uarurceds arlt paurat 'tn8 J?ln8uelJl aql uqll,{r Jmco {s -i1e:a1e1qe srsuro1,1 ltSotuopg erqunloC ng usrrquEc elpprw .(886I) surow ,{ea\uo) pue (Z/-6I) uollal{ ,{q suope4snll uro+ pelrdrxocielnossrw ,euEluol{l ,lZOg lun e yo srseqeql uo Jo,{lrsre^rufl Idn ou ueulrceds Jo nur^rsrc zx euetuol l euotsarllrr qcln9 leeg eql rllo{ 'BrJ riq pa1:oddnssea\ sal?-r Iertuac]o .elBproqcoluopouoJ,v_l r noc reql srseqloddqeql I sleroqoJrocol enssrt rgnoJ Jo 'se^{ araql reqssdnorS om1 aqt teql ,r ar3ql teql pa1sa88ns ryodorqcerqaleyncrlreut ciro luepne ere eJaql se.\r leuru€ luopouoJ [t'I tes I'qJ] eJ FJlJoNlq e :,{SolorqoaBFd luopouoJ ^ral zz [ch. 1 Sec.1.41 The nature of the conodont animal conodont animal was a lophophorate, then the fossil, although interesting,has shed little there are evident organizationalsimilarities to real light on the nature of the Conodonta. inarticulatebrachiopods,andLindstriim (1973) More recently, though, discoverieshave been suggestedthat there was a distinct possibility made that have revealedthe body plans of at that the two groupsshared a commonancestor. least some conodont animals, and have pro- There was, of course, no evidence of soft vided the first firm foundationsfor discussions tissue to corroborate Lindstrdm's model, but of their zoologicalaffinities. the hypothesisthat conodontswere lophopho- As part of an investigationof Lower Car- rateswas supportedby Conway Morris (1976) boniferous shrimp beds in Scotland,Dr Euan on the basis of a unique specimenfrom the Clarksonin early 1982 re-examinedlarge col- Middle Cambrian Burgess Shale of British lections of the Granton shrimp bed of Edin- Columbia. Odontogriphusomalw Conway burgh that are housedat the British Geological Morris is a bilaterallysymmetrical, compressed Survey in Edinburgh. Among the material -:-:: a rnd currently held animal, with a looped structure towards one were two slabs, part and counterpart, that { E =c'::sm. -Vadison, the end (Fig. 1.8). The singlefossil is represented preserved an elongate worm-like organism tfo*: :t:-:ments in which I by both part and counterpart, with different with conodontelements in the head region. Dr I :: a: :t-uents of various featuresof the loop apparenton each. On the Clarkson showed the specimento Dr Derek ,t r -t--,.:*_-e*ith our under- counterpart,regularly-spaced depressed areas Briggsand myself,and we wereboth convinced n-E: ::lparatu ses from were interpreted by Conway Morris as that this, at last, was truly a conodont animal. lrtr::- ?nd the combined retracted tentacles, while on the part are The specimenis 40.5 mm long and mostly less =.iicares that these moulds of 20-25 small spikes which he than 1.8 mm across,with a slightly expanded -ir'gl ln-^-o: ' ges- believedwere the remainsof rigid supportsfor head region comprisingtwo lobate structures li,::-.cirordates were not thesetentacles. He interpretedthe whole struc- flanking a central lumen (Fig. 1.9). The con- r jr:r -[a qr indirectly with ture as a tentacular feeding apparatus that odont apparatus lies behind this, with the atlr: --oDodonr palaeobiol- surroundeda mouth in life and was compar- ramiform group to the anterior and the Pa rh.-:r t 1971) recognized able with the lophophoresof other lophopho- elements at the posterior; the paired pec- cE'.r:.-D rophages to con- rate phyla. He also suggest€d,from two princi- tiniform elements are oriented transverseto f a :;:rtthetical conodont pal lines of reasoning,that O. omalus is an the long axisof the animal,while the internally E eTr -r€!ed the apparatus example of a conodont animal. Firstly, the aligned ramiform set are at a high oblique trr-;e ior a tentaculate spiky supportsof the feeding apparatusbear a angle.This arrangementof elementsis similar rE:rioD that the animal strongresemblance in outline and sizeto some to that shown by many bedding-planeassem- : fu!: :Een floated earlier simpleCambrian conodont elements, and, sec- blages.The soft tissueof the trunk, preserved lf-: - --ut rhe fact that it ondly, the whole lophophore showsconsider- in vivianite, revealsa linear axial feature, re- b :"1\i into the gut of a able agreementwith the hypothetical recon- peatedoblique structures that areprobably seg- b: L-:istrom to deduce struction proposedby Lindstrdm (1974). The ments,and, at the posteriorend, ray-supported It= _-: s:r rimes as long as evidence,however, rnust be consideredcircum- caudal and posterior fins on one side of the ht= :-:nenLr. In view of stantial. As only the moulds of the spiky animal only. A full descriptionof the specimen, (L-r-r.:.im 1911. p. 7 40) elements are preserved, we cannot test if assignedto Clydagnathus?cf. cavusformh,$tas tu-.i _f,el\ to have been their compositionand histologycompare with given by Briggs et al. (7983), who recognized d-s:-=: lather than long those of the conodont group, and, while the possiblesimilarities to two extant groups,the Ift:-:',::,r-rnsideration of convergenceof ideas represented by Lind- chaetognathsand the chordates. iiEE- :inerns and the strom's and Conway Morris's reconstructions Chaetognathsare elongate,dorso-ventrally 6 :i :_eleur. Litdstrdm is interesting, it certainly cannot be regarded flattened worms with lateral fins (Fig. 1.10A). I ::€ encircled as conclusive. Their generalfeatures match well with thoseof rr *::;ulated--iaeDr surfaces It now appearsunlikely that Odontogriphus the first Granton conodont animal if the latter k =-::g as passive pro- is closelyrelated to the conodonts(Briggs et al. is consideredto be compactedin dorso-ventral F:rc,--:: :-upports. If the 1983,Briggs and ConwayMorris 1986),and aspect.The axial tracecould then representthe I UPql Xelduloc aJor[ 'luopouor e [1aurnua8sr uaulcads purSrro eql t3o1oluoa1o4 awtqatntul uo asqDaq eqt ul F aerqt Ip puE 'raqlo t"ql slqnop alqrssodduu ladsrp o1 pe,r:aspue las Peeleql ,{rolloJo1 pealsur asoqr all '[Iur;;e lmr eqr Suop raqgny uorgod rogalue aq1 ur sasnleredd?luopouoc luopouoc ;o uralqord eql ol uo[nlos /fio1ceJ crp$ qroq uo s,{e-rur; pel?rJosse,{eldsrp o,r1 Jerllo eql lnq 'pe^res -sr1ese papr,rordIepou al€pJoqoeql Jou qluu rmnads eu6 'selnuos -ard sr prurue ue ;o ged rouelsod eq1 dpo -8ole"rIJeql Jeqlrau'aurl wql 1€'leqr apnlt rEalJ qlr,r\'pelueurEes suenrrcadseseqt Jo euo ug'(9961)'p n -uor or (9361) 1z1a s33ug pel prnue uoluerg Ip leqt unJuoJ relncu aSprrply fq pequcsappue paredardpue ryu13 eql ,{q pe,{eldsp seJnpn4s os eql Jo aJnleu pd-4os ;o yelep puorl ye51dq papeloc (ff f '8tS) suaurnedsarou aql SurpJeBeJserlur€Uerun qll,$ peulqluot mrads ,rau eql Jo IIV eerql;o ,fie,rocslpeq1 dq 7961 ur papr€aer ser\ slueuela luopouoJ Jo eJnleu anbrun aq1 sFLL 'pelunou sel\ peq duruqs uolu€JC eql Jo 'leportralEproqJ rn IIea{pelsporxurocre " qrJpes snonplssBup pup'eJuepl^e leuoqlppE 3q snql u"r enssll lJos eql Jo seJnl?eJpaNes D lql -v :q8rnqurpg ,(q pe,rloser,{qny aq dpo plnoc uopelardralur -erd aqa 'proqcolou eql s? qJns aJnlBeJreeuq ItEDp[tJ 6 I 8tI ur serueJaJlp e$ql lBql JBelc se,t\ lI Jerllou? ro 'ln8 aql Jo 1e,r e lueserder ppoc '(986I) rFq ecell I€xE aql pu"'snxolqdue ur saurolo,{ur dq pelueserdsE^A setepJorlc se sluopouoc 8ur psdeqs-4 eql qll,lr sno8oloruoq aq fqereqt -prz8e.rrog eseca,rrsueqardruoc ,{peprnred y lq8rur uorlelueru?as eqJ 'l€sJop srrrJeql qllt\ 'saurolsoqleuEaql ol dlqsuorlzyeresoyc e ?ursod 'pedse l?rel?l ur pspeduroc aq o1 pasoddnssr -ord dq8ql qtr,r\ 'sal€rqauaa ol saJu€lqruasal luopouos erlt Jo lunJl eql JI es?rsql q luePrae aql sserlso1 perra;erd (ggOt ,(qBIu'€86I rel^ ele sertrr?lurrs eqJ (S0I'I'Erg) sprourxdur -ue1)s-raqlg ',duroleuuqleuSota?qc ot spedse pu?'s,{aJdrx€l'(DutotsonlJuotg)snxorqdure ,{ueru os ur suro;uoJ Ierrnue luopouoJ /i\au eql s" qons'selepJoqc aldurrs q11,,,ruosFedruoc leql eJueplJuloJe ueql JJouraq fetu ll, leql el$ul osle purrus uolusrc eql Jo stJedsv pareprsuocpuB 'sqleuEolapqrpu€ stuopouoc 'luopouot eql Jo ;o sesnlerzddeSurpae; eq1 uee^\leq ,{Soleue {unJt aql Suop lueppra uorlelueru8asenbgqo esolce penSre(26961) uosfuag ernt?u sll ol aql qlra peredruoJdlrpEar eq uer leql seJnl se possardxaere,!\ sararl lueJeJl:rp,fiea 'Jeulru€ -culs ou lqrqxe sqleuSolaeqc1eq1 sr 'q?noqt luopouoc 1r\eueql Jo &Jrpueqlns eqt peldem? 'f1ruq;e qleuSolaeqce 8ur1sa??nsq1r,t urelqord lsour elrqA\ ',fue/iros tou aJa,t\rtoqtnn euo 'pooq elq?F€rler e dq tsar tz peraloc .ro11tlo, ere pu? qlnou eql Iu"U 1eq1saurds Surdser? -srp eJe,{ slred lyos pe,r:asardq1r.t suerurceds snorrr1rqcJo stes ssessodosp sqleuSolaeqc ororu Ulun '?luopouoC eqt'urn1,{qd eleredes alrq^{ 'sluaulruduroJ Izralpl olul IlIoleor IrEl B ol sluopouocu8rsse ol pue (T86I lleD) pu" IUn4 eql sephrp 1eq1,{relueseur uerpeur 'uo$slturedqlr^i'(9!6I) surol f€.{uoJ urorJuaerpeU leturueeteldtlloc Jo uot.nlsuocar 'g :J O 'uot8uqs€r{ 'drolsrH lemteN Jo unasnw puoqeN Sn '69196I WNSn ou ueuqcads'ued eql uo snlp&dde Sulpeo] eqt'V:su.roll Ael|.uo:.j-snpwo sm!&rSotLtopo-B I'Rlc [t'I res rqcl ,iaa!^er |eJlrolslq 8 :/t8ololqoaE|Bd luopouoJ nz :t lch. 1 Sec.1.41 The nature of the conodont animal z) Tt dII: : =:::n- spec'men !. - -_ r :a.'onstructronof a re1..: r: E :: .ri igl conodonts to a r -:'. C.rnodonta, until more F=-:-..i soft parts were dis- ;!rr:. -,-r::os'ary. While most lr-:;::r of the new conodont k-=:: -.-3\rs g'ere expressedas 4:-r:r 19S3a)argued a close r --:: :::dins apparatusesof dl-- : r--r eths. and considered rE::- --:.3na coincidencethat I conformsin so many u;-:-:: r-::tomy'. Others (Jan- t : : !-: , rreferred to stressthe r r.:::::3te.s. with Rigby pro- t=,:-.rf to rhegnathostomes. m,-::::sit e casefor regard- x :i':-:es s as presentedby r --::: -ie:e differences in Fig. L.9-ClJdognarhus? cf. cavusformisRhodes, Austin and Druce, from the Granton shrimp bed, d: 'ce fully resolvedby Edinburgh:A, the completespecimen, x7, IGSE 13821 (pafi), British GeologicalSurvey, Edinburgh; B, - -r head region of counrerpart,xl9, IGSE 13822 Er= :-: an assiduoussearch rft::: --edqas mounted.This r 1:-!- :'; ae discoveryof three All of the new specimens provide some addi- but still equivocal in nature. The new data r,i-g I 111.collected bY Neil tional detail of soft-part structures, and in par- were regardedby Aldridge etal. (1.986)as re: ;eicribed by Aldridge ticular confirm that the trunk of the animal is sufficient to negate the possibility of chaetog- G -:-,e ..f rhese specimens segmented, with clear evidence of V-shaped nath affinity, and, although still in need of fu :::: ..t an anirnal is pre- somites. One specimen shows closely-spaced some clarification, appearedconsistent with a r !'f::.: display associated fin rays on both sides of the tail, extending chordate model. The most attractive E-\- r:-o rhe anterior Portion further along the trunk on one side than the hypothesiscurrently is that the conodontsrep- jawless EE '-i. trossibledoubts that other, and all three sbow axial lines, rather resent a separategroup of craniates dr<-- : eenuinelya conodont. more complex than in the first specimen (Aldridge and Briggs 7986, Aldridge etal. rsab;ns (986I) sEEug 'tusru€qcetrrEurraqle8 '(2961 'P n eEPttPIV)aumlo^ 'uooJunJ ur pel?rlueJeJ -pooJpaJeloc-enss[ B se pel€Jedoa^eq plnot srql ;o 7 reldeq3 ur toJ PonEr?requnJ sr pue Ir:l: asaql lPql ,tlalllun snle-reddeeqt teql pePueluot llps sr ll eurnlo^ '(g3et) l11"qpu€'(S86I) rea^{S'(t86I) uoss IruoJ sseJoJd ur ,{3ue slql Jo (1861 11ocrp)9 rerdeq3 pue (1361 peor -ddaf pue e8prrply'(egg61)uosfueg 'sreqlo -qF sarJedsqcee Jo asoql -xag pue gocr51)g raldeq3 ur 'ra,raa\ol{'enssq SuorIIp',{q pesropues€A\,t\er^ sIqI e^IlsuJelle rr]]ruej Jo eft 'suorlrsod gos 1o SurraaorpJlElnlsod aql ,{q peqslululp elqrpeJJeJoru aql pepr^oJdsluauele IrrJoJIuIl lno;rsse esoql Surpnlcur eq pFo/t\ pooJ Supnurulurocur ,buerJlJleraql -cad :orrelsod eq1,(q pessecordeq o1 ,{erd 8ur lera JoJ 'eece opJeqqrH 13r{1pu€ 'e^ars e qcns dq pernldec eq ppo,n -dser8sluaruale ruJoIruEJ JorJalue eql qlh\ uel ioqdrolll lueraJJp ,{ra^ lEql selclu€d pooJ eql ol uolleler uI lsnqoJ -sfs pale-r8aluru€ l?qt pereprsuoc(886 'p P D -\ElU qJrqA\ saJnpnrls pue e8rel raqt€r ere stueuela qd pu€ €d aql sa8rrg'ue3-ro f,rolerrdsar/SutpealJElnJeluel dde o1 peqdde eq tq8rur l"ql peluerrrlxoc(9361) s33ug pu€ eSpuplv e palroddns feql t"ql Jo qlee] se pelrlroFod Far4IP eql ,!\oq raPrs 'lnE eql olur drlue o] roud sluaurelerlJoJIuIl ,{aql l?ql urerrrerseq4qrssod o,rrl eq1, uop I'ra-!a,roH'sa8elqurasse -ced rorrelsod eql ,{q peqsnJc ,{pue8 uaqt -JUnJluell.lals luopouoc;o uorlsanb eql J^losJl petuasarde-r,(11uepunqe s",$ r.lJrq,rrallptx pooJ:lulnctued l:os ol a,lats o1 i{lrurgeluopouot Jo uopdecradrno esn lou ! qJrI^{ 'exel ueetBql?u E se p€ plno^\'enssD pelellt ql4\ Pela^or -u€J enr'sernlJnJls elqeredurooi(llcertp f,ue q s:snleredde luopouoc 'slueluala uroJrurEJ Jo dnolS Jouelue eql Jo {r3l eql u1 'prcrpedns aq dpo f,uru sepuel sno EsnJSrp eqt Jo qJntrN 1eq1peSesr,rua aH IEIIrru€eqt Jo peeq eql ur -ltuls eql puz sa1e1d Surgoddns esuduroJ e,roof poo; I8rtue^ e Suop pa8uele ,{1.reeurl -urEepuecuo pelunou sdsnc{uroq eJea\slusrlala eql qJlql\ uI B esodold zurxtw ,rlleel' e$ lnq '(986I 'p p aaP\rPw r! aqI Iepour Jo Jqr qll,!\ lJqleSol 1111261'nL6I e1ppu4) elqereduroc tF:TPI;PIO-I1 I 3Ig ol 'ueulJ3dsuoluelo ISJIJ '986I 'srelsnlc u€ruoleq Jedd1 pea:aserd ,{1n; ,{ypzorq sr qsgBeq yo arnpruls pn8uq aq1 -uneequorJ acuepr,tepa,{oldure (986I) IIorlN 'snlereddeluopouoJne aql ql!r\ pozlSototuoq 'relncqred q '(S86I) locrg ,{q paSm ,(13uo4s aq acueprJuocelnlosqE qll^\ uD3 l€q] eJnl IIIls sr eepl goddns-anssrl e^4€uJeu? aql -cnlls Flela{s € ssessodselEploqJ u/aou)l oN pu€ 'uol"lardretur leuorpunJ slql dq PeJub '(Zf 'f 'AlS) sProurx,(ur -uoJeJE srJlJo^\ luopouoJ ll3 lou'la^a^loH oql ot drqsuogslerauos qth{ sdeqrad'(986I urePoure urElSep'V--OI 8lC IerelEI'proul).{ur uepour e Jo srEr8elp'g :i '/hel^IEslop 'qleu8ol€eqc Jo I llllsou V llEl lo pua tou eJuE snue uPlpaul Dr::co I oullsalur / uu te.ratelI soulos/ JO!i alsod 6uldser0 [t'I ces Lqcl ,nal^a.I lBtlJofslq E :,{3olo!qoeBpd luopouoJ 9Z 27 [ch. 1 Sec.1.41 The nature of the conodont animal trr ta -- t mould of f,TE: A- opporotus -ESenlery :=,if in B l n,:.:.--: l!\]noid. lateral I :, r:';--et rr otkets ate con- tSr : nlerPretation, and b-;-:-:gcn idea is still \-c: 19!5). in particular. pk"":: :r-,,ience from beauti- L:--. Delonian clusters, Fis'l1l-DiaglamsoflhfeespeclmensoftheconodontanimalfromtheGrantonshrimpbed,Edinburgh' specimen, to - r E:.r C::nton The remairls of shrimps lie across the left-hand and central sPecimens u r:,;:i rtre elements were rir.c-: = reirtral food groove prey which : .L::.,. He envisaged that Much of the discussionof the function of apparatusmay haveserved to grasp processing.A rc :: :=:niform elements, conodont apparatuseshas centred on polyg- was consumedwithout lurther of course,if tP: l::-3. \lould aCt as a nathacean taxa, which are by far the most similar interpretationcould apply, whole apparatusis presumedto haveacted srL:: . -.:.; matter which was abundantly represented group in natural the and perhaps in other ||r: :; ::e postenor Pec- assemblages.However! rt rs pertlnent to con- as a sieve. It lcriodus, apparatus t:r r::r snrn into the gut. sider how the different functional models members of the Icriodontidae, the and a :]r, commented that might be applied to apparatusesof different comprisesa pair of scaphatePa elements Es may have been k€: --i iather large and structures which may comprise elements of suite of cones,of which there than 140 in each individual (Nicoll r:: --: -: -.rrd Particles that very different morphological styles. In the more each of rd :-. .-:: a sie\e, and that Hibbardellacea,for example,all the elements' 1982). Nicoll (1982) suggestedthat a single I c;r- - _ food would be including those assignedto the Pa and Pb thesecones might be homologouswith -trns and that the r lr>---::id covering of soft positions, are of ramiform morphology with denticle in other apparatuses, be functionally I,l-:::i (\icoll and Rex- ihose of eachspecies showing internal consist- Icriodus apparatuscould thus -1 genera. bci:: 5 r\icoll 1987) of this ency in processform and denticulation. It is very similarto thoseof polygnathacean morphologi- crq- :e I that the aPParatus unlikely that these elementswere widely dif- The apparatusesmost remote are E r. : tissue-covered food- ferentiated in function, and Aldridge and cally from those of the polygnathaceans entirely of coniform elements. E Briggs (1986) suggestedthat the entire thosecomprised DISA\eruszSAq pepr^ord sr uorlueluoJ srql roJ -rr{oN'696I{rollod '896I a?u?'I'196I seureg 26r uoddns pf,fololsrq Jaqund .'uoEelnJrluepl?ea '3 e) peqr:csep ueeq e^eq sretsnlJ pralas rP r:aor lErlaqlrde pesee.icur -uft?qd pedola^appeq sqt€u?ole"rp arll qSnoqtp 'sa?elqruesse poo8 go pe1 pu? rDqr IErntEu 4\fuoJna luopouocEred qcuerq p a,rqzlaraql dq paradurequaaq seq sesnleredde rlE lEure]ur SuuaJJrp aql Jo lueseJderf,eu slerurueluopuoJne riFu: ro Iepou V-€l I BrC pue luopuocered,ueql 'en4 eJo^\sluopouoJne rrrJoJruos Jo uorFunJ pue arnpelqJJs ro; d:lsecuepelelnlsod sq F 'leql pelou (S d eql Jo Supu€tsrepun '8urdser8ot uoplppe ur 'q9961) uoslSuag pu€ 'sqteuSota?qr1o seutds uorlcun; Euueaqsro Euulnc e pau.royed eaeq ?urds?J8 aql aJ€ slueruele luopouocoloJd ,{eru esaqt teqt (986I) s33rlg pue a8puply luopouocnl reql ,{le{I tl lleJ (S86I laa.tg 'Ea) sreqrg fq pslseE8nss€^{ lr pu€ 'a"lsoJ Jo sleel 'f14uapr;o ;oord lou q?noqtp 'o/rl.taql ueeau 'sa8pe dreqs eleq sluetrreleluopouoJ rrrJoJ -aq drqsuorlele-r€ Jo eJueprle se srql pap.reSe-r -ruocatrtos '(gAOt {lzC '986I 'S86I lu€8,{r( oq^\'(€86I'286 I ) r{s^\Eruezs,{qpeluaurncop pu" {lzq '986I 'lo p a8pr.rply) sqleuSolazqc ser'r sqleuSoleeqclu?lxe Jo seurds Surdser8 ;o sesnlz:edde Surdser8 aql ur senSoleue :qt pup sluaurJJaluopouocolord uee,tlaq IeuorlrunJ lsesols Jreql a^eq ol peJeprsuoc ,{1r.re1rrursprnpnrls pue lzrrSoloqdroru3ur4r1s ueeqe,req pue ,{.rolsrqtuopouorne Jo ged ,{pza v esn ur se,n snleredde eql ueq^\ pesodxe eql ur flrsJonrp alqeJeprsuo3pe/(oqs eseqJ 3q plnoJ sluopouocne Jo u^\oJc el{] qolq/$ uro.r;'s1e1cod prleqlrda olur sluetllele aql 'pajeplsuoc srelcPreqc eql uolssncslP uoqo"Jlar e,rlo,rur01 peunserd s€a\ Jo Jo lenper8 IInJ e pepnord oq^\'(986I) p p eBpEplV reue :seterueD ecuenbasslql (€I I'3rg) e1elsluopouocetd reqto pu? etuopouoJ eqt uea^{laq sdrqsuoqpler eqt Jo elerpeuuelurue qSnoJqlpadole^ep ,{eql palse8 saseqlod,{q alqrssod oal8ultErNnl[ ulel8opelJ ZI I SIJ -8ns eq qcrq,n ruo{ 'sluopouocolord sq1 ut ElPr!olsoqlpu9 ,{e1{.r1s:rue reql leql pesodord(qt86l '91.61) rc9.rlsoalso uoslSueg 'uerJqrrr"Jal€l eql ur pareedde1s:r1 eptdsPateOn3 'f,poq pseq e pue ua\oJrpelercas-,(1p8n;u1uec e Jo Surlsrsuoc 'sluelxele luopouocng PplosBuv SJNOCONOJ JO NOIJN'IOAS (INY NICIUO gHI S'I 'padole,repsr sasnleredde Pp!luoz^urorlod prluoporepued;o ernlJalrr{3Jeaql Jo uoISSmsIp ereqa{'euInlol srqt ;o 9;ardeq3 ul (186I) "'12 la qtnug ,{q peqrJJseppu€ palerlsn requn} sr lpruruuBqseInB A :q1 'sasntereddeuro; -ruor Jo uorpunJ pue luauaSue:re eql uo lq6ll rceJlsorolaH elqsJeprsuoJs,no-rq1 uaurceds eql'pe^reseJd dpcayad sr lprxru? srql 1o snleredde p1e1a1s aql Jou anssrl tlos eql r:qlrtu qSnoqtly 'ursuorsrl\ 'eqselnel[ ut a8e nlurrs ;o luopouorolord elens ruo:; snue8eues aql Jo l?txrueluopouoc eluopouoc e pegoder (q'e9961) 1zra cqn4ryq 'ecuegodrut relnc4.red ]o 'pue 'eureDlll 1o ,fte,ropue11 eqt tlro{ s-trponpuodlo e8elquasseeleldruoc tsourle ue patuaurn3op(9961) tueEdrq pue Irzd ',{ltuerererol,,g'(zg6I e8puply'6461 ue1 eap!oulx^n t! I Jd) 1 r 'qcl ,aaFar lBqJolslq e :,(Bololqoaepd luopouoc 8Z ai [Ch. 1 The origin and eyolution of conodonts 29 !E -. - \l.rre recently, Dzik (1987) in Chapter 2 of this volume. How- l\- : ': -:r;nted an almost ever, an evolutionary link between proto- fut: : ?:,:,;erodrs from the conodonts and paraconodonts remains to be L-r-= : =:d. of particular conclusively established, although there is ft:.:,.. 1y:-ia.b)reported a strong morphological, histological, and strati- sr --:= r.::: _IenuSfIOm Stlata Protoconodont graphical evidence that eucondonts evolved r *:-se:ha. Wisconsin. from paracondonts by acquiring a crown f -: !::i tissue nor the (Bengtson 1983b). j E -' . =rrmal is perfectly An alternative hypothesis for the origin of pe,---: :-rorvs considerable conodonts involves phosphatic cones of the s:: : tunction of coni- ge )s Fomitchella, which has been found in -i s :.'.\':ukesha animal is Lower Cambrian strata in various parts of the i rc - ::.--:rt ed bl Smith et a/. world. Bengtson (1983b) illustrated specimens - -::: r :: rolume, where a and demonsuated that the cones are con- tr::: :: - 3,,ripanderodontid structed of lamellae that appear to have been :lE -:.:: added sequentially on the outer surface of the units. Thus, they compare with the mode of secretion of the euconodont crown, and it is L\ {\D E\ OLUTION OF conceivable that euconodonts may have evolved from Fomitchellq or a close relative and not from paraconodonts. Bengtson .---:. .olsisting of z (1983b, p. 12), however, preferred to consider H:: -- r-ln and a basalbody, the resemblances between Fomitchella and t:< .:: C: mbrran. Bengtson euconodonts as convergent or fortuitous, citing Tf,'l|-:.-: '--l3ttheir ancestry lay the lack of a known basal body and the ran- lu: :. ::t.rm which he sug- dom, finely granular apatite crystallities in the ncr:: :::-.iigh an intermediate e lamellae of the former as dissimilarities. In E , .. I 1-:). This sequence contrast, Dzik (1986) believed that it is plaus- r:, : : :radual retraction of ible to derive the regularly oriented crystallites r .: : |BJlro$Fl B :{Eotolqoa"led luopouoJ 0€ Lqcl ^ia!^eJ References 31 ir [Ch. 1 Proceedings pean ConodontSymposMm (ECOS IV), NottinS- f :--. l : G. 1986. Cono- Geological Society of America for 65 233 ham 1985,Abstracts, University of Southampton, rtu*- .-: \l H ,\itecki (eds): 1935, 436, alsoPan-American Geologist 1938 Conodont i ; r: :: L nl\ ersrty Prcss. Branson,E. B. and Mehl, M. G. (abstract). Society of Dzik, J. and Drygant,D. 1986.The apparatusof -: E. N. K., assemblages Geological =s - : : Clarkson, panderodontidconodonta. Lethaia 19 133-141. t ---. iiliniries of cono- Ameica Proceedingsfor 1937,210. --._ Clarkson,E. N. K., and Aldridge, Ellison, S. P., Jr. 1944. The compositionof cono- E|:: - --: ::. Carboniferous of Briggs,D. E. G., a\imal Lethaia 16 donts.Joumal of Paleontoloq! 18 133-140. ) t-9 291,. R. J. 1983. The conodont rri: -.---:-: 19 Ethington, R. L., Engel, K. M., and Elliott, K. L. g: , C . 3od Smith, M. P. I-14. D. and Monis, S. 1986. 1987.An abrupt changein conodontfaunas in the lE :-: .rn ol panderodon- Briggs, E. G. Conway -:: -- Burgess Lower Ordovicianof the Midcontinent Province. E-.:--- R {ldridge, R. L. Problematicafrom the Middle Cambrian -- -I In: R. J. Aldridge (ed.): Palaeobiologyof Cono- t l-'-. ::. : Faurth European Shale.In: A. Hoffman and M. H. Niteckr (eds): d.onts, Ellis Horwood, Chichester, Sussex, .'r,i I1 Nottingham Problematicfossil /l?,ra,Oxford Unive$ity Press. r- - ). LII_IZ1 . :: Sourhampton,2. Clark, D. L. 1981. Classification.In: R. A. Robi- L[ Fahlbusch,K. 1964. Die Stellungder Conodontida i --.:i ! 198-1. Ecological son (ed.): Trcatiseon InvertebratePaleontology, -tr':!.: im biologischenSystem. Palaeontographica, Abt. : .: .onodonts. sPeci4l Part W, Supplement2, Conodonta, Geological . - of Kansas A 123 137 201, pls. 16-22. El :. -: i - 1-+9. Society of America and University -:: Press,Lawrence, Kansas, W102-W103. Globensky,Y. 1970. The rrature of conodonts.,Le Dr: '-l :. \orb1. R. D., and Clark, D. L. 1983.Extinction of corrodonts.Iournal NaturulisteCanadien 97 213-228. lqs- l:. ::ihiecture and func- Gross, W. 1954. Zrr Conodonten-Frage.Sencken- Er:,-- :.- conodont of Paleontology 57 652-661,. -:t:ihacean fifty bergianalethaea 35 73-85, 5 pls. 1- . :-:,lse Palaeo- Clark, D. L. 1987.Conodonts: the final million led.): (ed.): Palaeobiology Gross,W. 1957.Uber die Basisder Conodonten. urri :. ,-1,r,1\ood.Chichester, years.In: R. J. Aldridge of Conodonts, Ellis Horwood, Chichester,Sussex, PaltiontoLogi.scheZeitschrift 31 7 8-91, pls. 7-9. - 1.65-1.74. Halstead, L. B. 1969. The paftern of vertebrate Arr:,:- i H T 1969.Acono- erolution, Oliver and Boyd, Edinburgh, 209 pp. : .-: -:. C.:rboniferousof the Conway Morris, S. 1976. A new Cambrian loph- pophorate of British Harley, J. 1861. On the Ludlow Bone-bedand F: : j 1:.,o-g_r12 400-405. ftom the Burgess Shale -: its Crustacean remains. Journal of I =. \lolnesian conodont Columbia. Palaeontology 19 1,99-222, Quarterly the Geological Society, London 17 542-552, r:rr--r- :-::s Bash Unpublished pls.30-34. fal- pt.62. art-. :: Ilhnois at Urbana- Conway Morris, S. 1980. Conodont function: Hass,W. H. 1941. Morphologyof conodonts../oul- trE::: laciesof the tooth model. Lethaia 13 707 7O8. nal of Paleontology15 71-81, pls.12-16. L- : ::e:tionable conodont Conway Morris, S. 1985. Conodontophorids or Hass,W. H. and Lindberg, M. L. 1946. Orientation r \:-:= O:dorician limestone, conodontophages?A review of the evidenceon of crystal units of cooodonts. of Paleon- le -:,. of Paleontology the 'Conodofltochordates'from the Bear Gulch "/orlnal " Limestone(Namurran) of Montana,U.S.A. In: J. tology 20 501,-504. Hinde, G. J' 1879. On conodontsftom the Chazy @: . \lcCracken, A. D., T. Dutro Jr andH. W.Pfefferkort (eds.).Palaeon- -- -' and Cincinnatigroup of the Cambro-Siludan,and Id,: G. A. 1979. The tology, Paleoecology, Paleogeography, Compte -.r:.::. from the Hamilton and Genesee-shaledivisions of q[: - .: O;do\ician conodoDt Rendu,Vol. 5, Neuviime Con|fts internationalde the Devonian in Canadaand the United States. Lc.i l: :i-151 stratigruphie et de g4ologie du Carbonifirc, Journal of the GeologicalSociety, Lon- -j :--: ar and GuadaluPian Washington and Champaign-Utbana, 1979, Quatterly 473-480, 2 don 35 357-369,pls. 15-17. ,trr- -: :::::::sraphy in Western Southern Illinois University Press, Hitchings,V. H. and Ramsay,A. T. S. 1978.Con- :',r --::: Stares. Journal of pls. - odont ass€mblages:a new functional model. 5- l-: l:.: Denham,R. L. 1944.Conodonts. Journal of Paleon- tology lE 216-218. Palaeogeography,Palaeoclimato logy, Palaeoecol- l:- -:---_:::e ot some Middle - 24l3'7-149. hf,:. .a: ::- aarl\ evolution of Domingo, X. 1983. Fosilesvivos en Lanzarote. ogy Cambio16 \o. 600,62 67. Huddle. J. W. 1934. Conodontsfrom the New Du Bois, E. P. 1943. Evidenceon the nature of Albany shale of Indiana. Bulletins of American 155-159, Paleontology2l no.'72,136pp., 12 pls. C r ,: :_,. I,reneed for a func- conodonts.Journal of Paleontologj l7 pl.25. Huddle. J. W. 1972. Historical introduction to the i.a; ll :l of Ordovi- problem of conodont taxonomy. Geologica et b, . :--:::..nal model for the Dzlk,J.I9'16. Remarks on the evolution PalaeonotologicaSBl 3-16, 1 pl. E- -,-i 16:E. cian conodonts.,4craPalaeontologica Polonica 2l -.r pls. Janvier, P. 1983. L"animal-conodonte' enfin h _:: ::r., nstory of the Con- 395-455, 41-44. conodonts. demasqud?La Recherche14 no. 145,832-833. r .-:,: 15 Dzik, J. 1986. Chordateaffinities of the -<-19. (eds.): P/ob- Elementarrangement in cono- ri L::r !1 G. 1933. Conodont ln: A. Hoffman and M. H. Nitecki Jeppsson,L. 1971. lematic fossil taxa, Oxtord University Prcss. dont apparatusesof Hindeodella type and in 4 101,-123. L= l:--: Dzik, J. and Drygant,D. 1985.The apparatusof similar forms..I-et aia Jeppsson,L. 1979- C !',=- \1 G. 1936. Geological panderodontidconodonts. ln: R. J. Aldridge, R. Conodont element function. Lethaia 12 153-171. r--,:: , :: ;onodonts (abstract). L. Austin and M. P. Smith (eds.)'.Fourth Euro- 'sldz 'zrr-0rl sld'9e6-626 pup ecrrerrjv Jo flarJos pcrSoloeg 'oJuopouoJ alotqaj sa4Jsllolslq)nJoN gj (8qotuoa\o4 Io lD tno[ Eu€Ipq tuorJ srel '7 ruarualddng 'A4 ltod 'f,SolotuoaPd :uI sluop F.I-;'inll lt) ol3ror5 ues -snlJluopouoJ uelJnlls pasnl bg6l V J'lJollod 1a^ul uo aslpatl:( pe) uostqou v u E{ | auozueru$rqzuerc ' LZ-1I 'sld '791-911 971 octtldot8 -ouo. Jo saurulJJeIecl8olooz 186I f ) 'relllhl Dl urf 086I H 'reqeru rep erSoloqdrouorT;''tr '89-gt 96I tadod ltnraas -oluoaDlod'ualuopouoC 'w.tlDtJut^ord 'tsn-8nv pun Sunzlesueuluesnz ueqcsruleqJtrlz 896I ottawy lo kaoo5 1rt.tt1o1oag -seurroJrluoporneN '.('pe) "I :uI rrr4l ' !tr'relSerz pu€' H'roured\' ['lqe^' H'reuzlerd puo salcololq uopouo) IleIC C |IrIISOTJIIIJ'UOqrSOdrUOJ 'sld /'16-I 'Smqsreled15 'ue{eqcsuessll[ 'ujsrlErcur,{ojdpu8'secEJolq'uollnlole luoPouoc b_11pue 1H C'sapoqU rap eruepu{V'sluauauD^no9 uaq)s-qlog uerchopro lseqreepue u€uqtuec t86I C I're[lrtr 'sldz - qJslssnY tap flaa$[S uaq)$Jnl1s sap aqJsq 's8-s1 llc I setes'uopuo-I ryE-'l srourll ureqtnos uafssol Dp anldDt8ouow 998l H C 'repued 'ttaDoSltttoA a|l'suo!DDsuDtf 1ott4doso1t44 q) 9uo uoiSutqsDlt'a14 '8rz-Erz 'V S n 'ursuocsr,/!\Jo uPunlIS Je,lr\o1eql uorJ 9 )rr.tdD.l8ttutsap puoltotl [a^lns elorq pe^.reserd,{lpuorldecxe V qs86t f '.VI-6, reded 'opouoJ lo Prlgopaj ^\eu -: npuav alautoJ 'J.lopuessenl)pue'5 g C's33ug'5 ('cln)lll J \ io| 'VtroJ 'rp,Qasav lua nJ uqlueq Jo lculsr(l , EJi.)apd '[8olo]uoapd 'puelq ereusellA'(uelct^opjo) uoqeruloC ulel LI L_9IL H rue r[ orlncl J I :uI -unotr{qurnql aqr uIoJJuo13ulql3 outpolag snuaB gzz arua',s'etotqperpoq-uosuElrnls v es86I I puE'C g (I'rlln{U A :qr uror; se8elquresse luopouoJaql Jo sralsnlJpasnl 6161 s c 'uEl^AoN 'Jropuessenl) O's33ug'C tr pue '99-I€ DdDd nlnutv GrlV J H d's€poqu 'sld IZ ' dd t0€ 'u8redru€qJ-eueqJn]e sroulll ItI lonads '8rnl\-89,^[ traoos Pueluol^I 'euolseut'I Jo frlsia^run 'slvTl Oltd pa4s4qndu2 stout111 lo In!8opaD ld :-€.iue) Jo {lrsJe^run pue eupluol^l ete4s (uetddrssrssrlq) ueuelsaqJ q.Fc rPeg eql tuojJ slelulue iulreeq-luopouoJ Jo 'eL6I 'M I IE foloeC 'otuopouoC uroJJsesnleredde luopouoJ 9161 6 1 '{q'to1 H 'ttoc5 puP rf c l[ uollentr 'Bg-59 rE+'o tu o a p 4 atu qaln iu I 'I9-6?'xessns'relseqclqC ' aJua .y ouol rF5 t{ :uI duouo poo^troH slllg 'sJuopouoC lo K8o1otqoao1ol -Dtuor plall pnuuo J|IZ 'ttalros loctSoloag pu€euols so:urela Jo se8elqtuesse :(pe) aBpupST f d :uI €uelpul ureqlrou -rorv sprurueBuueeq luopouoJ lsJlJeql rt Fy puE f H d 'sapoqt{ ruo.IJsrelsnlc luopouoc uBIrnlISJo uoneultu€xe -euI'I qclng reeg eq,L zL6I C i[ 'uollet{ . taqua' ='{uE){ aJuar^re'I'ssgrd -eu g J peorxe8 puE s u'lloJrN LOZI_ZOZI1 6961 186l uot lEs e.ueruv Jo &erJos '06 zI 'xessns'relseq.Iqc 'Poo^\roH -dag 'o3octt73 'uolwaiuo) 1ott3o1ouoa1o4 'tt quoN sSulpanotd elteTvowlelfue. i uDd 'tSolotuoaPd srllg' siuopouo) t o tSoplqoaDpd :fpe) e8prrply -uatuv aqJto qJlno rBag eql rt :r-ltt ejool,^trc d:uI 'I d :uI leturu€ luopouo. eql ul lueruale uo.rJeunej l16T D 3 'uollshl 'sld ott8o1oag D OOUTSOdJTluOUOXeI pUP eil eql Jo uoqcsnJ pue ruroJ /86I S U 'llotlN Z 06 I8 ISS DtSolouoaoPd n Fd ':96I Jt H d 'sepoqu 'LtI-EEI6 sJlstqdoag f,8opaC uDllo4snvJo 'euozolq pue sluopouoJ sll'sueredde (leFelz) Puo 'L ,G t.lrrJosloJll|dosolltld purnol erlelsnv uelsel 'ulsegSuluueJ slsuaJoquots oulpoltDzo 'zL6I L'e^oIqsBI l vws 'ntL-6zL Er-r.org sluopouoc eql iqt lo,t"tuoneq raddn eql ruorl (4561 'refetT ':!6I tuslu?qJeru SuueqleS-pooJ ril J H d'sepoqd ? Jloqcsrg) s(atq outpoltttzo pu€ 0t6I 'rel Ll [SoloruoaDpd ,Go1ot e se snt€lEddPluoPouoc eql I l 'tug4spurl 1-:.d. 196-999 97 -JnBlSsrllr: snlf,xsnqtou8f,1o4 selaeds luopouoc tr6l 'z0r-s8 trltls\_2 luopouot UPIU€A aqt Jo uoqrsodruocluetuelenFl l S86I S U 'lloclN ItI .;i6I [ntcog trBo tt J H C'sepoqd 'EIZ- L6I L s?ts[qdoa) Puo {8oPaD ndo1 lonadg Druauv Io loctSo1oag 'ELL_ILL :(pe) sepoqd :uI 8E uollousnvJo punoMg EIIeIsnVulelse^\ loozoapd luopouoC I H d 'stuopouoJ rql uo 'uglrspull r (!=1rUelal qlIAl IUOpOUOc 'urseg SutuueJ eq] Jo uEIuo^aCreddn aql ruo{ JosellrurHe €L6l hJ puB g 'dd 'uep lF\.r\ J 'pEorxeu IqeJ tr d, uosuejg snsuoclxa skpottJl Tvopouo. 961 E-_rl; S€'llotsuvsaPung eqt Jo uotllsodtuocluetueleqFl '286I S d'lloclN -retsurv 'rer^3slg 'sJuopouo) t96I I l'tuoqspul-I 'sLd9 oJlSolotuoaolod fun;pun4qy 'sito4sqv 'BZZ-LIZ,Z sctsf,tldoagpuo {3o1oag uollDusnv /9-l.C Z 21, untsodtutg tuop pulnof 'erLerlsnvuralsJ1\ 'ulseg Sulu la n!8opac uo,{.pleqo uejeJell seP uells) {o U/,{6r '9 S x pa) qnPluoqrs d H -uEC eql tuo{ qsIJposruoaeFd uEIuo,{eO leddn sneepun;uaddn:9-uetuopouoJ 3961 1'e8ue1 'Ed f,8o1uuoa1o7 Eo4 sjelsnp luopo uE ur sosnlendde luopouoJ ,16I S U 'llorlN Z '71OI-ZLOI 19 aql lrsl-:qr OgOt g f'DlsledeU '(74g1'uorlpe qrg 1/98r'uolrlpeqtt:6s8r .[o lDurnoI ;pow.qtoolirdns.luopouorolord aqestSolol puB ,{ruouoqdel'uoncn4suot :{lo^ urelsee J -iS4rtqJstnz 'uorllpapr€) 's1d26' dd 979 + t^x 1lt)oaaqlra^o ^{eN -uarue^\ols cruoJ€I eql uro:; sasnleredde(6561 Ji ur srelsnl. plog Io uoqnEustp aqi Io uo!|dlnsap tauq o q|Lt1 'uoqtqJollp Ftrll\ '816I v 's^otuQ{ sul,uat Jtu031o3u1u1D1UO) *l?d U$Obl lsaplo .le11n;lq),sranal snpopauootd, 1761 g'?wpue1 '9Z 'S'I-6ZI'xessns'ralseqJlqC'poo^uoH Id '862-18Z aqt Io [rots|| E\l PIrnllS tE8l I U 'uosrqJrnl\ e,iod DIJV sl€rn ure 'ZZ-I 'sld '88-I gg auag igo srll{'stuopouoJ Io t 7o1otqoao1o4 :(pe) aBpltply 'I eql buunP suorllpuoJcrueeco ,cdl1 wot! sldaptDtu -lD.ttu!Wpu, [7opag'tl$ta^tun o!o{X'aJuatJS l{ :uI uEunlrs '(I86I"qt seposlderoJ etuePr^e puoBnqul nE\ pelep)Z86I ll1ncog aqtlo stlouta7,t'1ueluopouoJ rep neq snolEruoueJo lo pue D -raddel) ' N A'^olqcnd -ureCuep reqn I 'ttue8o5 pue I X '.ra[ll^ -srp luoposoc lec€oloqll-J Z86I -I'uossddef 1161 '8ZZ olotqa'I sluetlt ,:-SSa III autzoSop1pa 'z I A\ -81,,4[ EI uoll.und 'uossddel BrtlnJ aqJ tr6l f 'appud 's€su€){'ecuolr\€'I'sseldsesue) Jo -ele luopouoJ eql Jo 9961 1 ^llsla^lun ,aaFar IEqJolsIl € :,(BololqoaBlBdluopouoJ I '{Jl aT [Ch. 1 References 33 .c-. l-:.. La\l rengs, Ktttaa, Priddle,J. 1974.The function of colodonts. Geolog- Rietschel,S. 1,9'73.Zv Deutung der Conodonten. ical Magalne lll 255-257 . Notur und Museum f03 409-418. -: :'l lnr-- - Uber den Fein- Puchkov,V. N., Klapper,G., and Mashkova,T. V. Rigby, J. K. Jr 1983. Conodonts and the early E:: l.- i.. of the Faculty of 1982 (dated1981). Natural assemblages of Pal, evolution .' of the vertebrates.Geobgical Society . ' . -- i:rr.agv and Mineral- matolepistrcm the Upper Devonianof the north- of Atkefica Abstlacts with Programs 15(6) ern Urals. Acta PalaeontologicaPolonica 26 671. 281-298,pl.25. Schmidt,H. 1934. Conodonten-Fundein urspriing- Ramovs,A. 1978. Mittelt assicheConodonten- lichen Zusammenhang. Paltiontologische clustersin Slowenien,NW Jugoslawien.Paldor- Zeitschrifi16 76-85, pI.6. tologische Zeitschrifr 52 129-137 . Schmidt,H. and Mriller, K. J. 1964. WeitereFunde Repetskr,J. E. 1980. Early Ordovician fused con- von Conodonten-Gruppenaus dem oberen Kar- odont clustersfrom the westernUnited States.In: bon des Sauerlandes.Paltiontologische Zeitschrift H. P. Schdnlaub(ed.): SecondEuropean Cono- 3E 105-135, dont Symposium (ECOS ), Guidebook, Scott,H. W. 1934.The zoologicalrelationships of Abstracts, Abhandlungen der Geologischen the conodonts. Journal of Paleontology 8 Bundesanstalt,35 207-209, 448 455,pls. 58-59. Rexroad,C. B. and Nicoll,R. S. 1964.A Silurian Scott, H. W. 1942. Conodontassemblages ftom the conodont with tetants? Journal of Paleontology Heath Formation, MoDtaDa.Journal of Paleon- 38 7',11-773. tology16 293-3O0,pls. 37-40. Rhodes,F. H. T. 1952. A classificationof Pennsyl- Scott, H. W. 1969. Discoveriesbearing on the vanian conodontassemblages. Jolrnal of Paleon- nature of the conodont animal. Micropaleontol- tology 26 886-901, pls. 126-129. ogy 15 420-126, 1 pl. Rhodes,F. H. T. 1954. The zoologicalaffinities of Scott,H. W. 1973.New Conodontochordatafrom the conodonts. Biological Reviews, Cambridge the Bear Gulch Limestone(Namurian, Montana). PhilosophicalSociety 29 419 452. MichiganState University, Paleontological Series I Rhodes,F. H.'L 1,962.Recognition, interpretation, 81-100,3 pls. and taxonomicposition of conodontassemblages. Smith,M. P., Briggs,D. E. c., and Aldridge,R. J. In: R. C. Moore (ed.): Treatiseon Invertebrate 1987.A conodontanimal from the lower Silurian Paleontolog),,Pat W, Miscellanea,Geological of Wisconsin,U.S.A., and the apparatusarchitec- Society of America and University of Kansas ture of panderodontidconodonts. In: R. J. Press,Lawrence, Kansas, W70-W83. Aldridge (ed.): Palaeobiologyof Conodo4b,Ellis Rhodes,F. H. T. and Austin, R. L. 1981.Natural Horwood, Chichester,Sussex, 91 104. assemblagesof elements:interpretation and tax- Sweet,W. C. 1985. Conodonts:those fascinating onomy. ln: R. A. Robison (ed,.): Treatiseon httle whatzits. Journal of Paleontology 59 InvertebrarcPaleontology, Part W, Supplement2, 485-494. Conodonta, Geological Society of America and Sweet,W. C. and Bergstrdm,S. M. 1969. The tir.ra i: :::3ia of Montana and University of Kansas Press,Lawrence, Kansas, genericconcept in conodonl taxonomy.Prcceed- : - _,rerrs. :grrj; --, University of w68 W78. ings of the North Ameican PaleontologicalCon- !F -'r--:-!:1. :03 pp., 21 pls. q Rhodes,F. H. T. andAustin, R. L. 1985.Conodont vention, Chicago, September 1969, Part C, :---=: : :i:er ofthe conodont assemblagesfrom the Carboniferousof Britain. 157-173. ::.-rr the Thumb Moun- L:=:: In: J. T. Dutro Jr and H. W. Pfefferkom(eds.): Sweet,W. C. and Bergstrdm,S. M. 1981. Bio- ,r :-: ::.:r. Ellesmere lsland, Paleontology, Paleoecology, Paleogeography, stratigraphy and evolution. In: R. A. Robison CompteRend.u, Vol.5, Neuviime Congris Inter- (ed.): Treatiseon InvertebratePaleonotology, Part 4rt --:r;;.i. Paper 79-14, nationalde slratigfaphie et tlegiologie du Carboni- W, Supplement2, Conodonta,Geological Society fere, Washingtonand Champaign Urbana,1979, of Ame ca and University of Kansas Press, Southern Illinois University Press, 287-300, Lawrence,Kansas, i..:;.lJ W92-W101. ,.i1.:", der Russisch- 2 pls. Szaniawski,H. 1982.Chaetognath grasping spines s, .--. Akademie der Rhodes,F. H. T. and Wingard, P. 1957 Chemical recognised -:::r among Cambrian protoconodonts. St i:::.:::s. I 91,7 pls. composition,microstructure and affinities of the Journal Paleontologl i.:--:: of 56 806-810. - "- H and Ziegler, W. Neurodontiformes.Journal of Paleontology 3l Szaniawski,H. 1983. Structure of protoconodont x-r{::- Z:s3mmensetzung und 448-45,1. elements.Fossils ond Sl/ata l5 21-2'1. Rieber,H. 1980.Ein Conodonten-clusteraus der Szaniawski,H. 1987. Preliminary structural com- 5- r- . ,:-:7. Grenzbitumenzone(Mittlere Tias) des Monte parisons of protoconodont, paraconodont and i{ l--:: Srlurianconodont clus- San Giorgio (Kt. Tessin/Schweiz).Annalen des euconodontelements. In: R. J. AId dge (ed.): DL -''::- of Paleontology 43 NaturhistorischesMuseums Wien 83 265 271, Palaebiology of Conodonts, Ellis Horwood, 2 pls. Chichester,Sussex, 35-47 ITEIOUOJO]OJd UEIJqU?J ) P-J6-ololsrq peqrr.ssP f6,I ) uosrBuog fq palues nld uro{ sluopouocne P lapou lerDeqlodf,q o! alq€u8rssEplu? (Z96I ll!1--' JnJl eql Jo seAn rFrsuoJ uaaq,{luoruuroc PUP SluopouocoloJd .\OLLJOOOUJNI I'Z '.{1a111peraprs F'uP sruopouoJ JoJ ,{Jl sgreu6otaeqc urepou ;o o fYl u-troqs el? sluauala { paurol ;o srred o,tr1 ul\-nIII aJE u^\ojc eql uI :':.1uaulela luoPouoJne :ql uo srncco Surleoc Eolouroq ;o suorlsaSSns IrqoJotord Jo f8o1olsrq tq1 idorso.rcruruo:pe1a res Eursn pele8rlse,rur IEOpOUOJne e,rqSUUd Er{Isslusurele luopouoc lxr? prnlsnJlsorcru eql JJVUJSflV r{s^\eruezs H luopouoceJsd ls AJ?ururleJd -ouoc urselc,{J t86I lI H'eue.I pue lL 'stdz'9€I-80I 'Zt-I std '90I-I 19 Sunlcstoluapog"ela.rz 9I uaLpunw nz ua{DqJsuasslA4Dp anaapDlv rlt saiuDsapuDl uaq)flssaq sap uaFunpuoqqv uaLlJslD[Dq X Dp alqcuaqs8urzrrs' ueluopo 'sInls sep ueluopouoJ ng6l H o 'resllF^l -uo3 .reqll 9881 A I 'uoqoll pue ){ 'tellz 'II std'Eq-I'ZI ry unamry puoabN Lq3l lBJlrolslq e :,{SololqoaBpd luopouoJ VE ^ro!^eJ i' [Ch. 1 :_ -: De\ooian to mid- h i. -I -\ldridge (ed.): f :. :,r;.. Ellis Horwood, r- --- ,^: kn': - -- \ 1886.Uber Con- gm.z-.:. : .iz. K. bayerischen E .-.:-::.,: :u Munchen 16 Preliminarystructural comparisons of protoconodont, paraconodont,and euconodontelements H. Szaniawski ABSTRACT grasping spines of recent chaetognaths (Szaniawski1980, 1982, 1983). The similarity The microstructureand composition of para- in shape between these two structures had conodont elementsand of the basal bodies of been noted earlier by Miiller and Andres primitive euconodont elements have been (1976) who consideredit to be a caseof con- investigatedusing scanningand transmission vergence,but the strong similarities in indi- electronmicroscopy. The closesimilarity to the vidual elementsand the almost identical con- histology of protoconodont elementssupports struction of apparatusesled me to interpret suggestionsof hornology.A thin, translucent, protoconodonts as close relatives of the coating occurs on the crown of primitive Chaetognatha.Thus, the relationshipbetween euconodontelements, and organic admixtures protoconodonts and euconodontsis of con- in the crown are illustrated for the first time. siderable biological importance and merits Two pairs of joined juvenile paraconodont detailed investigation. The imprints of com- elementsare shownto comparewith the teeth plete Carboniferous conodont animals dis- of modern chaetognaths,and a common ances- coveredrecently (Briggs et a\.1983, Aldridge try for conodonts and chaetognathsis con- etal. 1986) differ somewhat from recent sideredlikely. chaetognaths,but do no excludethe possibility of a common origin. Unfortunately, to date there is only one describedfossil specimen of a 2.1 INTRODUCTION probablechaetognath animal (Schram 1973), and its food-capturing apparatus is not pre- Protoconodonts and paraconodonts have served.However. some indication of the rela- commonly been consideredto be close rela- tionshipsbetween conodonts and chaetognaths tives of the true conodonts (Miiller 1959, may possibly be derived from comparative 1962) and assignableto the same phylum. A histological studies of protoconodont, para- hypothetical model of the evolution of conodont,and primitive euconodontelements. euconodontsfrom protoconodonts was pre- The preliminary results of my investigations sentedby Bengtson(1976), and I haverecently into this are presentedin this chapter. described histological comparisons between The material I have studied is from the Cambrian Drotoconodont elements and the Upper Cambrianof Poland,the Cambrianand :xnroJd Jo JadelJeuur Jo llmq erz,(aqt'(ggOt'gg6y) r1s,v'euezgJo alnl4sul aql uI pe^reseJd sl Pueleuj eqJ !+raltros $ uoll€uruI?l puE (916I) uoslEuag f,q peqrrcsep uaaq 'uo[€rrIroJep prole ol sueultads peqJle 8ul,ftp P Saurdsur seJnlcnJls s?q slueureleluopouocolord Jo aJnlJnJtserLL JoJ pESn urrq s"q poqtaur lurod tBf,lIJJJeql lapm a,reqI (C€ 3g sf,unrJnuJ,s 'sesEJauos ur 'pue 'AurqJlo roJ pasn ueaq e^"q Errru"I prce culu oTogpue epqdps runruoqC '0AISS) oslu sr saurds JNSIAIS.IS8O SNOIHIUJSSO Z'Z rp r€ql lreJ aql pepPe ,{docsorcrur uoJlJale SuruueJs pue '(WAJ) 51 qs,nztuezg)dlsnor,r ,{docsorcrru uoJlJale uorssrursueJl',{doJsoJcluJ @duroJ eql ol uotl 'uapa,{S ',&rs IpJqdo ,{q parynts ueeq e^"q sueulrJeds parBd mDrqJ 'Jru?8JodleJnd -Ja^run epsddn Jo faololuoeel?d Jo atnlqsul -aJd eql saprts euoloJJflIJ€Jtln pue 'seplls lmra 'seurdsSurdser8 eql ur sreqto pue 'de^uoN 'ueEJegJo,{lrsre^ euroloJcrrrJ'suorpes urql 'crls?ld u peppequra IPql ol l?lltuls ,fte^ -run ',(uroleuy Jo alnlqsul aql ur eperu aJe^{ sueurrJads Jo suorlJes peqrte pu€ peqsllod ld ;o arnpnrls aqa sJrpnlsaql Jo Jtuos u3{BllepunelJ^\ suorlEE 'sac€JJnsarnlJerJ peqcle pue qse4 Jo uollsul 'suaurrJeds -rlse^ureql Jo lsour ereq^\ '^\?sr?d\ uI seJue -rlex3 pe^Io^ur s?q uoq?Eqsa^ul aql '"Iuolsg qdomr aq ,{po uec -rJS Jo (ruap"Jv qsrlod eql lo fSolorqoe?Ied Jo coperlerl aql pu? 'uepe.{rs Jo JopeureJl r ral E Jo PelJnJlsuoc 'Ilaleurur€l UreA€ 'uql 'nuueo5Jelno - co xll aseqeql spr€,!\ol 'uarolc- IJ 'SuJ IJauor Jr 'auocFseq -.q 'la.(elratno - Io 're,{elolpplur - lur 'ra,(plrouur - II (g) lueruele rF wo4 pePP" ere^\ tuopouocnae^rlrLuud E pue (rg) tueuralaluopoumotord e Jo suoq.es pulpnr€uol Jueruoqss-I Z BI{ rE -(Iuepuocesueuo rc ratno eqt u€qt pal€ eq pue -ra11erucruzSro n aql qlla 'sauoz ra-ie1sqa'satllplsr{rr trsu f,ru?8JoJo pesod s n re,tel elpprul aql dho ,{1ernd'urql .{:e,r n'Z '8lC) s.re[21earql u,'i, t psp ruacnl ) -I\dz 'uape,rs ureqtnos ft -ds ouqnutns-17 tu9 lll fz z 'aes z'\Jl sluaruep luopouoJ go suos.rredurorlernlJn4s 9E [Ch.2 Sec.2.2] Descriptionsof elementstructur€s 37 The structureof paraconodontelements was documented by Miiller and Nogami (1971), but it is still possible to add some details. Etched longitudinal sectionsof specimensof Furnishina(Fig. 2.21 Pl.2.f , fig.5) showthat they are constructedof more than 30 quite long lamellae. The lamellae are thickest cen- trally and show the direction of most rapid growth, whereas towards the margins the lamellae are narrow and dense. Some of the initially-secreted larnellae are probably cir- cumferential.The basalcavity wasformed dur- ing ontogeny by the invagination of some lamellae;juvenile specimenshave no cavity at all (see also Szaniawski1971, p.404). Suc- ceedinglamellae grew mainly towardsthe base and are discontinuouswithin the cavity, but FlE. 2.2- Fw nishmasp., Upper Cambrian, Uddagerden, display growth both at the outside of the ele- southern Sweden,ZPAL C.1v135.1, x135. Note trans- lucent distal portion of cusp. ment and inside the cavity. The earliest of these lamellae are extensiveinside the cavity, three layers (Fig. 2.LA\, the outer of which is but later oneswere added mainly on the out- very thin, purely organic, and homogeneous. side. The rniddle layer is thick, laminated,and com- Transmission electron microscopy shows posedof organic matter and very fine apatite clearly that the paraconodont elements are crystallites.This layer can be divided into two composed predominantly of organic matter, zones, with the inner zone containing less with minor admixtures of phosphate. In organic matter and being lessregularly lamin- ultrathin sectionsof demineralizedspecimens ated than the outer one; the inner zone is also oI Fumishina the organicmatter is in very fine often secondarilymineralized. New lamellae concentrationsthat are denselyand apparently k; : :-:-:i\-' euconooont were added from the inner side and grew irregularlydistributed within eachof the lamel- a : - : :: :rr:lq. cr - crown, towardsthe base.The third, innermost,layer is lae (Pl. 2.1, figs 7, 8). However, in some sec- again thin, laminated,and mainly organic.It is tions the concentrationsare linear, suggesting constructedof a few very faint laminae which that the original structureof the organicmatter :E :r rsh Academy of Sci- can only be recognizedin very well presewed was fibrous. Sectionsof specimensthat have - r ::: :irost of the investi- specimens. not been dernineralizedshow that the phos- E-:r: - Some of the studies The structure of protoconodontelements is phate occurs as extremely fine, nearly e -:j: .-:e of Anatomy, Uni- very similar to that of Recent chaetognath equidirnensional crystallites (Pl. 2.1, fig. 9), c \: ^:r. and others in the graspingspines, except that the latter are of a apparentlyirregularly arrangedin the larnellae. aE-: : J,r of Uppsala Univer- purely organic,chitinous, composition. In addi- The similarity of the distribution of the crystal- tion to the comparisons documented pre- lites in the non-demineralizedsections to the viously(Szaniawski 1980, 1982, 1983), can be distribution of the organic concentrationsin addedthe fact that the inner layer of grasping the demineralized sections suggeststhat the N\S OF ELEMENT spines is also laminated (Bone etal. 1983, crystallites incorporate most of the organic fig.3G). I have independentlyobserved such matter in the element. Frr:: :r-rnodont elements has structuresin spinesof modern Sagittq,and. the Under the TEM it is possibleto differentiate :t 3-nglson (1976) and lamination is somewhatsimilar to that of the the lamellaewithin a singlespecimen, with the : : - ]-: ). They are built of inner laver of Drotoconodontelements. organic concentrationsbeing more dense in 000 otx I^lgJ '.uolres ssorcurqtElln 'pezrlerauruepueaq lou seq leql ueulceds E Jo lled Pseq eIIl Jo eerv 6n'IlLl'J'lvdz 'pun1o4ur.qirou '"1oqaroqcel,{oujpz 'euoz sltutollsld srysouSv'veuqueCreddn ' ds ,??4qurnl-6 BrJ '000zzx nar 'uoqcesssorc urqtprlF uI ueurlredspazlereultuep E Jodsnc eql Jo rerv €S I/AI J'MZ 'puelod ulequou '"1oq.loq 'aioz i*toitvd'srtsor3y'ueuqueJ reddn'DJs^\EIuezSDlolD Dul4slurnil-1 Bti! ""r^ouln7 '00t tx ,{AJ'uo$ces ssorculqiertln uI ualulcedspe4lereuurep e Jo ?d PsPq eql Jo eerv 8n'LlLI1 lVdZ ;pu?1o; rtutlollsld srysou8v'leuqueC reddn ' ds StJ ui"quou 'e1oq"ioqral^{our?Z 'euoz 'u?1s!urn!l-L '000 OIx wgs 'uoBcssIeulp$lbuol peqclepue peqsloo 31J e Jo ued IgglAtJ'Ivdz uepe,r\suraqlnos 'ueprgSeppn ',u?uqtuEcreddn',ds oul|nurnll-g ;u;;rola.lo peqcls pue 'obsx I,,tai itred 'c'q9 6g7x I,,{as 'ueulcedselatduioc'es uoqres leulpnllSuol li piqqfoi re/,trc fvdz 'uepe^\sureqlnos 'uePrgaeppn'ueuqrueJ leddn 'ds Dulllslut v-x-es 8tt '06x tlJgs luetue8uerft prnleuun uI pesnl stueuele elua^nl 'Br! o,y.J l 8/AI l lVaZ puslod urarDrou'eloqoroq cel^rouEz 'uelrqurEJ r.dd.r' ds Dulqstunl-r '0tIX WAS'tueuraguellPlErnleu ur pesnJslueruole eltue^nl Rrl o/'rJ 0I I/AI l 'MZ '.pu?lodurequou 'eloqeloq .el^{oujez 'ueuqureJ teddn'ds Dutqsluttlf-€ ;qri.t 8tg oqt"-y{gs o^r Jo orer,.'dlnec dFa € rl^tJ'tvdz'lrruepv quou'lueced ' ds o11t3o5-7 '09'x l^tas elelduocul auo lelno lvdz'4:lreDv qtrou lueceu ' ds I,r/l3l7s-I 3U 'qteetJo Jo ^rel^ 6I ZlLl J ^\or I Z erEld Jo llmq sr (aum PsBq :) fpoq IEseq aql Jo tred -BIereErcl f,1pnsnsr fpoq I"seq aql 'sluaurele aAIlIuIIId uI /rroqs eseqluI eJupuodur J?[ncIuEdJoere uzlc Ielno eql sluoPouo3na 13111 "lells suoqe8rlsa^ur,{ur'ra,ra.tog'luesald IEIJaI?u -rAopJOlse^\ol pu€ usrrqu€c (xo{ slueuele p f,ru€8ro Jo lunour? eql pue (896I lo nu luopouoJnaa^rl uud Jo sapoq Ieseqeql 'sluop -zlerd aas) setll"tsdIc a1r13dEeql ]o ezs eql ol -ouorne pu? sluopouosoloJd uae,nlaq drqs enp flur€ur Sureq o,n1 aql uee^laq secueJeJJlp -uou"ler fftuonnlo^e ue Jo ,Qpqrssodeq1 3ut eql qlra\ 'u^lon eql Jo esoql pue ,{poq pseq -1e8r1sa,ru1u1 u^\oJJeql Jo lsql se ul{roul a^\ q3noq11e eql Jo esllalrrEl eql uaa^{leq uop"nuBuoJ lue s" lou ll4s fl eJnpnrls s1r 'slueurele -pJoJuoc paauSoJeJ oqe ,(aql 'eJouueqlrnc luopouoceredpu? luopouocoloJdaloq,rn q1r,t " 'sref,el o^U otur fpoq PS?q eql elequaleJ snoSolouroqaq ol (886I'916I) uosr?ueg,{q -Jrp 01 uoseer ou,r\es [eql 'ecueH 'e?ueln"l peraprsuorsp^\ fpoq IEsEqeqJ '(1461 1ue?op eql Jo aJnla^rm fueqs e ruo]; pelFsel aJu? pu? JellntrN 't96I uroJlspurl) aull aluos -prossrp lueredd? eql l?ql punoJ pue'a? auIel JoJpezruSooal uaeq s€q slueurelaluopouocne snonu4uoJ eur?s aql Jo llmq eJE ,(poq pseq Jo dpoq I?seq eql Pu€ slueruele luoPouoJ aql eql Jo ,sraf,"l, qloq leql par?ls (IL6I) Iu?3oN -eJed uaa,t$aq .{1ue[urts leJnltnJls pue rellnl^l 'fpuenbasqns 'luEproJslP sI slai(€l 'A[Ip€eJ o.{$ eql Jo uorlsurlllel aql 1€r{l slseEEns(t 3q arorrre^losslp Pue '(ZZ'ErC) d elsrp cUeqd '1961 ssorg) uorlerlsnp sq pu? 'SuIlFI sll ueql -soqdaJorII pue luecnlsuEl aJ?(euoz sluttotlsld Paleulurel {laurJ pu€ f,lJ?lnEal arou aq ol euos snlsou8v et4!a^oq") 31?JlsJe8uno,{ tIIoJJ suaru eql patap$uoJ eH 'SUIIIIJ auoJ pue -neds auros sdsnf,aql 'ra^eaoH 'prceclJolqc I?seq " Jo euoc Pseq e 'sre,{el el"J€des o^\l Jo papruls -oJp(q Suorlsul aalossrytou op pue JnoloJ uI -uoc ale pelsErlsa^ur eq 13ql stueurele luop IJ€lq f,llpnsneJe uorEeJ crtpg eql Jo uerJqutuJ -ouorne urloFrrrBl pue luroJluoc uel^olpn'I Jo raddn eqf uro4 sualrlrredsluopouossl€d '(S '?tl serpoq lBS"q eql '(rs6I) ssolc 01 Sulproccv 't'Z '14) ernlcnrls 'prJs srJolqJorpfq uI e^losslp lou oP pue ua\orq ,{Euodse ,r\oqssuoJloas Jo serB;rnspeqJla pu? {JEp ro {3elq,(11sou are uoISeJ 3llleg aql peqsrJod'suorlsrlJru8eur q?rq tV seteds rel rrro{ sueurrJedsJo sapoq l?s?q eql (g'I sSrJ -Ieuelralu reFAaJ'urql dq peler€desuego are 'Z Z'ld) flr^?J l?s€q u,t\o sll qlr^l pedeqs qt^\or8 luauele Jo polred ral"I aql uI peleras 'ld) leuunJ f,luouruoc sI pu€ urt\olJ eql ol e^rl e" erlJelaqJ '(t'7rl't'Z sreqloueql auos lz'z'Jrs z qJl quetrrep luopouoi! ;o srrmgedruoe IBJnpn$S 8€ [ch.2 Sec.2.2l Descriptionsof elementstructures 39 L :-: : :Lrmmonly funnel- :; :..:l cavity (Pl.2.2, rr :, -:: ,.t specimensliom r-: -ri:l\ black or dark fr!:: : I rr drochloric acid. :-:.. :j- r.the basalbodies fc:-:: .:: ::miform eucono- !r -: -,.:!ilgated are con- =--.:: .'rers. a basalcone E rr:rlrdered the basal 3s-: - r :rd iinely laminated r-- -rr,rn t'Grncc 145? Ir --: ::::r:r3lion of the two !L : .-: --ntl]. Miiller and .if.: :.-:: :r.rh layers' of the ll1: : ::: :ame contlnuous lB: --:: ::: apparentdiscord- [ : : :-: aunafure of the =: -:: nO reaSOnto dif- air : :, rnto two layers. i :::..gnized a concord- -. ::r- lamellaeof the D:\:-l;::- : :1. Crown, with the -: :: :.' -r being mainly due l:llitcc lcac Pietz- nf nroqnic l--... = . n\ investigations fi : : -:.r:rodo[tsthe outel r:r:. = ::::l cone)is built of .i: :.r\ of teeth, iE\1 i,160 -::: -: - - l\ i 10 Two 1-:: l: a i\ E 1. Two - I - - l'. :- 3. Polished (-. :=: - : . SE\l x500 : :-: Part of a -- = - _:- : .r:lhern Poland, rr- _ - :- .. .eclron. TEM - -, I --:srec borehoie, _ rrE: - ::.::: arosssectlon! :: - _-- = ..lhernPoland, Er -. _ .: rlrrarhin cross 00I zx I lgs EereeruEs Jo dn-asolc'c9 qLix WgS 'u^lorc pue,{poq lesEqueealeq tceluoc8uB\oqs peiP'qS 09x f^lgs '!uoqces r'ds leurpntr8uolpeqcte pue peqsrlod'eS t 0S/ I J -MZ 'eluolsA]o ropetuerJ snpofplo)-7-Ei Brt '000zIX Ngs 'euoc rHuotsg I?seqJo eceJrnsralno Peqclapue peqsllod l 6t/AI J -IVdz Jo copr?uerJ' ds gtPoltp.toJ-n Bl& rq€ '000IIX wgs 'euoclPseq oql Jo oceJrnsrelno o/IX 'lcelul ,(lluerpdd€auoc eqt lnq tpe^losspSuIIrJIssEq puE u,{orr luopouorneeql Sul.{\oqsuoqces nAS lEs€cl leruotsA anbrlqopaq.ta pus peqsrtod'e€ I O9/AI J 'IVdZ Jo .operrrarL re}.i-ol'ds snpollptoJ-q-s€ 3!g '009 gx wgs 'euo. pseq eql Jo IIs^\ eqt 8ullroqs'uorlJesleuPnlr3uol peqrle pue peqsrtodeJo uPd z ogl|lJ lvaz 'eruolsg Jo.opeurerJ le^ol ' ds sngopouozoz-z 8rt oolx ngs 'ueurrrdsrues Jo eJ'le'qI 0'I. wgs peqJle ,{ltq6lls,{poqlEseq eqt pue pe^lossrp(latelduoc tsou{e u^ror. luopouorneeq13ur^\oqs 'uorlres Ipupnlr8uol peqclepuE peqsrtod'eI Eogl1llJ -lvdz 'eruotsgJo.opeluorJ ra,nol'ds snluopouocoz qeI 8IJ z z eleld 'JeAa,roH qd€rSoJcruiuoJpele uolssrulsup$ Jo pesodrrroJare Aeql 1?q1,{oqs sllnser waJ e uo'ds snpol(pto3;o uaurrcadspezqerautuep aql pootsJepunlou Uns sl urErro rreql 1nq ? rxo+ Jell?ru rlueEro palerlsnqr pue'u,{rorc '(1461) lueSop pue re1n11,(q pale$snlll erall luopouoJna Jql ul splJe ourue alqeqord pu? (9S8I) repue; dq pelou fleurSr.ro aran parlluapr (996l) 't, t" reuzterd plr? cuolqr sallpreqdsqcng'(7' 1's?r1' g' 7 ld) e"ulru?laql -orpf,qur luauala u€ Jo uopezrl"reururape1a1d uae^uaq JncJo sernlJnJtstntlnseqds 'snpol{p -ruoJur',t\ols relp ,lsoq8 luopouot, peAasqo -lo3 snua8eql Jo [llelcedse 'suaturcedsetuos " (7961) uolspurl ploa'(nn6l uoslqg) rellzru yo ?u11pyauor eql ur 'dleuoqlppy (q1 '8ty crue8ro auros uruluoJ sluetuele luopouocna 't Z 'ld) ur8uo d:epuores Jo setl1els,fir Jo u^rorJ eql l?ql ua{oul uaaq Suol seq tI elrledere8rc1 qonur uaqo aJEeJerll SuIIIIJ auoc 'pezrleqdsoqd,{pepuocas aql uI 'suorl€rluecuoccrue8to eql pepunoJJns ueeq fpeeyc s?q lr seseo,{ueurur pue 'ul8uo f,1qeqo:dpue aur; ,{.ra,rosp ara,t selqptsfrc aq ,{reur:d ;o sr ra,{e1eqt Jr peqsrTqetsaueaq -eqdsoqdaq1 teql aterrpul eleqdsoqdpa,rlossrp lou spqlI ae11eure1crleqdsoqd eql uqlt,t\ rxo4 aqr [q rJelsaceds aq; '(Z'?tl't'2 14) sra,{e1 -regerucrueS-ro Surdpapun aq1 ;o pue re,{e1stql eql urqll^\ flrep8errt palnqr-4srp ,{lqrssod yo uoqearaserdselqeue 'JeAa/i\oq '8ur,{rp lurod 'suoqe4uaJuocesuep 'eurJ ,ira,r ;o ruro; eql lB.rlrJC lB ur perrp ]r sesdeloc I lnq 'lue^ ur sr,(poq leseq eql Jo JeIErurruPEJo aqf 'ngJ, -los eql ur 1de1sr 11se 3uo1se 'uaurJads aql qlh\ pa^resqo suerurcedspazrlpJaulurep uI JOuOrlEZrlsJJulurep rJus 'u,lAorJJql Jo edeqs '(e ?lr eqt surcler1I prcecr-rolqoo-rpfq qllt\ lueurl?aJl 'Z Z'ld) pe^ressrd fporuuoc aJou qcnu InJeEJ ol luplsrseJ sr pu? 'aJ€Jrns qloous sr1r'(4961) ssorg f,q pa,rrasqose 'pue'spIJ" ol € spq 'f,poq l€seqaql Jo areJJnseql ol f,lps"q lu"lsrsaraJour qJnu sl auof,leseq :q1 ur8r:o spuelxara,{e1 slql'(Z 8lJ't'Z Id) ra,{BItuetnl f,repuocas;o ,{pueprlesr qclq.{ Jo tred 'lualuoc -su?rl urql e ,(q peleoc sP/t\ur\oJJ eJllua Jlaql 1u.taururraq8rq qcnur ? qlh\ ae11eure1reln8ar lsql pepe^er ,{lpnsn slueurala luopouocne sselpue relJrqr qJnu ]o pelrnlsuor sr 8ut115 alqrurgd eleldruoc Jo Surqcle 1(oIS ps?q eqJ 'eleqdsoqdJo slunoue roulru qlun 'euoJJql ur uEqlSulllrJ IESEq 'prreleru crueSro1o ,(1a3n1pasodruoc are ael eql ur eArsuelxeaJorx sr qJrq^\ 'uoD?zll€Jeulru -latrr€leseql '(Z'?tt 'Z'Z '14) ece;rnseuoc eql elrr,{d ,{repuores ur?luoJ uelJo osl? suaur o1 1e1e-redpeu8q" e€lleureleur; r{raa.'re1n8er '(I '14:1 '3r7 -rcads,{ueru Jo sepoq ps?q eqJ .{lsnoaue-rod Jo 1mq sr auo3ps€q eqt 3g 'g'7 -rrJeluof,peleJJes dlqeqord era^\ snql pu€ 'Z'7,'ld,:gI'Z 3rg) .ra,(e1aleredas e se pal?arl '(Z'zrJ'EZ '14) uorqselrelurs ur pa8uerre eq o1 (3u911 euoo =) f,lluanbesqnspeler " 'aeunuel reln8a.r aql s? pHel€ur aruBs eql -Jesesoql ruo4 ,{puarcrgnsreJJIp teql a€leluel [z z 'res z qJ) slueuala luopouot ;o suos.rredurorIsJnlJmtS lch.2 Sec.2.2l Descriptionsof elementstructures 47 | .:.. --: regular laminae, k :-.-.,n (PL 2.3, tjg.2), rr::: ', : ci -:::rplete primitive [. -' -rll\ revealed that Es :::::3d b\ a thin trans- =_- L i This layer extends E : ::3 basal body, has a : :i:istant to careful rr;--::-: J acid. It retains the r- -r-.: :emineralization of e . ::s kept in the sol- :::;d in air. Critical- E-s ::.tles preservationof Rr.E:a: irns organrcmatter r':::: -:mellae. It has not f -:< -=..er is of primary r Jl:si :: has clearly been lr--: t L. .: ihat the crown of G :;l:l:!ln SOme OrganiC ]r - .:: Lindstrom (196a) i :'1r::- after slow, incom- 1r :i .: element in hydro- E :- j- (1968) identified rr:. : :he euconodont EJ 1;;=1; matter from a s ::{ordtlodus sp.on a u : -- ..graph. However, '- :,: :-::: rnd etched - _-:: rnu r.i- r.rd! slightly {.i t ,!f : :.r r:.: 3nd etched ftre: :-: ::.hed oblique L-.:_1-::_ r rntaca.sEM I- &- : -_.. .:r:-Iaceof basal a i: ::::--: roneitudnal EE --r- -ri close-up of ------ '(ae!oc pseq) ,{poq pseq eqr Jo eerp Dqrou" loj ,eerea.ues eqr uou'rrJruaeu ,uorlres 66otl* Jo r,nd Jo req8lq 'qZ 008 €x WgJ ssor. urqle4ln ur ueurneds pezrlereuuDp yo auqi4 aubc jo .IVaZ Jo u^rorJeql Jo ?ulleoJJslno eql ol puodseuoJ 'u,{toJ3 feru s1uautelaluopouoc?Jed pue luopouoc eql Jo eEIIeur€Taql q1I^\ ,$rnurluoJ ou e^eq -oloJd Jo Surleoc crueEro relno eql (e) auoc les?q eql Jo ee eur?l palatJes-,{lpuur .sluauela luopouoJolold aqt Jo euos ls€el tE l€qt lseSSnsgrq^\ 'suorte ur ueql sepoq I€s€q luopouocna pup sluaur -rederddur ur elqrsrl lou sT(1961) rcfiar7 pue -ele luopouoJeJedur pa8uelr? flrepSer arour Ilrortspur.Ipue (116I) rureEo6lpue regn41f,q al? setqpls&r aql leql lseEEnssqd€JSoJrrur pegodar ,{poqIes?q eqt Jo esoql q}l^{ a"Uorrr?I .selrlplsdlJ .suaurrads uo4Jele uolssrusu?Jl pel?8uole u,rolJ Jo JJUBpJoJUoJeq; raqlo .eel .{pq8qsro puorsuaurpmba eurJdle^ Jo urroJ uI pauJrJuoJeq ot 1ef,szq uoqe,r:asqo srqJ aql ur slueurale11p ur srnJro aleqdsoqd (p) 1aure1cqzqedsoqd s1r qlt. reBur;re1ur,{lq€qotd 'uo4e,r-rsse.rd;o peJJeuB eq f?uJ ,(aq1areq,t (S'?rJ 'Z'2.14) uno:c aq1 olur stql lnq 'paEuerre,{peln8er erou eq o1s-reedde ecuslsrpauos JoJ enu[uoJ fpoq pseq eq1;o lr seryoqlpspq luopouocna pue sluopouocered aeleruel crueSloeqt lpql sa{oqssnpol(pto7lo uI 'suorlsllueJuoc aur.; dra,r Jo ulJoJ eql ur suorlJasurql paqJlepue peqsqodaq1 yo oug sr sluaurela3ql IF ur Jalleu JrueEloeqJ (r) 'uotl?rlruur 'seprsJalno pue dlr,rzceql qloq uro{ papp" f:epuocas u€ql ,{lal[ erou sr sql lsql " eJea aplleur€l eql sluopouocet?d ur laprs slset8nssluqg eql Jo uorsuaurprxJoJrun {lJpeu /bI^eJ er{l uorJ e€llarrl?l;o uoqlppe q8norql eql 'u^{oJc luopouo3 aql Jo l?ueteru JruEBJo parrnJco slgl sepoq lBsEq luopouoJn6 pu? {rzur:d sluasarderlto,r\lau aqt leql pel"Jls sluopocolord q sos"q ryeql sprz,nol qlror8 -uouep fllEro^rnbeun aq louuec ll q8noqllv ,\lor.lspue peleuruel ar? srueuela eql tlv (q) 'azs elqeIJEApuE edBqsJ?ln8aJJr Jo els lau 'uorlrsoduloc cru€EJoJo slueurale 3ql uI saloH'(Z't sBtl'V'Z14) slrrqg;o palcmrs .surJoJ ruo:; ur8rro ue EullsetSns pecuelpe ur -uoJ {Jo^uau l?uorsueurp-$Jql 'leln8eur u? crleqdsoqd,{preur o1 suro; e,r4rurrrdur Jrup8to sl eJeqlul{orc eql Jo Surleooqrql aqt Jepunlpql ,{pteur ruo4 seAlole sluaulale luopouocna pelEe^aruoqezrFreururap'suoqe8rlsa^ur,{ru Jo ,(poqlesEq eql Jo uorlrsodruocleJrrxoqc eqJ ut pe:edardsluaruela luopouoone eArlnurrd eql 'seJntxrwp€ Jllpqdsoqd qlr^\ Jell?ru Jru?3Jo u1 {docso:cnu uo-r1ce1aSuruuecs qll^\ paulu Jo pesoduor alE sad,(llueurela ealql [v (p) -exa uaaq flsnorr'ardlou seq pu€ 'u.r\ou{ Ile,tr lou llDssr slueuelaluopouo:;o uorllsoduro: pu? arnpnrls eqt adocs :s,u.olloJs? pezrJsrrrurnseq ,(?ul pu? letlrueqc 'ra,ra,ro11 -orcnu lerrldo aql qlr.ti parpnls eq r(eru qcrq,r 'sluopouoJnea,r4rurud Jo esEJaql ur pe{Jerx 'eJnpruls repllac purEuo eql uela upt flrelncrged eJ? senup[u]rs eql .queurale IeoAeJ uoqezrlureuurep Surrnp sluaurele luopouoc luopouorne Jo ,(poqps?q aql ol Jslru{s ,{leso[3 Jelleur crueEro eq1 8u1xr.yteql e^alleq eJe slueurale luopouoJejpd pu? luopouot Jo 1o (fSOl'SSO1) aeg-u€^-snerqeCpue snerqpd -olordleqt (9461uosfueg'1461 rureEogpue 'xrrleur crueEro ,{rerulrd eqt rrJo+ el"rluaJeJ ralln4 't96I ruorlspuq) suonsaSSnsrerllpa ot [.re,r aq ppo,t|. pu? eql urrJuoc suone8rlselur prrEololsq dW -Jlp rlmrJJrp aelleulBl cqeqdsoqd eqt epe^ur [Erx I€ueleur snourur SNOISO'ICNOJ CNY NOISSNCSIC g'Z -n1rq,{repuocesleqt peJaprsuoJstoqtnp eseql lt'z'ras 7',tlJl sluaruale luopouot Jo suoslrBduroJ lBJrqJulS zv .) Ich.2 sa^ 1l Discussion and conclusions i-$D CO\CLUSIONS lesiaions confirm the , ll-;s:rom 1964, Miiller ie:.*---.cr 1976) that proto- ll|;-'-acont elements are c |;:-- \1d) of euconodont ir::::s are particularly r cr r---.firive euconodonts, rie: iollows: -r Ec: :..-s are composed of I :r:sphatic admixtures. E--':: Ji the basal body of L< 3',.-ir's5 from mainly k:r :-. nainly phosphatic s$3::ng an origin from :cfi=alsrtron. G :.:-- :aminated and show i :-=s- In protocodonts E?- -1.'dies this occurred f ir=:-ae from the cavity tc-< :-:e lamellae were r =; -:r and outer sides. E:: :: all the elements is r-T :: concentrations. In d,ll-_!:\ront basal bodies it arranged, but this 'pr=-:r-5---:\ ation. c: -: :ll elements in the lc:::ensional or slightly ti -:-.rsmission electron I =i: -5e crystallites are : paraconodont ele- tE: r.rr3l bodies than in -F: GtrJ E=:-: !-oaring of proto- U'::!:!-\nt elements may r-:irs of the crown of -: '009 IX I IAS '!uaolc aql ur lpualpu-Icru€8ro pue Suqeocrelno eql 3ul{\oqs'pZ ur raddn ,{clpolErrpur eerE ]o dn-asolr'qZ OIZX ^ror& I aS '(arorle re^\ol) pe^oll]ej ueeq seq u^\o.roeql ereq^i alqrsr^oum leseqeqt pue BuqEoclatno aql Jo sluerxSerJqlrlrr u,\\o.n pezrJe.reuruap aql Jo slueuulerBura\oqs 'poqlaui lurod JearlrjJeql {q parlp ueulceds {ez paqcteJo rr€d lEseq I89/AIJ'IVdz 'eruotsg }o roppuarJ ra,{ol 'ds snuopouoroz-q-Ez ?rt lsenlxrrDpe OoeZX WgS 'u^rol] eqt ur lplla]etu crue8ro8ur^roqs dn-esolc 'p ! 000 IX WaS cruE8roqll^{ u,r\orceql puEeuoJ lesEq or{l Burlroqs'el ur ,\\ollp re,{roleql ,{q palouepsors aql Jo dn-esolr reuoo 'aI I gS eql SurtlrJ8ur^1oqs 'e m morE reddn eql ,(q petouepreIP eqt dn-esolr Ojt Zx lpe^losslplEspq Jo (poqtaurI Jo '!qI O€ZXI JgS etpqdsoqdeqt Jolsour turod lEcrrureqr [q paup r rpdlEseq eqt Jo uoq]es ssorrpeqcl€ pue peqsllod 'PI 'IVdZ ' eruotsa .opErrratI ds sryuopouorog-p-el RtJ I 09/i\I l Jo '"N\o!' l z eteld -uof, ur seJueJeJFp erxos er? eJaql qdnoqllv -oone pue sluopouocolord ueernlaq aEe1q1 '(q'eggOt uoslSuag osp ees) sqteuSoleeqc d:euorlnlola pesodord eql 'Jeq1e3o1ue)pl urePourol Satlllellurlsl?lJ^es /r\oqs(9861 'P D '(g'7 3rg) nrorc e8puply'9961 1z1a s88rrg) ,{l1uarer peqrrcsep pqrur Sururoreqeeleruel r{1ree;o s1-redplsrp (uoq"ruroJsuEJl slPlxrue luoPouoc snoJeJruoqJeJ aql (aJ eql qth{ r€lllurs e luasardar 'erT€rxruv Euarxala luopouocerederuos 1o sdrl pezpeqd eql Josraqureru Jaqlo 1e ruo4 lJurlsrp fre,r pue -soqd,{13uo:1s aqt t€ql elqrssodsr 11'fua8o1uo perue^pefpreuognlo,re are s4no-r8qlog (p) ?uunp plreleru ul\orc olut ur-rolsue-r1fpred o1 'eleJeuedel srceddeollapouoqdlg ;o lueruelee4 e ;o e1e1d o1 ,!111qeelq€reprsuor e ,{eldsrpseurds qteu Ieseq aqt t€qt pa^resqo(746y) eusraog -8o1oeqcpue stueuele tuopouoJ rllog (J) '(136y serpuy) '(9961 1ue3tuq pue ul\orJ slr uuql ,{ueSoluour JerlJ"eSurdole,rep {tz(I 'S86I 'pp eflp1tp1y'186I r{s^\?rupzs snpol{pto3 yo f,poq 1eseq aqt 'aldtu€xa pue qsladag '€16I leqrslerg 3'a) sroqlne JoJ'qlk\'uJe €d etx€saql s^roqssluopouocne ,{ueru fq patou ueeq seq sqleuSotaeqc;o a,rrlrruud ;o lueurdole,rep crleue8oluo eqa snle.reddeSurdserS eql pue sasnlereddeluopo 'sluopouocered Jo sluopouoJolord ur enssq -uoc eurosuaa^$aq ,{3o1eue puorlcun; y (q) Surpuodsauocou seq pup ,{poq pseq aql ueql ',s,{e,np:e,res ur sluopouoJro; en8opue uoqnlole ur relzl pedole,rapu,no-rc luopouoJne cr8olocad1a111 e sr l€qt runlf,r{dE elnlusuoJ ' , eqa a8eeuq ,{reuoqnlor'a auo o1 8uo1aq sqleuSoleeqcteql pelou (699'd'1461) laa,r\g pue snoSoloruoqe.re sluetuela luopouoJne pue pu€ uoppas',{lrur;p qleuSoleeqJ-tuopouoo 'luopouoceJ€d'tuopouorolordteqt srseqlod,(q 1osuorlseSSns peqsrJqnd ,{u? aretr ereql eJoJog (9461) s.uosfueg oddns serpnls ,(141 'uoqnqrrtsrp qldep-retea pue pqoy8 epr,n 'PAIJIIP[J ,{oqs q}oq sqteuSolo?rlJpue sluopouo3 (e) . aq o1 1e( sEq srrll Jo elnleu eql lnq 'euoz reuur pelpuruel f,gelnEar pue ,{1eu1; e:oru :suorl€Arasqoraqlo auos qlrnAlualsrsuoc osp e sr eJeql sueurrJadseruos ;o 3u119 auoc eql uI $ qcq/r\ 'srseqtod,{qsql slroddnsreldeqc srql 'sarpoq leseqluopouoJna ur.ro sluopouore.red ur palueuJnJoPeJuepr^e leJnlJnJls aql sluoPo uI u.rou)l lou sr leql re,{p[ rauu pel€lueJeJJrp -uoc ;o sdrqsuoqelaraq1 SupreEar seseqlod,{q - e,r e eaeq sluopouocolord leql sr sedfl Jeqlo fueur eql u?ql uoDuel]? eJour luauale eql uae/$leq aJueraJJlp urcul eqJ ou pelrerl€ etuq leql te tnq '(€L6I) Ieqrslerg '9 '?rJ'Z'Z'ld uo pet€rtsnlp uouete [q pesodord 1s]rJ sE,!\ .{1rur;p qleuEoleeqr -derd aql ,{q perseSSnssr slql ',{poq leseq eql y sqleuSoleeqcpuP sluopouoc uee,rleq f1r JO a?ururelaql ql!/r\ snonulluol a:a,r sSurluoc -ur;;e ue f ldurr (tg At' Zget'086 I Dls,t\eruezs) eqt leql pu" 'EuD€oc clue8ro ue qcns passessod sqleuSoleeqc pue sluopouoJolord uea,rgeq u^\oJc luopouocne erll Jo ?ulluel qcee l"ql drqsuorlelare Jo ecueprle eql pu€ sluopou alqrssod sr lI sluauale luopouorne e,lrlruud [€ z 'cas z qJl $ueuala luopouor ;o zuoq.redruoc1urn1rn4s nn lch.2 (cn I ?l Discussionand conclusions 45 cr-r:Er;e of a relationship frl --i 3nd chaetognaths 1::: -!!-:timply an affin- bc- .:; chaetognaths.A proposed E "=: ,lrst by trf :: --::i me attracted no rr --: ::larionships of con- r- =r ::lce documentedin rE ----r :\pothesis, which is I :.-r-' !i:hel observations: - t: : :-:rrgnaths bOth ShOW r::::-:epth distribution. r,. : ---:.hed suggestionsof m= . Seddon and -:rin 19 - --:=: ihat chaetognaths ri:: --::=:is a likely ecologic hr: _ _.:reral ways'. [i - :-.. ]it\\-een some con- r,: :- i :iasplng apparatus Es --i:: noted by many s-- 1.--i- Repetski and -{-:-:.: errz1. 1985, Dzik L E: : and chaetog- r : :::.:derable--:i3nts ability to r: :. - ::ronarily advanced r :::er membersof the -- lri:, -: aLrnodont animals B:.::. :: ;j. 1983, Aldridge rE-r i llanties to modern r-: 3ergtson 1983a,b). c ;:::= :Lft-erencesin COn- L r' _. .ai erchedcross E:. . :: !E\l Y130. lb, Er : _: >:\l .l i00 lc, EE :. r ::-, n:rn organlc dr-.l- ::\! ,: lll0 I : _... :=:: of erched rA: - =- ir:h irasments l]l :- :::: . 1rB). sEM !E:.: - :a: ;a materlal .NI_I fueru peplrord 'uurlpl 'seJueIJS;o ,(ruapecy 9I Dloqta.I leuru€ luopouoc eql €86I t l{ ueruolsA'f,8o1oag 1o elnlllsul 'ErlIA a^lIA J(I 'e8puplvpue') N g'uos{rel3'9 g g's83trg '686 626 SINIWSCqfl'IATON){JV gg utop8uty pa71u2aql lo uottotcossylottSolorg auuon aqt{o louno[ sqlBuSoleeqcJo seutds8ur -dspf pup qteel eql Jo uo4lsodtuot pue ernpn4s eqJ €86I V 'proJslndpu€'d )'ue,{U'O'euog 'sluopouoJpuu sqteuSotaeqr 'Lg-6e asplaI 6n uaSuqapapawaqxs8opag uBrjquec uoe,rueq dels f-reuor1nlo,raa1e 't€6I '(tqen pue uosuerg) oloqol onapouolldtg ue SurluesaJdalsleurrue selsllueP pup 696I 'sseH uadooc o\Tapouorldtg -rpeurelur Jo Jo arnlJnrlsleseq aqr uO ft6l J ) 'elusJsog pe8udr?qd aql uaeq e^eq plnoc slue(Ilale '6I-S 9I UD4Spuo sl?ssoJsluopo luopouocPEd lstll pelse?8ns (q9361) uosl rq€86I I _g gt KSopnutw puo -uoJ eqt Jo .{.ro1srqdlrea aq1'q€86I S 'uos18ueg -8ueg (7961 nls^reluezs uoslSuegees) .8t ototx '8E gl ofptqa1 snlErEdde 7 'aJualcs to luopouoc spurrus ,{JeAJo 'eru"s aql [{lael aql EqopouoJ r3p nEqsrec aql roJ e€861 'S 'rElrurrs Jo lepour lEuorlrunJV 'uosl8ua8 pue SutdsurB 6o_1 pue f X 'rellg6 '902-98r ol slueuraleluopouoJeEd 'saurds G {4tpsqaz aq)stSoloj 6 OtoqtaT uoncunJ pue elnpruls luopouoc eql ol puodseuoJ sluatI]eleluopouoJolold r tunuquet uereqo Jo uonnlo^a dll€e eql puE 'sluopouocueIJqu€C ]?ql elqrssodq tI '(Z'I S3$ 'I'Z'16) slueru 7 uoqrrpnleuur (6961 alpprw euos Jo ernlrnrs eqJ 916I S 'uosl8ueg -a?u€Jtzr€lrurrs ur rnmo sqlauSoleeqJlueJag 'l€-61 nra' uoa eddn:8ue1uop ZfV uagurypuoqqv qleol eqJ, (t't s8g'I'z'ld) ses"qrleql 13 puv pue f ) 'renBl tr aq)q{oqJsuassloag nu1lag (Sunlertlld Jo uart\e-I 'sserdsesue) e?qneFo71)ueluopouocna pun uetuopouoJ ueqc tc?luoJ ur slueluela relulls ,{re,r Sursuduroc v Jo iarJos IP.rSoloaC -suqlue)lueqJsr,{rz ua8unqcrezeg 136I C 'serpuv fllensn '(196J srlpuv '086I Dts,t'PIuEzS) lrytuoa1o4 a$tqatotul 'z 'uolqruE sJelsnlssE Jncco seIxIlaIIIos slueuele luopouoc IJU:qsluopouor -qlnos Jo dllsJe^Iun 's|zb4sqv '5861 lu0q -eJ?deIueAnI'seurds Surds?J? Jleql ueql reql€r 're unsoduttrg uDad dets Z96I I ) thtr -Sutnon'(tt SOC luopouoC 'sq1eu8o1aeqc qlaal eql ol ed?qsu lelurls III asaC -on4 qunod:(spe) qrrus pue 'u4snv 'I Jo lHSi {otrs d I I oft slueuJele u?rJqrx?3eITIOS lecimens for his- Dzik,J. andDrygant, D. M. 1986.The apparatusof Miiller, K. J. and Nogami, Y. panderodontidconodonts. Lethaia 19 133-141. function of conodonts.24th IntetnationalGeolog' ft:=:: GrsleFosse, Univer- Ellison, S. P. h 1944. The composition of cono- ical Congress,Montrea| Section7, 20-27 n: l: Srefan Bengtson, donls.Iournal of Paleontology18 133-140. Pander, C. H. 1856. Monogtaphie der fossilen r-r::: =railable their SEM Fehraus, L. E. and Fihreus-van Ree, G. E. 1985. Fische des Silurischen Systems der Russisch- Akademie der Wis- r. --: l::etson. Dr Dietmar Histomorphologyof soft tissueconodont matrix. BahischenGouvememenls, senschaften,St. Petersburg1-91, 7 pls. :r=:- 3:rlin. and Dr Jerzy In: R. J. Aldridge,R. L. Austin,and M. P. Smith (eds\ Foutth European Conodont Symposium Pietzner,H., Vahl, J., Werner, H., and Ziegler, W. kr :::- \\'arsaw,read the (ECOS N), Nottingham, 1985, Abstructs, Uni- 1968. Ztu chemischenZusammensetzung und Er-- r:::t! r aluable com- versityof Southampton,10-11. Mikromorphologie der Conodonten.Palaeonto- [ nsuisticallyby Dr Fihreus, L. E. and FAhrreus-vanRee, G. E. BraphicaA l2a 11,5-1,52. L-=-:-:: " ::.,r of \ottingham. I 1987.Soft tissuematrix of decalcifiedpectiniform Repetski, J. E. and Szaniawski,H. 1981. Paleobiologic interpretation of Cambrian and | -.: r [hem. elementsof Hindeodella confluens(Conodotta, --i Silurim).In: R. J. Aldridge (ed.):Palaeobiology of earliestOrdovician conodont natural assemblages. Conodonts, Ellis Horwood, Chichester,Sussex, In: M. E. Taylor (ed.): Shortpapen for thesecond 105-110. International Symposiumon the Cambrian Sys- Gross,W. 1957.Uber die Basisder Conodonten. tem, 1981, U.S. Geological Survey Open-file PaliiontologischeZeitschrifi 31 78 91. Report 81-7 43, 169-172. Hyman, L. H. 1959. The inveftebrutes:Volume 5, Rietschel,S. 19'73.Zv Deutung der Conodonten. F.- --- E G. 1986. Cono- Smaller CoelomateGroaps. McGraw-Hill Book Natur und Museum103 409-418. Ei: .- : \1 H. \itecki (eds): Co., New York, 783pp. Schram,F. R. 1973. Pseudocoelomatesarld a EJ l=;:r; Unilersity Press. Lindstrom, M. 1964. Conodonts.Elsevier, Amster- nemertinefrom the Illinois Pennsylvanian.Joal- I = :l . Cl3rkson,E. N. K., pp. nal of Paleontology47 985-989. --:= dam, 196 ll!: attinities of cono- Seddon,G. and Sweet,W. C. 1971.An ecologic - _-:: Lindstrom, M. and Ziegler, W. 1971. Feinstruk- EE Carboniferous of turellg Ultersuchungen an Conodonten. 1. Die model for aotodonLs.Journal of Paleontology45 E- -::'::--.: l9 :i9-29L Uberfamilie Panderodontacea. Geologica et 869-880. +l: C. and Smith, M. P. Palaeo nto lo pica 5 9-33. Szaniawski,H. 1971. New speciesof Upper Cam- E r:: a-:-_::on ol panderodon- Miiller. K. J. 1959. Kambrische Conodonten. brian conodonts from Poland. Acta Palaeon- ac:-: .:: R. J. Aldridge, R. Zeitschrilt der Deutschen Geologischen tologicaPolonica 16 401-413. P :----: red:)r Fourth Euro- Gesellschaltlll 434-485. Szaniawski,H. 1980. Fused clustersof paracono- E, -.-. ECOS IV), NottinT- Miiller, K. J. 1962. Supplementto systematicsof donts.In: H. P. Schonlaub(ed.): Guidebook,ab- l:- - :r:l ersir-,* of south- conodonts. In: R. C. Moore (ed.): Treatiseoh sbacls, SecondEuropean Conodont Symposium Inv ertebrcte Paleontology, Part W, Miscellanea. (EC)S II), Abhandlungen der Geologischen E : --i:: z\ischen kambris- GeologicalSociety of America and University of Bufldesanstalt35 211. D: . -: - :..ionren (Vorleufige KansasPress, Lawrence, Kansas. W246 W249. Szaniawski.H. 1982.Chaetognath grasping spines ;i,F".- C:oxissenschaftliche Miiller, K. J. and Andres,D. 1976. Eine Cono- recognized among Cambrian protoconodonts. t:: : dontengruppevorl Prooneotodustenuis (|u{:ijller, Joumal of Paleontology56 806 810. fh ,-----.: e of some Middle 1959) in natiirlichen Zusammenhangaus dem Szaniawski,H. 1983. Structureof protoconodonts. G- r:: ::e earl\ evolution of Oberen Kambrium von Schweden. Palaon- Fossilsand Strata 15 2i,-2'7. E .-:1.don Lethaia 9 possible --: tologischeZeitschrift 50 193-200. Szaniawski,H. 1984. Structureand origin paraconodonts.27 th Intemational Geological j- Miiller, K. J. and Nogami, Y. 1971. Uber den of a-::-:--!al model for the Feinbau der Conodonter Memoirs of the Faculty Congress,Moscow, Abstacts 9 (supplemelt 2), r -z:--- 16 33. of Science, Kyoto Uhiversity, Series of Geology 64-65. I're :.: r :i:ron of the Con- and Mineralogy3E 87 pp. l-i--.-: 15,<-19. ! l-: -:: :asal structure of tlt- :J5:. 1959 and r =---ir:r :nd Vehl), 1934. ' ll : : t :..:'::zn J9 39-57. t- L:: l-::-.iord.,\. 1983. The |!FEr:: l: _le rieth and grasp- E;-:. .- :,-|nl of the Maine t, .!E .: ':::ed kngdom 63 \ K. and Aldridge, Lethaia 16 -:.:=al Re-examinationof Silurian conodontclusters from Northern Indiana R. S. Nicolland C. B. Rexroad ABSTRACT tional 54 conodont clusters from six rnore Examination of fused conodont clusters localities in northern Indiana. All but five of belonging to the genus Ozarkod.ina from Pollock's specimenswere obtained from the Silurian strata in northern Indiana, including Kokomo Limestone Member of the Salina those describedby Pollock (1969), demon- Formation at the abandoned France Stone stratesthat thesespecies conform to the general Company quarry (Fig. 3.1) at Kenneth, Cass septimembrateapparatus structure described County,Indiana, E1l2 SWU4 SE1/4 sec.30, for most middle Palaeozoicconodont species. T.27N., R. 1E. In this studywe haveexamined original collec- The present study is basedon extensivere- tions and new collections of both fused and collecting of the quarry site (Fig. 3.2) from discreteelements. Both typesof material indi- which more than 700 additional-clusterswere cate that Silurian speciesof Ozarkodina pos- recovered.Some of the material is very frag- sessedseptimembrate apparatuses containing mented, and element identificationis difficult. 15 elements. The clusters have an anterior Elements of Ozsrkodina are relatively thin, complex of discernenselements followed by and sedimentcompaction appears to havebeen the contundenselements. Crystal overgrowths responsiblefor much of the elementbreakage. obscurethe fine detail of someof the clusters, Other specimensare so overgrownby second- but enough good specimens have been ary phosphatecrystals that it is not possibleto examined to provide an interpretation of the accuratelycount or identify the elementsin the apparans structure. clusters.A generalproblem with all studiesof cluster material is the obscuringof diagnostic parts of some elementsby the stackingof the elementsin the cluster,and the more complete 3.1 INTRODUCTION clustersof this study all presentthis problem. Fused conodont clusterswere first described To circumvent some of these problems we from Silurian rocks of northern Indiana (Rex- have examinedboth clustersand discreteel€- road and Nicoll 1964) with the recoveryoftwo mentsfrom samplesused in this study.Because fused element pairs. Both of these pairs are the conodontfauna is of limited diversity (only now assignedto Ozarkodina eosteinhornensis five species)we have been able to determine (Walliser). Pollock (1969) describedan addi- the multielement associations using mor- q or lEedd€ eJeql sJel eleredes ol alqeun ueeq e^eq aA\ slueruele Jer{to aql pue 'luepunqe ,{1enba lnoqe al€ Aou riaserd ra$eq eql uI qd ourospup slueur:LJe4 aql:o; tdacx3 'ds p pue rtp[ous ' g 'satreds luepunqz lsoru 'sluaurele '(Of 3$'f e 14)ralsnlc arll s ssuautotlulatsoa Dulpouvzo uollnqul I arB stuatuale perJ4uepl pr1-redauo f,1uouror; ua\ou{ $ lI sluetrrala -srp sarradsuo suollP^lasqo.rraqlpaSueq.r tou psorxe)I puE I B SluasaJdeJsluaurela suouJecslpeql Jo selcrluapSurpuels ee4 'punol seq Eurlcelloc-arrno pu?'(€16I) E s.:lJollod sad,(l luaur pue ,(Soloqdrorulsnqor aql ,{q peunuralep 1co11o4,{q peqlJcsep ueeq s?q ,{lq"Jol ralsnlc qs €uusrp o1 11ncr15rpf,rea ,{gsee,{1aa.r1e1ar sr snleredde 'nou ds nurpol aql uro{ ?un?J lueuele elaJcslP aqJ, Aou '(r:slflElA\ 'O '(lesrl r sluaurele Jo qsnJJ eql -.{rzo eql Jo uolllsodurocsql alqrssoduruoIlIS 'O pu? vsu)utotluPtsoa eS aq o1 lqSnoqt -oduroc luarualegFuJ Jreqt Jo srs^1eu?ap"ur -1e1,6)ttp\ous outpo1tozg "ds Zsnpollsdo1 ''ds auo ol Erllela-tr31a raqlo eql pu? amq 2snpoltsdoqpue D apopg lo acuepunq€ ollapopg :sercedslueuralalllnur aAIJJo iro azruSocer a,tr saseJ ,ro1 ,{re,r eql pue sretsnJt pesnJ Jo Ire'I PeJreJeJeq u"3 peuul?xe slualuele aleJcslp llv :l pelJlluapl slueur 'oJBJsl.^ou'ds'O pue'sJelsnlceql uI lue "\Bq SJNSI,{U'IS SIf,U]SI(I Z'€ I Pue slua{xele DS eeJql -ururopsr s$uautotlula$oao aJEl 313selJads ft) tJolod f,q lq8noqt '(sretsnlrrll$ seldues ) locrp fq peqlrrsep selErrpur*) ,{rrEnb r(uedulo3 euols eruelC PeuopuPqe se ,{cuanb 'sueturcads ro; + m PunoJ eql Jo eceJ uretsEoquou eql uo{ uollelllro{ ?ulJES Pern8r; 1e rE aql sr srql z: ed :qt urs1e,u:;ur :1dues Suraoqs uortsss leuurnloJ Z f 3!l suolluueldxe a1e1daqt ut pepnlrur e:e (sasaqt z:qS't:rs'I=3s -uared ur) suoqeu8tsep aldues pue ,$rpco1 { :ret{ tsnu (g g 8ts) aA€l .r0lEM pue (slr8rp er'r3) sraqurnu f,rolrsodag suorl 4 ur ssd,ft luaurele -ca1oc fe,ung lectSoloag euztpul-(lrsre,r t snoqs qcrqlAIeIJeleuI ; -ru1 ?uerpq aql ur pelrsoder sr ptreleru aq1 11 -sad,{tluaruela ma,res e 'ser:eds luereglp Jo Eueur 0 JD snleJ?dd? eqt t"ql -a1e;o ?urxrur i(ue paAJesqo 3,{\ a^eq sJelsn[3 E 'srelsnlc g pue (796y) oua,{n uotlPullo-.1 9ql Jo euou uI aql ul slueuale Jo 9 r agr ;o uorldn-rsrpa1a1d eulles ecuesard eql ol uoIlIPP" uI BIJoITJJpcrSoloqd 'uorlrsod L ;r FuorpunJ 8 rp eJu"qrnlsp lo Frrr 6 'Euerpulprluec-quou 'fulenb ,{uEd srD 3ruosleql salBJrpuT -uroJeuors oruErl pruopueqe rql lo uorleJo-l-If 3ll aql 0l 1a 1o Suruorlrsod ti l ir ipo Sursrldurocasoql f ----r-----r ---- |^]I r.us sntereddealalduroc -f I ur) 9t 0 7. -|' --t rg a3uer sJalsnlceql t 6 oulolo) 7 o s-\OIJYUSCISNOC q t \J :SUSISO'IJ g'€ -r i lI|5 I l OUVMO--I H --l I I I 6 x ssalun , ed rn?luoc,{eql OL a ilJe:aue3 satcedso,nl -uaql alerzdes o1 pele tl lxr arEslueuele sueuJeJ l nla,{ zl - q t! 'as€r aqr sI slql Jl uodsue60-j Eoloqfuoruyo uorleper8 et lpsr (Er6I p€orxed pue , Jo sluauraleed eql Jo 3 qlcaa$o?'O pua rtp[Dus 3o stuauodruocelercsrp ll lV I U\l Jd) uBlrnlls uollBu.rluBxa-eu It t € qJl BrrBlpul uJequou u.lo{ sralsnlJ luopouoJ Jo 09 r|Lfftr Indiana [ch.3 s-n ? 1l Clusters: general considerations 51 discretecomponents of the apparatusesof O. snajdri and O. eosteinhornensr's.Examination of the Pa elementsof these species(Pollock andRexroad 1973)indicates a nearly complete gradationof morphologybetween the two, and if this is the case,it is probable that the dis- cernenselements are not sufficientlydifferenti- ated to separatethem. Thus clustersof these two speciesgenerally cannot be distinguished unlessthey contain Pa elements. Fig. 3 3-Plan showing atrangementof discernensand contundenselements ln Ozarkodmd 3.3 CLUSTERS:GENERAL CONSIDERATIONS apparatus preservation: piled up, opened out, The clustersrange from those that contain a or chaotic. We think that the first two of these completeapparatus structure of 15 elementsto are a reflection of the position of the animal thosecompdsing only a singlepair of elements. after its death, and that clusters of these types The positioning of elements in the clusters have been subjected to only minimal disturb- indicatesthat somemust havehad only a rnini- ance (Fig. 3.4). Chaotic preservation reflects mal disturbance of the elements from their disturbance of the elements either by biological functional position, while others show com- activity or cuffent action. plete disruption of the element sequence. In cross-section the conodont animal was Uyeno (1982) and Nicoll (1985) suggested most probably laterally compressed (Briggs that the apparatvs oI Ozarkodind contains etal- 1983). Thus, if after death the animal sevenelement types. This is confirmed by our settled on a flat bottom, it will usually have material which shows that the distribution of come to rest on its side. Sometimes, however, element types in the Ozqrkodina animal the animal will have settled on its dorsal or (Fig. 3.3) must have beenas follows:M : 2, ventral surface. As the soft tissue of the animal Sa ,l,Sc=4, Sb =2,5d=2. Pb=2.and decayed the elements will gradually have col- !e, : .:: .::3mple intervalsin the Pa = 2. This is the sameratio of element fre- lapsed onto the surface of the sediment. If the - ft -: .:: ieastern face of the quency as found in the Polygnqthus clvsterc animal was restins on its dorsal or ventral ts,|IE r-:=.! quarry (* indicates _:_.:ar!,. (1985). clusters fr:: i described by Nicoll Those thought by Pollock (1969) to contain two or O :: ::2,:hornensis is domin- three Sa elementsand that with three Pb ele- I i -- -r. sp. nov. is rare. mentsidentified havebeen re-examined.In all o s--..=:. and the very low caseswe recognizeonly a single Sa element, oo h;e =d Dapsilod.us? have -: and the other elements that Pollock had \rll 'lTII i :< - ::ultielement compo- thought to be Sa elementsare Sb elements.In ltri/il T':t :,'=prosition of the Ozar- the crushof elementsin someclusters it can be Ttt .-i:r is relatively easily very difficult to distinguishbetween these ele- k : r --rsr morphology and ment types.Pollock's example with multiple Pb sc Sc Sb Sd SdSbScSc 4z rs :,- :iles of the discernens elementsrepresents a normal cluster; the mis- \\-//zz: sb r:-"8: :r!rm only one paftial identified elementsare broken Sc and Sb or Sd !:: elements. r :-- :::irents and some Pb Fig. 3.4-Cross-section of dead conodont animal showing In the better preserved,more complete,clus- method of formation of (a) opened out and (b) piled E :Ea:r unable to separate ters there aDDearto be three basic forms of clusters. 'sluaruele ,A\ah cs o/l! pue qs euo sepnpur lelel€l ,.^ou '[JS 'I] '0t89I 'sluarrjelasueurersrp Jo retsnp Jo sqdprSolrruoloqdoerels -ds Durpo\rozo-\| .?tt <6289Ilrled .[-Is !I] '^{er^Ieralel'[-Ig.I] luJruslsed.6.^\rr^ leralet '8289I 'rred lueurale EiJ'g sqdeforJruotoqd orlols (t96I rasrtle^\) rrplrus outpolbzo-6-g sAtJ rlt ,I] ,/zggl ,rred '^,\ar^Ierarel tuetueleed sqdslSolsnuoloqdoeJels_/ .8rc .szggl Jo,luetuala PrelEI gzggl luetuele rs 'g ,{er^ Ieral€l cs 'g ,rer^ louelsod ,tzggl lueuela ^{er^ ,g lluaurele W'i Ieretpl'€Zg9I lueulela qS ,{rer^lprelEl,ZZggI pS,Z lueuala ^rar^ ^{ehlouatsod,lzggl BS 'I ['I9 'I] Ip 'stuatualesueurersrp elerrsrp '(7961 req1pA) esuautoquD\oaDupol.tozo-g I sB]J 'lS eldlllEs'I ,qJlecolseteJrpur [-IS'I] 'requnu tuot$oder SCI-nI sr requrnu16rp o^tg 's9x [v-I € alPld Jeqlo er{l Jo flr^€c pseq eqt Jo drl Supu€dxe ueql uourrxoJ erour er€ srelsnlc pelrd .sJelsnlJ eql Suqcnol lsourp luorrrela euo Jo salJ paJe^oret eql ur suorlnqrJlsrp pa^Jasqo eql -nuap eql qlur pedd?IJe^oele.ezs eurEseql qlrd\ lualsrsuoc eJ" lueula^orrj lueuJele IIle out eJe JO Slueuele o^u eql'pauruexe seldu"xe -1sod yo suoqelardralur lecrleroaql aseqa IIE uI stueuale ?d Jo red B Jo pasodruoc '(tt'Eg'f f f 1d'6961 qco11o4)stuoruele sr sed,{1 relsnlc uoururoc alour eql Jo euo ed eql puqeq pelerol sluaurole qd eql e^?q 'relsnls o1 reedde lpqt sluauala qd pu? pd Jo sratsnlJ eleldtuoc ? ur punoJ eq pFo^\ leql sluauele ;o ,{rerrocar uotuutoJ ,{le^rlElar aq1 sureldxa 9I eqr Jo zI roJ slunocrs srqJ relsnlJ eql Jo srql sluJu:lJ Jql Jo lsa: aql uo.r1,(e,tre pe;oed pu9_Jouelsod aql leeu punoJ sr slueuela qd Jo Buraq ;o sseroJd eql ut pe,rraserd lueruele JrPd3 pue 'aprs Uel eql uo pa^JesaJdsr lueurela qd 9uo pu? slueuela ed o^u aql qlr/il peJeAoJ Ii{ el8urs v slueuela reqlo eql qteaueq rxo{ -ar sp^\ relsnp pegd sy11 Z'zrJ 'ZE ld, lno iur{Jrls uoes eq ueJ lueurela eql Jo lJ?d uo u^roqs relsnp eql ,{q pe,ryosar sr sdqs i(1uolece;:ns -repuneqt uo snteJeddBaqt Jo pue -uorl?loJ ]uerrrela ;o uorleueldxa raqlo auo JOrJelu€aql lE palesol sI lueruala e5 el3urs eqa 'suoqrsod Jo 'slueruela cs Jo sll"d olal ,(q pa,nolo; sluaruale e olur uell"J e^Eq plno^\ 'purru€ eql qs o1(rl eqt or3 sluelxale ps eql qleaurepun JO^lelre^ Srx" eql Ol esrelsusJl sex€ 8uo[ ]reql qlr,!\ pue aprslng snteJ€ddp aqt Jo auq l"rlueJ aql 'Eueurele es pu€ I^{ eql asPFns luatulpas aql Jo eprs reqlre uo pel?col slueruale p5 ;o led e ot ,{llBJalsl uelleJ e^eq osl? plno^{ tnq tueurale s"q pue 'I'?tJ'Z'E'I1, uo uaoqs sr eldruexa Sutsoddo oqt spr€^lot pe,roru e,req dlqrssod lseJ?elr aql sralsnlJ lno pauado eq1 ur uaas pFo^l stueuele suepunluor aql '(q? € 3lg) lseq sr slueuale sueu.racsrp;olueruaSueJJe eqJ uorpaJrp euo ur Surlurod seurl€r Ips€q ar{l s?'J'7't 'td uo u^roqs saldl lvxfl :suglsn.IS Jo IIe w\N"L'g'z t'€ sJelsnlf,o] repurs e1d z olur ,{l1erelpl sserduroJ ol pePuel e^Pq plno^\ sluelllele sueuJeJsrp '{coJ aql uro{ uraql alorrreJ ol Sursseco:d eql 'aprs slr uo lseJ ol aues lpurru? aql ueql\ Surrnp uelorq aq {pur fueur pue 'elr8er; . Jo^ '(y '7 '1 s3t1'7y 1 14 '6961) po1o4 osle eJ" sJalsnls aql sJelsnlc Jo uorlprxJoJ aql ,{q pue 'y '€ 'I s8g 'Z'E Id. ao u^\oqs esoql lueAerd pue sluetuela aql aleredss ol sluerJnc se pea,reserdsralsnlJ ur IFSoJ plno^{ slql 'suoll to s.taSue,lecsfq acueqrnlsrp Jo lunouJe II€us -tsod puorpun; raql ur peuErxeJ e^?q pJno,!\ e el€l r(luo pFoa lr'stueulela Sururo[pe qtr^\ 'dlesrarrsuerl peluer-ro Sureq'sluaurela suap ur gql lt€luoJ Sureq sluauala uo spuedep Jel -unluor eql (el'g ?tS) snleredde eqt Jo eurT -snlJE sEslueuralJ otll Jo uorle,r:ssardasner:g eJlueo eql tuo{ ,{e,,!\€II€J ol papuet e^eg plno^\ 'pe,uesard sluerrrole ,rreJ e ,{po qllt a1a1d anssq ,(q peurpJlsuoJ lou ueq/'(\slueuala sueu -uroJurJo JlloeqJ elg slelsnlJ Jo ftuoleu aq1 -rersrp eql ueqt '(E961) 1ocr1,q,(q 1ptxrupeql Jo 'a1qeue,r,{p3rq sr sralsnp pald ur stueruele eS uorpnJlsuoJeJ eql uI pesodoJd lpql ol lplurs puE tr^lJo uortBJo[ eq] pu? 'srelsnlo lno peuedo uorlrsod e ur eJar{ slueuale eql F pu? 'ecEJJns 'Jrs [€ t € qcl euBlpul uJoquou rllo{ sJalsnlJ luopouoJ uplrn|ls Jo uoIlBulutBxo-au z9 r*rE Itrdiana lch.3 Clusters:general considerations r-<, r--: :he location of M and b: : -':ers is highly variable. fu-.': : :re chaotic or incom- E :-: elements preseNed. h-'r r: :ie elementsas a clus- Le . ::::nrs being in contact r[.:--> r: sould only take a &:--:::.-e bv scavengersor t --:a :-3rnentsand prevent &u:::. The clusters are also ali: 1:\ be broken during F. a :::r-: trom the rock. i E\{\[PLES cr :----;-ens elements is best c :-: : :sters. The clearest r :. ? -: i. fig. 1, and has rr--' t::ed on either side of { == .::::arus. Outside and lc:..:::::: are the two Sb l:". - ::irs of Sc elements. E: -r: t..:ted at the anterior lG _:]]i under surface;only !!r: := re :een sticking out cr- = : iments. A single M ic; . '-:e leit side, and a pair fr,-:: ::ar rhe posterior end E i::- -:tis for i2 of the 15 rr- :r :.rund in a complete re ::r=:Don cluster tyPes is F._- : - Pa elements. In all ![- -ia ,-r o elements are of : -.: .=::ed with the denti- E : t::lost touching the dE ::!r !'alitv of the other Er.1- =-,rle 5L. r:c:--:::. -:-l 1. 5Ll. 1, Sa _::: E . .-rieralriew 4,M lfiE:_ : :l!, lareral view. l- :. : =-:t: pair. 16828, :E : :-: :i 16,!10.[1,5L], ti , :-urboqs ? :-,:"o:ttozo- L'9 sBtJ r-::: :l sapnlcul relsnlc | -=.'! i ralsnlJ-t'€ sBI{ a-: ::aq a^Eq luetuele :.u:u.lale suepunluoa ILE-:lrp-:':! eruPsq poluauo lI .! -, I ) I,J :lueueSuExe I ::i;: rersnll-S'Z SBIC * = rrueluale r.f-!i:oqd oarels-I Brc 11to:llDzo-9-I s8 g -e ;t | -5rlt€id ,{q peuqlno sear n--i-::s sue:edde clseq eqJ TNAOnNZO tO SSI(IruS ErL$ \Osruvdhloc s'€ r;'1 sBg'711 yd '6961 F:!a :llueJ aql uI luauale F ::uu eql uo $uaurala qs l :{ lo srred o^u ulgluoc leql p ::;-re1'.raaar,ro11 uoululoJ uE:prs reuul aql uo lueul EXDetr?JS 01$ qlIA{ SIaISnIC E rrrrrn aqt ,{lpcr3o1oqd:oru !$$dulr sr lr sas?J qJns uI .rr rq;u o^u reqlls Jo slslsuoJ 'uI uouruot f1a,lt1e1erere d z _gopasodruoc s.re1sn13 nuauela qd puE ed aql eIe em]eur ur f,po pue 'slueurale ra-n sluauala ed ar{l sebeds m.j sr 11 slueruala e4 lse8J"l uE sruaurele q4 1se3-re1agt qi ur pue 'ezls lElrurrs Jo eJ€ oqro u1 rred lueruela qd eql 1 lred tueurela eg eql popnts auros uI slueluele Bd erll 6al 'Pulqeq PeleJol eq ,(?tu Euaurale qd pue ed Jo qcea 6Ie SrelsnlJ Jo Jequnu Y 1'c 14) uoqra4p etrr?s eqt uI Enue lleql e^eq pue 'lueulels suBlpul uraqlJou uro{ sJalsnlJ luopouoJ uB!JnIIS Jo uopeuluBxa-ou hoJ [S E res s'qJl previous Indiara lch.3 Sec.3.5l Comparison with stlu,diesof Ozarkod.ina 55 -kD element,and have their anterior endspointing ognizedApparat H (O. excavata(Branson and in the samedirection (P1.3.1, figs 7-9). Mehl)) and Apparat J (O. steinhornensis A number of clustersalso contain one pair (Zlegler)), Jeppsson (1969, 1.971., 1974) eachof Pa and Pb elements.The Pb elements refined and elaboratedthe multielementcom- may be located behind, beside,or in front of position of several species of Ozarkod.ina the Pa elements. In some of the examples (Hindeodella). Both Walliser and Jeppsson studiedthe Pa elementpair is much larger than thought that the apparatus consisted of six the Pb elementpair. In other clustersthe pairs element types,neither recognizingthe Sd ele- are of similar size,and in the discreteelements ment. the largest Pb elements are as large as the Mashkova(1972) describeda beddingplane largestPa elements.It is possiblethat in these assemblage of Ozarkodina steinhornensis speciesthe Pa elementsgrew fasterthan the Pb which she interpreted as consistingof 13 ele- elements,and only in mature or geronticforms mentswith the Sa elementmissing. With refer- are the Pa and Pb elementsof similar size. ence to Mashkova'sspecimen (1972, fig.Z, Clusterscomposed of a pair of Sc elements pl. 1) we would interpret the assemblageas are relatively common. In all casesth€ pair follows: M elements-l2, 13; Sa ele- consistsof either two right or two left elements. ment-missing; Sc elements-8,9,10,11;Sb In such casesit is impossible to distinguish element-anterior part of 5; Sd element-6; rnorphologicallythe inner and outer elements. Pb elements1,2; Pa elements-3,4.The post- Clusterswith two Sc elementsand an Sb ele- erior paft of element 5 and element 7 are the ment on the inner side are present but not broken posterior parts of the left Sb and Sd common. However, larger clustersare present elements.Elements 5 (anterior),6,8,10and 12 that contain two pairs of Sc elements,a pair of are right side elementsand elements7,5 (post- Sb elementson the inner sides,and a singleSa erior), 9,11 and 13 are left side elements. element in the centre of the group (Pollock Assumingthat the Sa elementwas presentbut 1969,pl. LL2,figs. 1,2). not recovered,the multielement structure of O. steinhornensisis identical to that of O. eosteinhomensis. 3.5 COMPARISON WITH PREVIOUS Uyeno (1982) recognized that O. brevis (Bischoff Ziegler) had a septimembrate STTJDIESOF OZ.INXODINA and apparatus. This was confirmed by Nicoll The basic apparatusstructure of Ozarkodino (1985) from a study of fused clustersand dis- was outlined by Walliser (1964) when he rec- crete elements. Plate3.2-All x65. FrEs 1-5- Ozarkodina eosbinhomensis (Walliser 1964). Fig. l-Stereo photomicrographsof elemenr clusrer, 16831, [1,5L]; inner lateral view showing 12 elementsin openedout arrangement:M (i), Sa(1), Sc (4), Sb (2), Sd (2), Pb (2). Figs.z,5-Cluster 16832,11, 41.2, stereophotomicrographs of lateral view, showrng14 elementsin piled arrangement:M (1), Sa(1), Sc (a), Sb (2), Sd (2), Pb (2), Pa (2); basalcavities of all discernenselements oriented in samedirection (left) but thoseof contundeff elementsdirected away ftom opposingelement; contundenselements show ovidenceof having been partially peeledback, both Pa elementsand one Pb eledent havebeen flipped poste orly but are still attachedto the rest of the cluster.5, basalview. Figs. 3,4 Cluster 16833, [1, 5L]. 3, stereophotomicrographs of inner lateral view. 4, outer lateral view. Cluster includes 13 elementsin opened out *rT"t:ff:*, M (1), Sa (1), Sc (a), Sb (2), Sd (2), Pb (2), Frgs.6,7 -Ozorkodina cl. O. eosteinhonensir(Watliser 1964), Iaterat views of cluster 16834, [1, 5U], showing10 elementsin piled arrangement:M (2), Sa(1), Sc ( ), Sb (1), Sd (2). '(I) qd '(I) r,qs'(Z) cS :srueureto t qlh\'.r3r^ Ieretelrelno'[-]s'I] '0t89I relsnlc'(t96I tes\]..$\)slsuau&qulatsoz oulporlrozo-z'3u '(z)Bd '(z) qd '(z) ps '(z)qs '(r) cs '(r) es '(I) I^l :lueure8uelplno pauedour puepa^rels {pq8rts 'sluetuela sapnlcurrotsnlJ lalno tI ^rer^lpretpl .s?r! '€ Aer^Fretsl rauur'l ['IS 'I] '6t89I retsnlc'(t96I nu11e11)stsuautotlut4soa Dutpoltozo-t I '99x IIV-S g aletd '(z)ps '(r) qs'(r) cs ,(l) nS,(Z) :luerua8uerrepald ur stueurele0I sepnlrulrelsnlJ Fralsl 't IErelelJo sqdEr5or l,,f ^{ar^ ^rar^ -cruroloqdoaDls'z [n9'I] '8€89I .Iotsnl.'(t96I ns41e1j,)stsuautoqutatsoa o lc outpo4tozo +'zs3tf '(z)ps'(z) qs'(r) w :ruatu -e8u€rE pelrd ur sluelllale6 sepnl.urralsnlJ pjelel's'(r) 'S psBq '€ aar^ lelotel Jo sqdel8olcltu ^\ar^ ^1er^ -oloqdoerels'I ln'Il'LtBgI relsnlc'(t96I resle$r) slsuauloqulatsoaO lx oulpo4lDzo ;'E'l s8:c 's9x IIV-+€eleld '(z) ea '(z) qa '(+s) lapu s '(I) es '(z) w :rueue8ue P tno peuedouI slueuele'9r {lqeqord'tI tseel}e sepnlrurrelsnl3 Arer^lerotel Ieuur Jo sqdplSolrruotoqdoerets't lereleltalno Jo ^{er^ sqdErSorrrrroloqdoerels'€ [9'I] '99991 rersnlc'(7961 nst11ep,,\nsuauolul4soa ourpo4nzo---V,t s?rl '(z) ea'(z) qa '(z) ps'(z) qs'(t) cs :ruetue8uerlepelld uI slueuele sapnlJurrelsnlC reuur;o sqderSorJ[uoloqdoerels .Z .^\eh pre1e1relno sqdvr8 ZI ^rer^Ierelel 1o -o.lcltuotoqdo.re$ 'I [.Ig 'I] '9€89I relsnp'(1961 .reqp6) slsuauto4ul4soanupoYtoz6-7'1 s1rg 's9x llv{ € ereH AlluanDa4AJ" SluerrJAlesuepun]uoJ eql oulpol ourpulDzo. 'stuatxale qS eql eprsur 'sesseJ -tozg an tqm uos€er eql eq feur sql ulaqt -ord Ieratel petJemp f,lpft^\ur rreql qtra\ 'Eueur 01resolc palecol eq ol slueuala suepunluoJaql -ala pS eqt ro; eceds al"q lou saop 'lueuala t ,rollu oslepue slueuelosueuleJsrP srll Jo 8ul es eql uo sserord Jouelsod E tllr/r\ snYlDu -dnor8 lcedurocaJorrr e ur llnsal plno,r sluau -BK1o41uaue1a pS eql uo ssaJoJd Jouelsod -ela ps puB qs er{l Jo drqsuorlelerpuoqrsod p Jo eJuesqe -ro ecuaserd eq] ol pat?lal ,(lq? srql 'lueurele qS eql sdEIJe^oteqt uoursod -qord are sasnleredde snlpou?Klod pua ourpol e ur Pat€collueuale ps aql e^eq ueJ lueuala -tozo uee,^laq sacualeJJrp IeJnpnJls esaql €S eql uo ssacordroualsod B {JBI leql sabeds '(S8et floctN) s]uauala sueurersrp Jo 8ur -dnor8 roleru aql purqeq petecol eq o1 'nqtou3(1o4 pue Durpottozo uI lqSnoql pg srupoug{1od uI '(g'I SEIJ surellEduoqnqulsrp tuauiele eql Jo uosueduoJ-S € 3lJ ar" slueuala erp 'Z €'ld) xelduroc tueurala suauracslp sorullsoJ lcedruoc e Jo U?d I?r8elur uB s? sluerrJale qS aql aprsur peterol aJe sluauala ps eql oulpol -toz6 u1 'srugou8tlod ur pe^rasqo l"ql ulo4 lueJeJJrp l€rl/i\auros eq 01 sJ€edde slueruele S eql Jo drqsuon€ler aql 're^eaoH (S'g'Atg) roualsod sluollIale Pd aqt pue JoHelu€ slueur -ela IAI eql wa mqtDuStlod ur s3 aJuenbes luauele aurBs eql A{oqs sreTsnlx Dulpo4tozo -Jqs's,s eq1 sadft luaurala ueles stueuele qlra\ "s,sqsl- [_1qs qs Jo SI cs.s rs .s (EA6f IIo"lN) sla1x1t snqlouS(.1od ur punoJ lILlll lpql ol ftIrurs ,{p€oJq sr Du?poltDzo lo uJ?l qAT P?P -1ed uoqnqr:1srp pu€ ecu€punqe lueuele eql SNHJVNOTTOI H itl snqleu6Alod :flunJJnurssnrvuYddY 9'g ;:reddy [9 € 'res € qcl BrrElpul uraq ou urorJ sJalsqJ luopouoJ rrBlJnlls Jo uollBulru"xa-ou 99 rrlEftr lndiana [ch.3 Sec.3.6l Apparatus structure: Ozarkodina compared wiahpoltgnathus ozarkod ina ..1,-::: Y-2 o-{ 'oo / \\ \ _r'? ??? r-?|lllll | : ::sbl sbscsc tsd sd r x -:. :::::tr dlsrributronpatterns 6--: ,:: .,.!nathus. - l:r:i:-::or processon the sa r E S: locatedin a d;:s --::-lement Sb element. This r: :: :he Sb and Sd ele- t . : =tre compactgroup- E-* .--:tenrs and alsoallow ca ,. --r:.rbe locatedcloser to be --:: ::ason why tn Ozar- rE-. -=.::tentsare frequently -:- - .::reo photomicro- C!---:- ::.-dei 12elements t- - : . =-.: :\olomicrographs !t:;::::: earl13, probably F.:::.P3(l) il:- : : 1. irereo photo * : ::::'r ur piledarrange- :: :-- _ :. !:3reo photomic- E :--::-3:rr: \1 (2),Sa (1), t GI , :!: . :::ral view, with 4 r|BJeddV [9 € 'Jrs € qcl ?UB|pUJ UJAqIJOUrrro{ sJolSnIJ luOpOuOJUEIJnIIS Jo uOrlBU[UBxo-alI rthtrn Indiana fch. 3 Sec.3.61 Apparatus structure: Omrkodina compared with Polygnathus rno ut uoruuror ,{1a,rr1e1arare sralsnlo'e1a1d eseql'sel"lo pssq eql punor" eql Jo "erE -urot flJeeu Jo 'elalduof, pue i snqlou8,t1o4 1o fi1ppe eq1pas?anur ,(1lqE1s e,req plnoc 'e1eyd snleredde aql u€ql lr"duroJ eroru sr snleredde pseq eql Jo ernlcn4s crleqdsoqdeql uqll^\ aql slueuele 9I surctuor pu€ alerqurarugdes pualeur rrue8ro aql uo .{fletJedse'leurrue luop sr DurpotltDzo Jo eJnpruls snl€J?dde eql -ouor aql Jo sureuraleql uo {celle puepBg 'ed{l luaurala qJEa Jo sJequrnu peFadxe 'sursrueEroreFllerp1nru,{q 8ur8ua,,lecs eql usql eJorrr 8ururcluor srslsnlJ puq lou op elqrssodpue uollr? lueJrnc qloq o1 salldde e^{ JsneJeqpue sJalsnlt eql ul yred luauele sql peuroJ aaeq lou pFm sJelsnlc eql Jo {tuelulrs eZIs eql IIlo{ e8pnl e.t\ srrlJ Teur pu? 'sluaureleaql paleredesaaeq plnoa{ uoq? -ruE Juo uErlt erour ruo{ sluaulale 3o Surxrtu -lueluof, oroJeq pue I?lurrJs eql Jo qleep eql Jo ecuapr^e du" s,t\oqs stalsnlJ erll Jo euoN JauEslueuele luoPoum eql Jo ecuPqrnlslpl"c 'loorulpeseql ur uorlelodroJur -rueqf,au ,{uV 'l?unq raUE aunl errJosluaulpes raql relJ? errq erros 1e palelrdrcard uaeq spq aq1ur aoeld1oo1 sralsnlJ aql Jo uo9?lueueJ teql aleqdsoqd d:epuoces dq raq1a3o1plaq ere 'IeuJru?eql UI peuorlJunJsluaruele eql ralsnlJ eql Jo slueIUelaaqJ'suatUlJeds ala:cslp ueq,r luasarduaaq e,reqlsnur salqd Fs?q leql se peJe^ocer sq IIrAl rapueuel eql pue 'srel elpJrpur IBrJal?u clraua8uoc ur sa1e1dpseq -snlc eql ur pe,uese-rdaJp lueurpes dq peunq go uorle,rreserdaql pu€ ,{SoloqdJorrrlueruelg ueaq e^"q,{eq1 rage stuelIlola luecefpe q1r,n 'Surssrurare sa1e1dpseq eql pue'pa,r:aserd tc?luoo ur eJp leql slueuele esoql dpo pur eJe suarorceq1 dpo'sralsn1c aql uI slueuela -rue euo,{u? Ixorg eql eeft qlnou elrdsaq |, lrDus puo s/r,rsoJ sluop lEruruu Jo luopouoJJo uo4e,r:esard1ue11ecxe eql .I D i]-alilsv t/6I 'uossddef eql ur punoJ ,{llzurErro stuauralo aql Jo '^\eJ olDIta1 sulroJ t€F(urs e ,{luo Jo 'll? urEluoc fzur relsnlc qcea sl?rllluE SSIY'Id 'IYSVtr CNISSIW fol t ''€ trdH Jo sesnlPPdde luop luopouoJ Jo sulBUeJ Ju1e1a1spa,r.resetd aqt r n::Belf .I ruossddet It6I luesaJdeJ,{pnls sql ur pautrxexe sJalsnlc eql E.'rtlpuottJol a aloqlJols I 'uoqrsod 19 -!-ruopouor lueulelerllnur SNOISn'ICNOC 8't r! i:ro-\ '696I 'I 'uossddef rplnurs e ur pauoqJunJ sluelllele eql aJII uI l"ql 'nvr uoDpf,rpuluE sr srrll a^JrlJq alA 'deFa,rosace,; eql lrT\1ooJ €86I f'x -Jnsl3ro eql leql os pel"col eft aseql 'paNes = -Eoqretl'c a c's33ug 'uot1e31lsa,rut.;o -erd s1uaruelaqd ro/pue EA W^ oulpolqozo ssJNsf,s.{flu ee.re Surlseralur uE sE srql lsaEEns ar11'se1e1d Jo srelsnp eqt Jo Ip lsourleul '(9961) locrp dq pseq Sursstu aql Jo el"J er{t lnoqe uorsnlJ peururexe snqlDuStlod lo stetsnlc eql elrTun -uoJ ou qceer u€J aA{ IEllal€lu Jno lloJd 'uo4ern8rluocalqels e ur sluerrrele 'sa1e1d '!-rrsiqdoec pue pseq pa,r-resardseq,(1arer qcq,n aql ploq ol goddns 1zra1e1ol{l qll/$ tueurala 'Durpoliozo ueql uor1€zr1eJeururororu ea?q ou s?/( eJeqlasneJeq's$l lo] elqlsuodsar'u?d ll ro nBaJng u"rlBJlsnv^Eoloac f?Ix 'lprJeleul a1e1dpseq surc1er,{puanber; uI .eq f,etu eS aq1 uo ssacordro a Fe -iaa-rng pcrSoyoeg luaurala r]f,lrlrr,l.'snpo!,rJl 'snql 'seneds Jeqlo uI ueql -tsode;o qcel aq1 'suoDrsod palxns ) nPrs aql fq pazuoqlne leuoucunJ pezqereuru ssal eJeA\serceds -ro etaua8 aruos -ard lno pu" pauolsp readde slueur ll.rr,luef 'seJJnosell l"re 4aql Jo sa1e1d eql leql sq ferll JotJ"J -ele eql'dpnls srqt sratsnlf,po^resord-lseq {il\.I rnI{UV Jo {ro.{r eIIl ;o leseq leuonlppP Jo uY 'su.noJr eql ueql pe^lossp ,(1rse::.rou aqt ur 'uooJelloJ u"rl?Jlsnv aql ul p$ pue ,{qdefoloqd WgS ldarxg uaeq al?r{ ppom se1eldps?q aql Jo ernttnrls ra11aqsr sdrqsuogelerluauale yo uorle,r-reserd sL\fhrscos'IAtoN)rJY snoroduado arour aql pue'crprcsdllq8rls ueaq oql elrqa\ ,1eualeuteuerpul aql ur tuepunqe er'eq feru {coJ eqt Jo sacedserod aq1qEnorql eJorr aJe srelsnlcalayduoc leqf s,noqs(9961 ,9961) ?ur,rour sprng lzql sr,{1pq1ssod raqtouy 11ocrp[q peqursepasoql qlr^\,{pnts srql aliE rrroltoq eas eql uo PrrJ 'raqlaSol srelsnlJeql Jo slueluele uI srelsnlcJo uoq?,rrasaldeql 1ouoslreduro3 p uoorsod Surlser pury aq1 aql sPurq tBql eleqdsoqd aql Jo eJrnos eql sru4ou3K1o4ut arEr er? sral q rreJ sJelsnlc eql Jo aruos flqrssod sr l€ueterx pe^lossrpsrqJ 'paalosslp -sn1calalduroc elq^r'sJetsnlc elalduroc,(Feau $uoqeler eq1'suouceloc 'fletalduro3 Jo'^ ulJ€d ueeq e^pq snql lq8!u uI slueuale sueurecsrpol paq33 ? pareAooer € 'qcl euBlpul uJaqlou urorJ sJelmlJ luopouot uBlrqls Jo uogeulurBxa-au 09 tutr Indiana [Ch. 3 References 6l a :;:tially, or completely, collections.The relationshipof the elementsin Mashkova,T. V. 7972. Ozarkodinasteinhotnensis rf,'--;=i material is possibly some of the clusterscan be used to interpret (Ziegler) Apparatus,its Conodontsand Biozone. : E-:hate that binds the the final resting position of the conodont ani- Geologicaet PalaeontoloBicoSBI 81-90. Nicoll, R. S. 1983. Multielementcomposition of the F.-:ogethel. mal on the seabottom after death. conodont lcriodus expansus Branson and Mehl ift- -. that fluids moving ftom the Upper Devonian of the CanningBasin, pr-:- .-rithe rock may have Western Australia. BMR Journal of Austrulian 7 L "t:c --:e more open Porous ACKNOWLEDGEMENTS Geologyand Geophysics 187-213. t- :.:ies rrould have been Nicoll, R. S, 1985. Multielementcomposition of the conodont species Polygnathus xylus tylus Stafi- k: --:al the crowns. An SEM photographyand plate preparationwere the work of Arthur T. Wilson (Bureau of Min- fer, 1940 and Ozarkodina brevis (Bischoff and Er --:ar the basal plates of Ziegle4 1951) from the Upper Devonian of the E:d - rere less mineralized eral Resources, Canberra). Publication is CanningBasin, WesternAustralia. BMR Jou al E+. Ttus. Icriodus, which authorized by the State Geologist, Indiana ofAusttalian Geologyand Geophysics9 733-147. , b,i:-- :late material, may Geological Survey, and the Director, Pollock, C. A. 1969. FusedSilurian conodont clus- ters from Indrara. Journal of Paleontology 43 r[r:r:-.] than Ozarkodina, Australian Bureau of Mineral Resources, Geology and Geophysics. 929-935. R=:-. basal plates. Pollock. C. A. and Rexroad.C. B. 1973. Conodonts d -.i .-al reach no con- from the SalinaFormation and the upper part of -: r fu:= ..i rhe missing basal the WabashFormation (Silurian)in north-central '77-92. a ai 1i an interesting area REFERENCES Indiajia. Geologicaet Palaeontologica7 Rexroad. C. B. and Nicoll. R. S. 1964. A Silurian Briggs,D. E. G., Clarkson,E. N. K., andAldridge, conodont with tetan.us?loumal of Paleontology R. J. 1983.The conodontanimal. Lethaiq 16 3E771-773. 1.-14. Uyeno, T. T. 1982. Systematicconodont paleontol- Jeppsson,L. 1969.Notes on someUpper Silurian ogy.In: Norris, A. W., Uyeno,T. T., and McCabe, f,s multielementconodonts. Geologis ka Fdrcningens H. R. Devonian rocks of the Lake win- i StockholmFdrhandlingar 91 12-24. nipegosis-Lake Manitoba outcrop belt, Man- rhis study represent -: Jeppsson,L. 1971. Element arrangementin cono- iroba. Geobgical Surveyof Canada,Memoir 392 H-: :emains of conodont dont apparatusesof Hindeodella type and in 280pp. E =:r contain all, or only a similar forms. Lethaia 4 lDl-723. Walliser, O. H. 1964. Conodonten des Silurs. rc :::srnall,"" found in the Jeppsson,L. 1974. Aspects of Late Silu an cono- Abharu)lungendes HessischenLandesamtes fi)r donts. r r---. From any one ani- Fossllsand Struta61-54. Bodenforschung41 1-106. kF-- lhat are in contact e-r ier rhey have been I i:: :ieserved in the clus- in:e gill be recovered as . Tre elements of the cluster ry =r',.ndarv phosphate that E .:: :ome time after their ! s-i:Ient. t= ::oqs any evidence of t:-l more than one ani- E i:: rhe size similarity of b ::srers and because we E J:tr3rmng more than the c- :-:ci element type. &-.!-.-:e of Ozarkodina is I c':---:hs 15 elements. The E-- R. J. Aldridge.M. P. Smith.R. D. Norby.and D. E. G. Briggs ABSTRACT Pa element which can be derived, directly or indirectly, from the Pa elernentof Ozarkodina Bedding plane assemblagesof Carboniferous (Klapper, In: Robison 1981, p. W157). The polygnathaceansdisplay a limited number of complete apparatusis normally seximembrate recurrentpatterns, The most common is an or septimembratewith the ramiform elements arrangementin which the ramiform S and M multidenticulate.There is ample evidenceof elements form a sub-parallel cluster set the multielement compositionof membersof obliquely to parallel pairs of pectiniform Pa this group, as the vast majority of known bed- and Pb elements.This pattern is evident in the ding plane assernblagesand many clustersare conodontanimals from Granton, Scotland,and of polygnathaceangenera. Additional import- is hereintermed the'standard pattern'. Other ant information is provided by the first com- repeated arrangementsare designated'paral- plete conodontanimal, from the Carboniferous lel','perpendicular',and'linear'. Assemblages of Scotland, which is a cavusgnathidpolyg- of all thesetypes can be producedby collapse nathacean(Briggs et al.1.983).Hence, much of and flattening of the same three-dimensional the discussionof apparatusstructure and func- structure.In this architecture,the elementsare tion has centred on the polygnathaceanplan. arrangedabout a dorso-ventralplane of sym- The elemental composition of the polyg- metry, with the ramiform elements at the nathaceans may be exemplified by the antedor and set at a steepangle to the long axis apparatusof Gnathodusbilineatus (Roundy), of the animal; the Pb and Pa pairs follow which occursas bedding plane assemblagesin posteriorly, lying vertically and nearly normal the Tyler Formation of central Montana to the long axis of the trunk. An integrated (Norby 1976). This apparatus is septimem- functional system may be postulated for the brate (Fig.4.1), comprisingone pair eachof apparatus,with the anterior elementsgrasping platform (Pa), angulate (Pb), and dolabrate food to be processedby a shearingand grind- ramiform (M) elements,and a set of ramiform ing action of the pectiniform elements. S elements.Within the group of S elementsare four pairs with bipennatemorphology; two of these pairs are designatedSc elements, the 4.1 INTRODUCTION other two Sb and Sd respectively.There is a single, bilaterally symmetrical, alate Sa ele- The superfamily Polygnathaceaencompasses ment. In somerelated species and generaan Sd those conodontswhose aDDaratusesinclude a elementis not differentiated,and there may be :::-.irur (2161) lDta uos -rue uoluerg eq1 ur sasnle:eddeeq1 .{1p1srp -rrroc Jo suorl€4snllr eql leqt pelou eq Plnorls : . r]'qsBtrr'\J'r'9 s8g't96I lsourpelenlrs stuJurele W aql q1r,n':ledq4 aqt 11 snle-reddu aql Jo eJnlJelrqJJ" Ieuorsueurp -_: _t-E SBTJ'92Id'€16I ruo-q,{e,,ne ace; fpouluoc 1nqpasoddo ,{1ensn -eaJrll eql paga: ,{1aso1carou 1q3ru (aot 'Alr s:: "i ilepr^\ uaaq eAEq lou aJe sluaurele urJoJrr["J eql Jo seJSJJns 'ZL6I'p p uosu\llo3) uralled ,uoqelor eldrurs, ::: ': Jo urelledpJ€puBls pel"F rluep aqJ stuelxela qd eql 01 lsasolr aql teql pe1se88ns,{puanbesqns (Arct 'gtOt) r,{-:!se uEec"qleu8,{lod sessocordror:e1sod eql;o sdrl Iplsp eql qlr,tr ,iqro51 (yg1 '3r1 ''lrc do) ,(ele reau11 ,{y1e 'spr€^\ 'el8ue u" le peluerJo sr dno:8 ruro;rurer eq1 -ur8lro ue uro{ pelrJep se suraUed aq1 pelerd =_:. :ql Jo esoqlsEelaq^l\ tnq 'reqto qceeo1 lelpred are s-uedq4 pue e4 -retur oq,ln '(Zt6I\ '1, t" uosurlloJ ,{q f,1snor,,r -: -:-L)J 3q1 JO SelClluap eq1 lueruuSrle1a1lered-qns ur sessacordror-re1 -ard pesn asoql uror] reJJrp pue 'e^rtduosap =': i rue w eql Jo sesseJ -sod Jreql qlrl\ stuerueleS pu€ W ruJoJrrueJ e:e sluatue8uerte uottlttroc eql roJ srrr.relJnO r: :a[elnJrluep Jo suJel ;o dnor8 E pue 'pesoddo selJrlueppue lueJ 'uorlcedruoc/asduyloJ pue qleep ;o sseco.rd -i _ :: aql Jo srxE leurpnl -elpe spua rorJaluer{1rA\ IaIIEred ureBE'sluotu aldrurs e q8norql su-re11edeseql JoJ lunocJe E --: - lere 3uo1 rreql q1r,n -ela qd Jo:rcd e 'uorlrsoddour sap€lqoer1 lsmu sasnle-reddeJo ernlJetrqJre leuorsuaunp u- -::irtr:d aql rorralue palelnJrlueplreqt qlrAr'sluaualae4 lelpred -aerrlt eql Jo lepour e^rt?lerdretu .(uy qleep Lr!--.i eql leql IEe^eJPue ;o :rcd z :slas eeJql olur paleSarSessluetrrele s.lErrjrueaql uo uorlc€]luoc JElnrsnrrrJo unseJ -z: :.r ieldsrp (SZ 8tg eql (VZ t 3lS) ad,fi srql;o sa8elqruessy e se ro asdelloc Surrnp par:ncco l€ql sluauela l ^\oqs v :iAI l, la s88rrg) sleu ;o uorlelor ,{ue [q pagrpour '8urue11eg;o eueld NUSIJVd CUVCNVIS gHI Z'n eql 01 e^Il?ler snleredde eql Jo uollEluauo :ql slJJUJr sluJUr:[a 1o uolgern8r;uor aq1 '.Jeaurl,pu€',JelncrpuedJed,',1e1pred, perurel -::: :UrluPluoPouos 'I: tL_. oU ueulrads 'se8elqruesse uo pes€q eJe sJeqlo aql se uoujurot - a:-\ IEnpe 'urellpd ,pJ€pu?ls, :t!.ql-z 3ll ti!: -: t I1etou ere pu? pazlpapr e:e (91 3q'7t61) lsoru eql eleu8rsep e^\ eJeq :Jeqrunu ur pelrunl '/rra uosuqloJ dq ua,rr8 sluarueSueueuoru e-re su-re11edpeleeder eseqa suorle-rn8guoc pauell?U prnleu se palardralur aq,{eru qcrq,r ' tnuaulllq snpoqlouDuo peseq sluetrrele ;o slueureSue:re luelsrsuoc 1t\oqs rleJ8elp cDerueqcs:snl?luddp uparlqlsuS,{lod snoraJ leql I€JaAes sl?e^er 'JeAe^\oq 'seSeyqruesse -ruoqrEJ E Jo uorlsoduoc lueuelerllnru aqJ-I t 8lc eueld Surppaq pa,uesardlla.tr Jo suo4relloc Jo uorlpulusxg ernleu peaue8rosp rreqt fq pezruSocer eq f,11erurouuec aseql lnq 'senprser culse? pa1e1r3rn8al Jo seraeJ se elzur8rro euros sJeA\oJrnqpuz s-rs8ua,recsJo sarlrluJ€ eql ,(q pue sessaco:d .{ecep ,(q palcagz eq ,{€ur sJeqlo snleredde puorsueurp-eeJql qd aql Jo 'qleep reue 'osdBIIoJ aql uror; Suqlnser slueuela Jo sf,erre puorsueurrp-oarq ,{1pr1 -uasse eJ€ se8eyqruesse,iue6 (9 3g 'ZSOf { 1 b aSprlpyy) pelerlsn r ere etep ot pasodord ueaq e,req 1eq1sueld plelels snorJEAeleqa\ 'elunlo^ srql Jo I reldeq3 ur peuqlno sr ernlcolrqJr€ snleredde uo seapr;o luaudoya,rap eql stueur -ele eqt Jo luerue8uel-re leuorsuerrJrp-aaJql pur8lro eql 1e-rdrelur o1 se8elqruesse aueld Surppaq aq1 asn ot llncrgrp pa,rord seq 1l 'pools -Jepun e,!\ sr ereue8 ueeceqleu8flod pre,res ;o uorlrsodruoc tueruelalt1nlrI eq1 q8noqlly 'stuerrrela cg ;o srred :no; '.(1qr -ssod 'ro slueuela rS pu" qS ;o qoea srred o,,.r1 lZ t'ras t ttJl sasnlBJsddBluopouoJ ueareqleuS,{1odsnoJaJluoqJeJ LCh.4 Sec.4.2l The standardpattern 65 !"'-qh .-: -" Sc r '':. i EE ::-:r::::ron of a Carboni- i.hematic diagram ft/':,:--!..L-._-.: a.:neatu' g:. =: )r Collinson etal. Ei=-=d and are not all Fig. 4.2-The standardpattern; A, beddingplane assemblagefrom Bailey Falls,La SalleCounry, lllinois, !l[::::eS- specimenno. X-6377, Universityof llhnois, x25; B, headregion of counterpartof the frrstspecimen of the conodontammal, from the Granton shrjmpbed, Edinburgh,specimen no. IGSE 13822,Britjsh Geological SurveyEdinburgh, x30. ID P\TTERN mals (Briggs et al.7983, Aldrrdgeet al. 7986; type within their 'simple rotation' category,to qF Fig. -1.2A)show tbe Fig. a.zB) display typical standard patterns, whichAvcin (1974) assigneda9 (48%) of his I ic:: :hree sets: a pair of and reveal that the ramiform group is at the ordered assemblagesfrom Illinois. Norby L .r -- rheir denticulated anterior. The pectiniform pairs are aligned (1976) recognizedeight specimensof this type Frr- a pair of Pb ele- with their long axesnear normal to the longi- and a further eight that were broadly similar I r-:: alrerior ends adja- tudinal axis of the animal. The differing pat- and could be ascribedto 'rotation' in his collec- qrr:: NusJJ,vd uv.IoclqNfldusd sHI t't .(OSd ,SZOr,iq;o51) pcael i\ aq f,eur ,uoqcerluoc eldurrs, se palerdralur .lr seldruuxeeruos 1eq1 alqrssod sr 11 puulsJapun e,t se lt--- urellzd lalpred aq1;o se:nlea;aql qJl€ru asar{lJo [E lou lnq ',uorlJE]luoJol anp sq feru, / ro sa?elqruasseslq Jo 0S teqr pareprsuoc(9161) dqroli ,{ro8alzcsrqt ol pa8uoleqse8elqruesse parepro srq Jo (%I€) zt l€qr punoJ (tr6l) ces [s't rtJl sasnlBJBddBluopouor ueaJBqleuE,{lodsnoJaJluoqJBJ 99 rDs{s [Ch.4 Sec.4.5l The linear pattern 67 r :: -:1,") of his ordered p: :-. rhis category. Norby IEi: :,r of his assemblages E-3----r:r'. bur not all of these or ---: larallel pattern as we F.! :-r rhat some examples E, :ontraction' may be f-: - -' r. SICTL{R PATTERN E:_:-:'r-nt is typified by the 5- .:: :ectiniform elements .r pairs, with --:--:dinal rS ::::isd. Their posterior F--:: rhe ramiform set, in r .r-: at a high "-igred angle *:--i- :it ro the axesof the E r-- fe$ other known r:.:t .:nd all show differ- B Fig. 4.6-The linearpattem:. A, linear l, beddingplane assemblagefrom the ltlsalcaram-Kieselschiefer, Hemer, westphalia.iedrawn after Schmidt1934, fig. 5b, x13; B, linear 2, beddingplane assemblage from Bailey Falls,La SalleCounty, Illinois, specimenDo. X-1480, University of Illinois, xl9 There are two distinct types of linear the assemblage,but, in contrast to linear 1, arrangement.The first (linear 1) is exemplified conventional orientation would place the Pa by a specimenillustrated by Schmidt (1934, elements at the antedor, followed by the Pb fig. 5a, b; pl. 6, fig. 1). All the elementsare elements. The ramiform group is somewhat aligned so that their posterior processespoint disrupted, but is commonly illustrated as lbl- in the same direction (Fig.4.64). If conven- lowingthe Pb pairwith the posteriorprocesses tional element odentations were used as an directedbackwards (e.g. Rhodes 1952,frg Z) indication of the orientation of the apparatus, Linear arrays of both types are rare, but the ramiform elements would be at the have attractedparticular attention becauseof rb ::i:m- bedding plane -- F- , i:lte Counry, Ilinois, anterior, followed by the Pb elements and, their bilateral symmetry.Many workers have - [ : . Srale Geological Sur- finally, the Pa elements.The second type of stated or implied that such patterns most linear array (linear 2) is that shown by the closely reflect the original arrangement,and specimenfigured by Du Bois (1943, p1.25, have used the resultant concept of the fig. 14, see also Fig.4.68) and subsequently apparatus architecture to develop functional II.\TTER\ re-illustratedseveral tim€s (e.g. Rhodes 1952, models (e.g. Schmidt 1934, Hitchings and pl.126, fig. 11, Sweet1985, fig. 1.,von Bitter Ramsay 1978, Nicoll 1985). Further, in their t:=; r-rs of elementsare and Merrill 1985,pl. 1, fig. 1, Aldridge 1987, categorizationof assemblagepatterns, Collin- !E :r.r'ii a line of bilateral fig. 1.6 this volume). The anterior-posterior son et al. (1972) considered all other con- tbc: :ie denticulated sur- orientation of the element pairs, as far as can figurationsto be displacementsfrom an origin- rs ::i lrpposed. be determined,is againconcordant throughout ally linear architecture. --': 'sluauale 'aue1d Suppeq eqt otuo eql p€eq :- .q, o, ,o,r"tu" '^",n Ienpr^rpur eql aurllapou Iururue Jo :-r: - :li :luaLUele ruro]lurxl roJ srseq e sE (9L6I ,{q:oy ees) sntoauryrq eql Jo asd€llol luenbesqns pu" uortJzrtuoc rel :---:-.i,r? eqI-/ t 8lJ snpoqlout Jo snlPr?ddB u^\ou{-JaDaq -nrsnrrJurelJorr-lsod ,{ue yo slraga aql 8u!€rod relqrurecsrp aql pesn e,\€q e^\ {peeyc 1ou sr -roJul'uonetuJuo ql"Jp teJnlBUe stuesarder Frrrru€ uolu?rc eql ur sluoII-IeleJo elrns IInJ eql urelled srqa '(gZ l 31.{) pruIUE qcee Jo {unrl sy ,{rleruurfs 1o aueld prlua,r-osrop ? sapnlJur Jql ol leuuou Jeau slueuelJ ruroltutt.r:d aqt Ieporx rno os 'l€culauufs flerelelq se,r pue enbllqo slueuele uroJruer aql qlr,t'urel snleredde luopouoJ eqi leql saSelquesseeuBld -1ed prepuels e ur 1nq'f,1reeu11pe8uerre 1ou Surppeq pue Surrmd lueruele ruoJJ aoueprle ere sueulcads eseql ur seSelqurasseeql esec eldrue sr e:eqa (S86I lto"lN '1461 uossddel aql f,Ir?ssaJau lou se^\ srql l?rll u^\oqs s€q osle aas) sluaruala JS aql ol :orralue ,{11q8qs q8rnqurpg ruo:; syeurue elelduroc 1o fre,rocstp pue Jplslno sluJrxJlJ ;41 aqt areld ot srat JqJ [Bu.ru"luopouoJ aql Jo slxe aql o1 1a11ered -sn1cruo-r; pue se8elqruessuaueyd Sulppeq ruo{ pau81p slueuele eqt Jo sexe Suol aql qtr,,rr ecuepr^a pesn eABq e,r ',(Felurrs lapoIII ]no '.lEJurlse,r :rnlJn[s eql leqt uorldunsse aql I ut Suruorlrsod sql pe^\olloJ ,{18uprome a,r.eq ueeq seq aJnpolrq3:p snlBredde lJnrlsuoJeJ I e14 se8elqruasseaueld Suppaq:eqlo ur uor1€l ol sldurallB snorle:d uo lure.4suoc roleu y -uarJo JelrrursB lE pezuSocer eq osle um qtrr{^\ 'I3nPr^rpur auo ueql erour 'luaruele eS eq1 ,{lqeqord sr srqJ'repureureJ Jo surerueJ eql asr:druoc IqAIur e8elqurass€ eql ueqt srxe eqt ot elSue -req8rq,{pq8qs e 1e srql 'azrs ur ulroJrun lou eJEsluauela eql ecuts pauogrsod 'peleuruucsrp eq uEr uortElnJrluep 'luql pasodord i(lelrleluel uele aI{ uone:3elut uela erour ,{pq8qs q1r,r ssecord rouelsod e las -sp rualroru-lSod Suunp pelecolsrp uaeq a^€q rrrJoJrrxsr srql urqll N Isruru" uoluPJc eql ur feru snleredde .srog nd ;o slred 1eq1 pe1se8 slueuala rrrJoJrrueraq1 ;o sassecord rouelsod -8ns pue ',pe1sv,r1pu" paulnl, ueeq eleq sa8elq eql Jo uoueluarro aAere E eql ruorJ palelncleJ -uressEl€iJ^es ur slu:r!JI: lEql pelou'Ja^: el8ue enbqqo u€ 1€ eJ€ slueurelo S pue W -/(oq '(IZI d 'ffOf) uossddel elqtledruocut aql seeJeq^\ 'snler"dde aql Jo srx? Suol aqi ol se,t (y1 3ry '92'ld't 6I) srog nq ,{q parn8r; asrelsupJl peuoqrsod e:e s.rtedlueruela qd pue unr;ed Treeurl Jo uaurcads eql leql pozru pd aql (SZ t Srg) leulue uotuerC tsrrJ aql -8ocar 1nq'(VS t'3r.1 '4961 e8prrpSTosp aes ur peareserd snleredde eql uo ,{prerurrd peseq :7961) lpruqcg [q adft 1 .reeu11;ose8elqlxess" sB^1lJpour inO sluJurelJ 1o 1u:uraceldsrp-ro uoJJ peJnpap slnlJJllqJie reaurl :ql pat:od uorlrerluoc urauour-1sod Eur4o,,rurlnoqtr,rr pue -dns srelsnlc pue se8elqruasse u,rou{ lsour Eurualleg .{q f,ldrurs a8elqurasse;o seuo8alec teqt pepnlruoc (1161) uossddel Z reeull pu€ uretrl eql y1e ureldxe o1 tduetie plnoc e.tr I Je:url sB ulaJ:q paquJsJp 'suJalted .Ieau!l qrqn qtr,!\ (f f ?lS) Iapour Ieuorsueurp-eerr{l o.{\l aql ur,{tHedsrp eql ,{q pesneJ ueeq s€q elo; ele1drue1e se urelted prepuels eql esn fllncrgrp JEInJTtJEd ,uorlrsod oJII Jreql uro{ ot ueeq seq snleredde ueeceqteuS,{1odaqt;o roEep reus slueuala pal?lsu€rl tsqt slJsJJe e:npelrqcle eql Surle,rerun o1 qceordde rng uorlrsodruocap-uorsseJduroJ (q ro spege 'snleredde aql ;o ,{:laruoe8 pur8uo aql peger uonJeiluoJ-uorlexBlJr eJJsnu,{q pauteldxa saop lI leql SutlsaSSns'sad,(1 .rofzur ragto eql eq seseclsour ur uec, sluaura8ue:re reeurl rrro4 IIe ol uorlrsuel s^\oqs osl" lr lnq 'sluerueAuerr€ sa.rngedepleql peleprsuoc 'alduexe rc1 (y7'd eueld Surppeq eql Jo uorurxoo tsotrr eql 'ZL6I) p p uosuITIoJ alnpnrts snleredde urelled prepuels eql sr fluo tou'peepul ernl leuorsuourrp-aorql 'elAurs e qlr^\ surelled e8elq -Jnrls l"uolsuJurrp-a:Jql:ues eql Jo Surualreu -ruesse eueld Surppaq go,{lau€A eql elrcuocer pue asdelloc f,q pecnpord aq qtoq u€c suJer€d ol llnJrJJrP 1r punoJ aAEq sJsIroA\ snol^eJd pJEpu€lS pue JeeuIJ l"ql A\or{s e^t lnq ^\olaq 'adfl slqt;o sa8eyqruesseacnpo.rd o1 stuerrrele SSSNIVUVddV NYSCYHIVNCI'IO ilqJl sasnlEJBddBluopouor uBaJeql8uSf,lodsnoJaJluoqJBJ lch.4 -Fr! 9-l :::roked 'rotation' of Ea ::\:nblages of this type, r -::: inear and standard tx :::,:-ced by collapseand re -:-::-drmensional struc- [-. : ::]e staldard pattern n :: :re bedding plane t ii: i'r!-r\1stransition to all EEl: :-_aiesringthat it does rFr -:::-\ of the apparatus. r--. : -ing the architecture E= ::liratus has been to EL-. -- :t a template for a m:':-. Fis. 1.7) with which : : : -,:lain all the main r,,=-: ::nplv by flattening lf :r:s:-10rtem contfactron i :::::ts. Our model -= was :l:f,ratus preServedin m_----': Fig. 1.28). The Pa r= :.r.itioned transverse --: E< =:::ratus. whereasthe t:-::: an oblique angle : :;.::-ie orientation of the i :: -:; ramiform elements 'I ![Ii -lhin this ramiform =:. = -: slightly more even E : :inared, positioned --: E:: :.r rhe axis than the r=:-\ the Sa element, =-"=::;- at a similar orien- ir- :..::e assernblages.We qf':;:: lhis positioning in i- rare used evidence -: .rs- -::lages and from clus- \1 : ::rents outside and =: -t elements (see also ;c. - .:-< ). There is ample lr:: ::inng and bedding i,i: -- : .onodont apparatus [::=,'a]- so our model r:-- : ane of symmetry. As x:--. :: ihe Granton animal =-_r:- lre have used the tr;:::-! of Gnathod.us j-6) Fig 4.7 The architecture of a polygnathacean apparatus, reconstructed using modelling clay. The r:". as a basis for ramiform elements are more widely spaced than in nature to emphasize thefu relative positioni. A. Iateral L---:-aments. view, anterior to the left; B, view ftom above the model. ln succeeding tustrations. rh; photographs have been retouched to remove the background and supporting pins. uuoJrrxeJ eql uorldnrsrp eql e8€lquresse uP uro4 srxP slr Suole peqdErSotoqd sI 1:ir al!rEJ-orelsod Jo lepoul n(I ur uaas sB'sred qg puD ed egl paleJrTder u-ra11ud eqJ F:-l :lers sroullll I08I-dz9 ou .srog Jo aql uaq,r sr Iell€red r: ;--s e I slleJ {elrPg ruorJeSPl depa,ro o1 sp€el uorl€luarJo srql ur SuruatjelC '(g l AlS) fpoualue (spre,tdn ro) spre^\u,lrop -V :i:!ed eqI 3rJ lellssd 6 t '(Zt t BtS) fpouelue spre,,,rdnpa111 sr peeq puB Jprs euo ol pJtlll snlsJ€dde eql qll\\ eql ueq^\ sllnser (sluaruela eql Jo uorlelueuo 3urue11egenbqqo,{q peureldxa oq uet uJell€d pesraleJ e seJnpoJdles uloJruer eql;o Suruorl ,prepuets lesJJo, eql 'relnolu€d u1 Sutppaq -rsod roualue ue aroqm) luaute8uerle .Z raautl, eql ol a13ueue le Peeq egl Jo 3uI1[I13 Suldldu]I oIII '(t t f 8tS) fl.rorrelue sprc,tlu.rop ':elncrpuadred aql uror] sprzl({Jeq ro spJ€,{ dlrq8lts parlu p€aq aql 'SuruolleU IErluel -ro; 'spre.tdn 1as;1o 'se18ue anbqqo f11q311s > -osrop ,{q pecnpord f,ppear sr ruorl Suruepeg ,{q pecnpord e:e urelled eq1 uo 'suorlelu3uo I€UOrlUaAuOJpeJJoc Jreql ur slue[xele eql qlr,|A suoq€rrp1 ed,q t"qt ;o sa8elqruessesatectJdar F 'lueure8uBJIe .l Jeeurl, ar{I su-re11edreaurl ,(11ern1eu',{r1aruru,{s ;o eueyd aqt ot relnf,Ip elqrledruocur ,{lSurruaes o,tr1 aql ureldxa o1 -uedred dlreeu '8urua11eg l"lel?l lieJlp 'uret ,{1rpqzs1r ur sel lapou rno Jo lsel IennJJ V -1ed prepuels aql uo pes"q sr Iepou rno sV 'ereJ eq ol u,,!\ou{ 'pslecqdar eq plnoqs stuerrrele aql eJe soS"lquress€ :eynorpuedred tnq 'suorlel ;o suorlrsod e,rrleyereqt 1nq 'sa3€lquresseeueld -uarro elqrssod yo a8uzr pelrurq e urqlr,{ ,{luo Surppeq ur luaredde aq 1ou 11r,rsa8eur ctqder8 Jnrco plnor!\ slql (0I t Br4) s.{emaprse1ddo1 -o1oqd eql ur ueas SurualroqseroJ eql snql ol les uJoJrrrrEreql SursnEJ'eprs euo o1 f11q311s 'eue1dSurppaq eqt ot 1e11eredsaceyns:eueyd pu" 'sIx€snl"J"dde aq1o1 1e11ered elelor sluau laqt Surrq ol {ressaceu el8u" tunrururtu eql -a1e u-ro;rurpad eql leql os spl€^\roJ pallq q8nolql aletor ol puel plno^\ stuauala oql leql pEaq eql qlr^\ pe^erqcB aq plnoJ srql slualx ur.{1uo sragrp 8urue1legpue asdeloc;o ssec -ele uuo]rrrJeJ eql [xo{ uorlceJrP lualaJJrp -o:d lernleu eqJ suorleluerJo 1ua:a;;rp ut 8ur E uI 1pJ ol stuetuele uroJrurlced eqt sesnec -uauBU Jo stlnsar eql Suqcrurlu-qderSogoqd tEr{l snl?J?dde eql Jo esdelloJ e,{q peureydxe eq1-eue1d leuorsuaurp-o^\l e oluo eJntJnJls eq {eru tnq 'elecqder ot tFcrJJrp erou leuorsuerurp-eerqt aql laalord ol pesn sEA\ sr u:e11ed ;elncrpuedled eqa seSeyqruasseeqt se13uesnoue,r uorJ Iepou aqt go ,{qder8o1oq6 ur se pepa\oJc auro3eq f,aql tnq 'llnseJ stueluele '(1961) sue11p16 pue s88trg ,{q peqursep eql ;o 1uaruu311eue seop ,(1uo 1ou :(6 p 3rg) qceordde eq1 paf,oldua eA\ ernlonrts srql uo uoqrsod (leret?l-oralsod ro) le]alel-oretue 3urue11eg;o sloaJJeeql euru"xa ol Japro uI Iapoulo enbrlqo'g :SIx '{o^rns IerrSoloeCetels sroullll'IZZ-d/9 ou ueturceds'srourlll ^\er^ '{tunoJ ellss E'I'slleC ,{alrpgruorJ a8plqtuessB auEld Burppeq 'V :urat}edp:epuets tasJlo aql-8 t BrJ g i.t [L t 'tes ilqcl sesnleredde luopouoJ ueaaeqluu8,{1odsnoJaJJuoqJEC OL lE€s [ch.4 Sec.4.7l The function of the polygnathaceanapparatus 7l group in this assemblagesuggests that these elementsmay have been oriented almost nor- mal to the bedding plane. A reducedangle of upward tilting would result in a linear arrangementin which the pectiniformelements are oriented in the oppositedirection from the ramiform elements. An assemblage that approachesthis arrangementwas illustrated by Schmidt(1934, fig.3), but a componentof lateral tilting has produced a pattern that is transitionalbetween linear and perpendicular. Flatteningof our model thus accountsfor all B the repeatedarrangements shown by bedding plane assemblages.The standard pattern, in ta :: - -. Sa e County, particular, is shown to be produced by lateral Itir :; .ri model - compaction of a bilaterally symmetrical apparatus,without any 'rotation' ol 'contrac- :'-<::ro-lareral) position tion'. It follows that the head, at least, of the f .l:€: an alienment of the first Granton animal is presewed in lateral lE:i :acome crowded aSin aspect,and the apparentbilateral symmetryof E ndicular pattern is the soft parts in that specimenis not indicative | ::::.-3te--::e but may be of dorso-ventralcompaction (contrq Bengtson r--+ :: the apparatusthat 1983).lt isprobable that the entire specimen is hr: :lements to fall in a in lateral compaction,confirming suggestions ::= :he ramiform ele- that the structuresof the trunk are comparable k .:::ered rvith the head with thoseof chordates(see also Aldridge and c;: :re pectiniform ele- Briggs 1986, Aldridge etal. L986). The pre- d:: : '3 apparatusaxis, and dominanceof standardpatterns among known e-. :: rhe ramiform set to bedding plane assemblagesindicates that, on E - - r. f115 would occur death, polygnathacean conodont animals E r.:ii of possible orien- usually came to rest on the sedimentlying on r:-:,:: assemblagesare their sides.This implies that the animalswere laterally flattened in life. q-- :--,.iel lies in its ability o =::]]lnglv incompatible h -::eer l arrangement, 4.1 TIJE FUNCTION OF THE I E-:: --!rrrectconventional POLYGNATHACEAN APPARATUS E. ' :roduced by dorso- '.:. :r-ad tilted slightly The architecture of polygnathaceanappar- mr; (Fis.4.11). The atusesdeduced herein and repr€sentedby our :r ;:are an antenor posi- model is radically different from most recon- r= producesa reversed structionsproposed hitherto. Previousauthors *:-,',s-: ) results when the B have envisagedthe elementsas longitudinally rcs =::enorly (Fig. a.l2). Fig. 4.9-The parallel pattem: A, beddingplane assemb- aligned,often in a linear series(see Aldridge rEr-:: lrn leads to overlap lagefrom Bailey Falls,La SalleCounty, lllmois, specimen 1987, fig.1.5 this volume).The evidenceof E!- :: seen in Du Bois' no 57P-1801, lllinois State GeologicalSurey, x25; B, beddingplane assemblages,on the other hand, posterolateralview of model, ir::: ,rn of the ramiform indicatesthat the pectiniform elementswere IF1,a!\) aPnlul? srqt ur 8uI 'rolretue eqt ot ,{!q8!ls pup e^oqe tuo{ pe^ler^ Iepou :urelrpd I rpaul eqJ-II t BrC 4 :1eq lou PFoc pue 'uolll I l-r a:e sluaruala IrJroJIureJ Fl.-nJrtuep pue sdsnc eql hrnr srred eql Jo seJeyns o:cr aiesr,rua o1 elqtssod st rL->1 aql ,{q pessaco:d se,rr = srueurele urJoJrrrrcraql -nc,.ung pale:8e1ul alqrpeJc o: q9961)'p 7as??rtg rr-.rinl Jql roJ luaura8ue-ue r :qr arE l"qa\ 'rouelus r!-::ruEr aqt qtl, N (ng d Eli JrrrruE eql Jo sIxE eql ot Ib--:rurEr eql pu? esJe^suPJl @:: :?.urtaql-zI f 8Ic 'lepour rouelsodenblqo'g :€ZX 'IIol,I-dZg ou uaurcods :urelled lelncpuedled eqJ-ol 3lg Jo ^ler^ 'V t E {.r ll t 'ces ilqJl sesnleredde fuopouoJ ueareqleuS,{pd snoJaJluoqreC ZL lch.4 Sec.4.71 Th€ [unction of the polygnalhac€an apparatus 73 .$.1 L :r: :-: :'o:terior view of B Frg. 4.12-The hnear 2 pattern: A, specimen no X-1480, x25; B, model vjewed from above and somewhat to the postenor, transverseand the ramiform elementsoblique grasp they must have adopted a different posi- to the axisof the animal (seealso Norby 1976, tion from that in which they are found in p.54). With the ramiform elementsat the bedding plane assemblages and clusters. This anterior, what are the implications of this idea has been forwarded several times; for arrangementfor the function of the apparatus? example, Jeppsson(1971, p. 107) stated that Bnggset al. (1983) consideredthat the most 'it may be difficult to define one exact position credible integratedfunctional systeminvolved of each element, but it will perhaps appear that the ramiform elementsin graspingprey which they had different resting-growing positions was processedby the pectiniform elements.It and working positions'. To bring the cusps of is possibleto envisagethe opposeddenticulate the ramiform elements in our rnodel into surfacesof the pairs cutting and grinding, but opposition would require a rotation of each the cusps and denticulated processesof the side of this part of the apparatus inwards ramiform elementsare not preservedin oppos- through about 90'. This action could have ition, and could not have functioned in grasp- been accompanied by extrusion of the cusps h.r -: _-:e anterior. ing in this attitude(Nicoll 1985).In order to from the enclosing epithelium in the manner paqsuqndun urseg sroul[I eqt uoq seBelquess€ ueeJer{lEu8flod JoJ Iapou puoq.un} E pesod uerseurohtr tuopouoc se(I tL6I f w 'urc^v -ord pu€ aJnpelrqJr" uE peqsrJqetsea^eg elA ' 16Z-6LZ61 ltDtlrd-lpue':oJs, 'q8rnqu rp3 JOSnOreJrUOqrEC er{l ruorJ ecuepr^e ./g6I .d .trl ^\eu-sluop.qrnus HCUVSSSU -ouoc Jo senrurJJEeqJ pue '>I N A'uos{relJ'g A C's33ug't 11'e8puply UUHIUTL{ IIOJ SSIJI'IItrISSOd 8'' 'sser4,{lrsre,rrunpJolx},Dxot l\sso! tDualqota .I .v :(spe) rlcelrN H trpue ueuJJoH :q .$uop ,s33ug .1 .g .e8prrply -ouoJ 9g61 9 g q pue 't€-II'xessns'retseqrrqS 'i(J1e:1ua,rpalJeJrp eq ot readde 'poo^uoH stlla 'suopouoJ tSolotqoabpd plno^\ stuauele ed eql Jo sruroJlsld eql puB .u Io :{ pe, Jipllplv f :ul sluerrJeleuJoJrrrrsJ aql sdsnJ er (6'1 8rc) ^lJt^Jl leJuolsrq Jo e :fSolorqoeelpdluopouoC ,g6I f d 'a8puplv purue aqt Jo peaq aql ol ur8reru srql acerl a,r saJNsus.{f,u Jr pue 'IunJl eql Jo Surlsl^u rof?(x Jo eJuapr^a ou sr eJarll IESrop eql lueselder ,{eru ur8teru slql (9't s8g '€86I 'pp sBBttg) uaurceds lsJrJ aql uo luapr^e aJ€ 'luaurdole^ep JaleeJS eql qlrx\ sdeqred euo esoql ,(1ug '1J[.4ASV rW pu? seuof 13qt 'eprs ;o tu=:;iod sercedsluopouoc 'C rI{ papr^ordse.r ecuelsrssecrqderSotor{d '(t 3U 'SgOt 'p p e?p.upl17)Iun:l eqr Jo ,{rr EE:ierrlnl^l s86I s d'lloctN tuosurllrl^d^q -Ixa4x3 rorJalsod aql Jo aprs Jsqlra uo sfPJ urJ r-:..:t" otuoaolo| D octSoloag pue I sJW [q pelJ"Jp eJel{ sSur rt rrsnteredde (:ay?er7) -^\eJpaur-J 'suorlseJloJ ,{qloN pu€ ,urJA\/,srog Jo lueudola^ep leorrlarurudse ue a{oqs aseql 'slzrxrue uolu?rg eql {q pephord sr uorlpf, 4]c-:OaL6lAIe^o{qsPI J nO eql ol sseJreroJ',{a,trng pcr8o1oeg ale1g -:: pepeJrp ere^\ I - I0I t D1Dl,fla.Is\\ttol slourllJ'lua) '-I {I pue 'suo4Jallos38"[quassB -rpur uB lnq Jo f,llPsJop pi::urp.;o sesnlereddeluop sluauela urroJnueJ"{lJeJlue^ eql sdsnc aql Jeqleqlr sepoqu pu? llocs aql uro{ prrel?ur }o uEoI Jo rf, ::i:rrr.lg 1161 1 'uossddel ,d8ol ot se ecuapha pco^rnbeun ou $ eleqJ ' t8o eqt Sur8u?Jr?JoJ'srourlll Jo,$rsla^run 611-1gy tr7 .e{elg .g .(I 'rrrJopeld rolJalsod pue epplq ae{ L-.:"i.olod'{qd BoaBoaoPd -oec Jo luaur edeQ rc {ueql .J.U.!I.N JorJalue -r>-- e :se8elquesse luop elA '90IS/E/UC tuer0 qcr?aseu ,{q olur slueureJePd Jo uorlBrlueJeJJrpeql y -i"!EEl{ pue ureldxe plnoc srq; uoqrod ror:e1ue eql purqaq H A's8ulq.trH papun; s rffiolorqoeepdluopouor uo IJo,r rno 'sz td ,{e1 uorpod rouelsod eql leqt os paullJur sluopouoc lo /Dlllrol^ SINSI IS9CS'IIITONXJV erour ,(pq8qs uaeq e^€q ,{eur lueruele ruroyurl E:f:\f-.+-:d 'stog nc Et6I d a -Jed qc€e',(1errqeure11y'sp-ren1ceq,tylq311s €: -J iO raBeuo]slelrsre^Iun pelJarrp eq ot uJoJt€ld ro ap€lq rorratsod I ,F.j :( spe) e8ueC U Pue E- Jo ql.el duJoqeql eql SursneJ'srxE fpoq eql ol lprxJou ueeq e^eq ":.lr-il{ 'sluarrrola a i:"-Lrlql €96I V f'uos^{Pc tuereJJrpdlSurtuts epnlcurqJrq^\ ,(eru slrun eql Jo uoqJod rouelue .(1euorlua,r a;: sauas _a ol looqapnc Jo fu€ul 'sesnlerudde ruro;ruoc-uouJaqlo Jo -uoc eql Ieuueqr pooJ eql ol relnJrpuafuad t ,'i?r.s sloulllJ sroullll uorl?Jedopu? eJnFalrqJrpaql Surpnrlsuocar ueaq a^"q lou plnoJ luaurale arqua aql l"ql os {:\ e'I uorJ saBelquresse uI alzlduel ? sp alJes u€c u?ld uear?ql€u eql arnl? rnJ erxos xroqs E:d L/61 ) C 'lllllehl 'apelq Jo lnq 'I?ueld [ -I 'I|urr^v J 'uosullloc -B,{lod eqt luexa leq^r ol ees ol Suqse-re1urf,1arerere edfl srql Jo Euaruele 'Jole,rroH uor1 'n9rL9r eq 1,t\tl puz 'relJo^\ luopouoJ aqt ot uedo -cunJ ol uoqeluerro purpnlrSuoy e olur pel?loJ uoll€rolsel | ::uauEg Jo ,r!ou $ ,{SoloqdroruIBuoqJunJ Jo plerJ eJaA{slueurele ru.royu4cad aql leql asodotd ol E{lr\\ pu€ 9 g 6's33ug .NI_I eql^ln4 'sdqsuonElatpuz sar8oloqdroruIelueluele uosearou eos eAA sluauJla rxJollurlf,Jdarp ol qlra sasnlsr?dd€Sururtupxe fq pe1se1 poo; ! rEoFlosor aqJ €86I I )I luarellp lceq ssed o1 ,rno1po1 uorlrsod purpnlfuol t1-:I 'uoslrelJ ' C g O's33ug puP pepuexa eq osl€ upJ laporrr3r{I selssnur otur slueruele aql Jo sexp 3uo1 aq1 tq8norq " E gI DrDqDT snlPrPdde luop luoreJJrp[q pete]ado uoeq e^eq ,(?ru foql osp lueruelorrr eqt ,{le)H sr rI (586I ueple1 rFroLirDunJ V t86I S'uols8uag l€ql SuqseA8ns eqt uro{ eJuap '€96I uos^req) suorlrsod Surlser pue puorl '902-s8I 'slueuale S -uedepur u? (eldsrp ,{luourutoJ slueuale -cunJ ol"rodJocu pu€ slueulele pered uorle + PoP elnpn4s luopouoc tr41 ;o eqt uI paulJereq ol uleuler -redo rp t,uE stuopouo. ueuqu9c 'J€lnJrlled 'sluetu FJelelrq eql e^Io^ul qloq l€ql ur 'sprour rrYlas aqJ. 916I S 'UoslSueg -ele rxJoJrrrreJeql Jo suoqrsod e^rlsleJ 3ql -x,{uI eql Jo snleJedde IenSull aql Jo leql ol -:1d 5-''dd 791 'u8ledtueqJ ,{flerJedse'lepotu eql Jo slp]e(I ecuepr^ealqe snoSopue ,{peo-rq sr uelsfs srql Jo uonerado irsre^run I lL.r 's!sa4t Oqd lrp^" rllrlr lualsrsuoJ'sesnlereddeluopouoc Jo epour eql (9461) uoslSuag fq pelelnlsod ilqJl sasnlBrBddB luopouor uBerBrlfBuS,(lod snoJeJluoqrBJ VL frEs6 [ch.4 References 15 Ea-:- consistent with avail- PhD thesis, University of Illinois at Urbana- 1940 alndOzorkodina brevis (Bischofl and Ziegler n:. erfthe model, especially Champaign,152 pp., 5 pls. 1957) from the Upper Devonian of the Canning of the ramiform ele- Bengtson,S. 1976. Tl'e skucture of some Middle Basin, Western Austmlia. BMR Journal of conodonts,and the early evolution of Australian Geologyand Geophysics9 133-147. --"-ilr =ired. In particular, the Cambrian conodont structure and function. Lethaia 9 Norby, R. D. 1976. Conodont apparatuseslrom n:-r display an indepen- 18s-206. Chesterian(Mississippian) strata of Montana and 5 . ::nents, suggesting that Bengston,S. 1983.A functionalmodel for the cono- lllinors. UnpublishedPhD lhesis, University of krr operated by different dont apparatus.Lethaia 16 38. Illinoisat Urbana-Champaign,303 pp.,21 pts. H :-: also be extended and Briggs,D. E. G., Clarkson,E. N. K., and Aldridge, Norby, R. D. 1979. Elemental architecture of E 1::.:.ratuses with different R. J. 1983. The conodont a maL Lethaia 16 natural platform conodont apparatusesof Missis- l-14. sippian and Pennsylvanianage. Ninth Inrcrna- |or,e-- and relationships. The Briggs,D. E. c. and Williams,S. H. 1981.The tional Congressof Carboniferousst'atigraphy and | :r-hology is now truly restoration of flattened fossils. Lethqie 14 geology, Urbona-Champaign,1979, Abstructsof l'':: e 31[s1, and it will be 15'7-164. papers,249. : L- a tat extent the polyg- Collinson,C., Avcin, M. J., Norby, R. D., and Rhodes,F. H. T. 1952. A classificationof Pennsyl- vanian lournal Paleon- rr e as a template in Merrill, G. K. 1972. Pennsylvanianconodont conodontassemblages. of assemblagesfrom La Salle County, northern tology26 886-901,pls.126 129. : r--::::ecture-.1 and operation Illinois. Illinois State Geological Survey Robison,R. A. (ed.):1981. Treanse on Invertebrate t[ -: :Pparatuses, many of GuidebookSeries l0 37 pp. Paleontology,Paft W, Supplement2, Conodonta, tn: -. liJferent elements. Dawson,J. A. 1963. The oral cavity, the 'jaws' and GeologicalSociety of America and University of the horny teeth of Mlxine glutinosa.lni A. Brodal KansasPress, Lawrence, Kansas, 202 pp. and R. Fange (eds.): The biology of Myxi\e, Schmidt,H. 1934. Conodonten-Fundein urspding- Universitetsforlaget,Oslo, 231-255. lichen Zusammenhar'g.Peliiontologische Zeit- Du Bois, E. P. 1943.Evidence on the nature of schrift16 76-85, p|.6. DTE\IS conodonts.Journal of Paleontology17 155-159, Schmidt,H. and Miiller, K. J. 1964. Weitere Funde pI.25. von Conodonten-Gruppenaus dem oberen Kar- :Jaeobiology is funded Hitchings,V. H. and Ramsay,A. T. S. 1978.Cono- bon des Sauerlandes.Paldontologische Zeitschrijl -=:z:: Grant GR/3/5105. We dont assemblages:a new functional model. 38 105-135. ll.ii:- Department of Geo- Palaeogeography,Palaeoclimatology, Palaeoecol- Sweet, W. C. 1985. Conodonts: those fascinating little Journal Paleontology 59 f f :..i.. for arranging the ogy 24 t37-149. whatzits. of Jeppsson,L. 1971. Element arrangementin cono- 485-494. It'- ::e Scott and Rhodes dont apparatusesoflJindeodella type and it srmllar Von Bitter. P. H. and Merrill. G. K. 1985.llin- ra*- .rd Dr L. Kent, Illinois torms. Lethaia 4l0l 123. deodus, Diplognathodus and Ellisonia revi- k :. - ior accessto the Du Mashkova,'1. V. 1972. Ozarkodina steinhornensis sited-an identity crisis in Permian conodonts. ic: . :,-rllections.Line draw- (Ziegler) apparatus,its conodontsand biozone. Geologicaet Palaeontologica19 81 96, 1 pl. Yalden,D. W. 1985.Feeding mechanisms as evi- b'- \L.r J. Wilkinson, and Geologicaet PalaeontologicaSBI 81-90, 2 pls. Nicoll, R. S. 1985. Multielementcomposrtion of the dence for cyclostome monophyly. Zoological EE!-. ;:s proyided by Mr D. conodontspecies Polygnathus rylus rylus Staulfer Iournal of the Linnean Societv84 291-300. ta= palaeobiology: a - --:::,iont a R J. Aldridge (ed.): L . :tw\. Ellis Horwood, _ : - _:r. B:--=.-..-D E. G. 1986. Cono- hi: .rc U H. Nitecki (eds): Orrbrd Unive$ity press. s--.. '-- E G . Clarkson, E. N. K., 1:]- The affinities of cono- E ::rn the Carboniferousof t -.:jt,iia 19 279-291,. ! J:: \foinesian conodont fE - rnojs Basin. Unpublished Form and function of the Pa elementin the conodont animal R. S. Nicoll ABSTRACT 5.1 INTRODUCTION Blade and platform type Pa elements,both as The relationship between conodont element fused pairs and in pairs reconstructedfrom morphology and function is an area of study discreteelements. have been examinedin their that has been given relatively little emphasis presumedfunctional position with oral surfaces compared with taxonomic, biostratigraphical, intermeshed.This has provided clues to the or ecologicalaspects. There hasbeen a general function performed by these elementsin the tendency among those who study conodont conodont animal. Based mainly on Upper elementsto regard them as objectsto be clas- Devonian or Lower Carboniferousconodont sified,and not to speculateabout their possible species,four major categoriesof morphologi- function. In addition, most papers that have cal relationship betweenPa element pairs are discussedthe possible affinities of the cono- recognized and defined: blades, plates, hol- dont elements or animal have not carefully lows, and crests. Cone and bar type Pa ele- related element rnorphology to functional ments have not been studied. hypotheses. Pa elementsof the typesstudied are believed There have been someexceptions, including to have performed a crushingfunction on fine the studiesof Jeppsson(L97l, 1-979,1980), food particlespassing through the mouth of the Lindstr 17 s J") s qcl Ielxlue luopouor aql q luauolo Bd aql Jo uollJunJ puB rrrJod 8L h a-nimal [ch. 5 The Pa element 79 SQLE\CE types is seven. Some apparatuses may contain the one on which most conodont biostrati- m= ..; elements in the cono- lewer element types, but it is doubtful that graphy has been based. Morphologically the Pa t =:::er of dispute until the apparatuses will be found with more than element is usually more complex than the other unless there is a differentiation of the apparatus, and is the most F--en from the Granton seven, elements of the fu:::sh. Scotland (Briggs two pairs of Sc elements. Thus many mid- differentiated. Frequently the Pa element has a--:::es had been generally Palaeozoic conodont apparatuses, exclusive of developed lateral processes and/or expanded I --:; Pa elements anterior the Icriodontidae, consist of 15 elements rep- laterally to form a platform. rfr. ::-S) and the other with resenting seven element types as follows: M The Pb element is structurally linked with k::--s anterior (Jeppsson (2), Sa (1), Sc (a), Sb (2), Sd (2), Pb (2), and the Pa element. It sharesa similar transverse .:-\-ribed by Blggs et al. Pa (2). orientation (Nicoll, 1985) and appears to have b: ::: discernens elements The earliest conodonts, proto-, para-, oreu-! been located slightly anterior to the Pa ele- rt:-..: end of the apparatus have simple undifferentiated cone mor- ments, but may have been partly overlapping. : :rents at the posterior phologies. By the late Cambrian there was a The function of the Pb element is not the same - differentiation of element types within the as that of the Pa element, and it has not lC-r :ugsested that the apparatus structure, and by the Early Ordovi- developed as great a range of morphologies. It tF--=-n from the Granton cian it is possible to distinguish as many as is thus relatively conseruative. distinctive element types in each preferential morphologi- i :--. -:nctional position but seven The reason for the t l:serer. the positionsof apparatus.This number appearsto hold for the cal malleability of the Pa element over the rest i ::s::nen are very similar rest of the time rangeof the Conodontaas the of the elements in the apparatus structure is i =.:r oi the fused clusters maximum found in any one animal. not totally understood. I believe that it is prob- d :irm Indiana (Pollock The differentiation of element morphology ably a response to variation in the prey that the -r krr-:: 1987). This suggests within each individual animal probably repre- animal was feeding upon. There may be an { =: ;lernents in the Gran- sentsthe initiation of specializationof element analogy between conodont elements and teeth t*<::r a normal distortion of function within the apparatusstructure. Some in those vertebrates that chew or grind their I =:i ha\e resulted from element types appearto be very conservative; food, where the rear teeth are usually the most n .: :.rrm factorsreldted to for example, it is possibleto recognisean M complex because they are assigned the task of r cr: ---: rissues of the organ- elementbecause there is a consistencyabout its breaking down the lumps that are passed back morphologyregardless of the speciesto which by the gathering and cutting front teeth. Rear lhi: :iusters and the Gran- it is assigned.This conservatismof morpholog- teeth reflect closely the type of food eaten by good the function to { 1y!r ) suggestedthat the ical type is a indication that the animal, and are modified from species f = ::r3 apparatus was M of the M element did not changesignificantly species as eating habits vary. after the basic form evolved in the Earlv There is a broad evolulionary progressionin c-r: - tbllowed by a com- [:r: Pb element pair, and Ordovician. the morphological change of the Pa element. It n. ::ir (Fig.5.1). This is started in the Cambrian where, if it can be E r{ :1e sequence proposed differentiated, the element had a coniform It- =: has been confirmed morphology. In the Ordovician the denticu- lated bar was introduced. and this was soon r c': -:idirional cluster mater- 5.4 THE Pa ELEMENT Ecr:1987). followed by the blade. The platform was the If the M element representsa conservative last general type to be introduced as the elementform, then the Pa elementis the radi- denticulated bars and blades developed lateral cal element. No other element types evolved expansions. By the late Devonian or early Car- rr-PES anywherenear the number of morphological boniferous the coniform and denticulated bar stylesthat can be found in this group. Pa element morphologies w€re rare, and the r =:Jual expansion of the The Pa element is usually consideredto be blade and platforrn morphologies were domin- G =iluded within the cono- the most important element in the conodont ant till the late Triassic. The discussion that d :: i: now generally recog- apparatus.It appearsto be the element that follows deals only with the blades and plat- m--n number of element evolvedmost rapidly, and hasthus come to be forms. r sluauele I o^u eql 'sl"rrJrur?luopouot ;o sdnor8 tueragrp -uo4eler Suryo,tr luauralo-Jalur xaldruoc e:oru r s---:j ur alqrssodsl ll '(€86I ,{q pessecord are,t saprlred poo; fe,n eq1 ut qcnur e^eq sluerrJale e; edfl-ru:o11e1d eq1 rE:J aql uorJ peuElqo uorl€uel ouros alecrpur o1 q8noua luaraJyp eJB '1uerue1aSursoddo d ::rsnlc snoJerrJnu eql Jo ,{eql lnq 'leer8 lou ere sed,4taerql eseql uee.tltl aql qlr^\ lceluoJ ur lou sBA\ tuaurala qcee Jo perrl q1,.ror8 6 _ - .r1!\) e se -eq,{Soloqdrour luaurale ur se8ueqc aqa ec?Jrns prEq eql snql pue 'pera^oJ-anssrl ,{[qe r lo a\oJ esla^Su?Jl P 'slurod-pue eqt Jo IIe uee,\\leq -qord lsoru eJe,r sluauela aql leql requleuaJ x -i-ul::: rr ' iSoloqd:orrJ eJellns tsrxa sruJol leuoqepur8ralur pue 'sat:o8elec o1 luegodurr sr lI 'leurue luopouoJ aql tr : \€r{ aepquopolrcl aI{J peurJep ,{pre.t1rqre ate aseql luelxole Jo ,{erd elqeqoJd eql are.{ lBql uo11ue1daqt 1o - Sursoddo aq1 uo:; ,{eare3ur,rlnc luaruele qcea slsel Jo sll€a IIer Jelno eq1 Surqsn:r snql 'sat?J sur8reru eql pu? uI slueul eql uee^\teq zztg snsuodra snpotlcl (q) ;o 1€Jal€l lJeluo, tualrrela tueqt Suqyor .{q salcrpud -ele eql Jo eullpltu aql 3uo1e ea:e ?!\orJzu ro pooJ eql peqsBu a^Bq plnoc uorlJs srql slusur leurs e ,{yuo qlr,r\ ruJoJ aql ;o alrsoddo -ele eql Jo uorlrs a^rlJaJJa 3ql se1( uortorrJ ^\o1oq r- '. --::Ip a.re,nereua8 aldls Jqt sr uJroJ tsJrJ Jqf lleluor reeu ut sutS SulqsnJt aqt leql raprsuoJ I 1reluoc reau aql qlr^\ slueuele eql eullp[u ur sJJe] Jeuur JLll qtu'l uorloru Surqsn:r q--J ol ueunls eql Jo 'lsour -Jetu IeJal"l Jo lErJlBl r[ r:lrls aql oslg se , Dutpol eq1 3uole eere uedo ue s"q txroJ 1'to11oqeqJ ? pEr{ e^Eq Jo 'uorl3? ?urllnc e qlr,n raqtaSol '-reln8e-r.rr FN-J aql ol uslf,IAopro flq8rq ro qloorus reqlre eq ueJ soceJ pue lrede pa^ou oleq plnor slueuele eql puP 'lJ?luof, BeJ€ rs-\:,rtns \-rsuorlnlo^e lJeJrP -lns luelxele eqt eceJrns Jo 'palJoAe lou ara^\ pu? 'prurue aq1 urqlrariaceld e8rel e seq uro; a1e1deql'(2 8rC) eql ul peuotlJunJ slueulJlJ :ql Surutnssy F.::q paurErxeJ 1l esneteq 9 tserc leqt eq '(eZ =::l! \Je \ E uAeq e Eq lSntII pue '.{,'olloq eql 'alEld eqt :paauSoce: uec S 3lS) Surddepe,ro sareJ l"relel reuur peo;q -:C.",7 DulpoltDzo eql seee;rns :eddn u.ropeld ;o sarro8elzc Jreql qlra\ s3^\ slueurale eseql Jo uorleluerro alrl :E(i:: pue tueruela Sutsoddo aa:q1 xaldruoc e:oru qcnu sr dtqsuotleya.t alqeqord lsour eql l"ql .{roqs (1361 peorxag r ::: \ lErl] asoql ueel$3q sl eqt lrBtap ur 1nq 'ruro;1e1d eqt Jo e3eJ aql puE IIo3rN '6961 )1co11o4)stuewele Ed Durpul F :-1 _IOSaSEJJnS Jalno puE serrroJaqaJeJJns Surlro^\ eql leraua8 u1 sdrqs -tozg e:d.f1-epe1querJnlrs Jo srotsnlc pesnC 3--::jaJJrp ,{ue aq o1 rzedde r:.::::r srql go de1-rer'oereg p::;.ir aqt ,t\oqs qloq (dsnc snpoqlDu[) - ruroJ lsarr '!:snqtouS[lod - LuroJ 'e :sn4uu8tlod rrlro] el€ld 'p :fidaptDrultq wto! eteld ^\ollor] -Lr :aGoloqdloru lueJaglp '.:snpourl tJj.lol eluld'q:Dutpol)D2o - ruro} apelq 'E :s,,,ro11o;se e:au:8 yo slueuelo ed uo pespq ere lnq -: .ig I S 14) u,,ttoqsare cr!BrueqaseJe suortJes-ssorJeql Jo sedeqs u^\oqs 1ou sr uAo:o aql Suue^oc enssrJ enssrt ,{poq q peppequa N S:[dUEXeO^\J seJPJJnS (seurl) anssl e^[.euuor pu? (lc€lq prtos) etpld l?sBq eq] qlLh Suruedo qlnour aqt ur u^\orr luerxele eql s^{oqs eldluexe qreg txet aqt uI paqFcsep lueruale ed Jo seuo8eteJ rofEtU eql Jo suopres-ssora)-z s Brc li, :rrq$ sluauele Suldd?l lE-::i?l eql ueeq e^eq plno.tr F rauorpunJ slueuelo aql p.-urioddo eqt qlr^\ qseul ol u::t lEr{l suEerllsluaurale Ed br,,.qd-rour apelq eldrurs aq1 E tuoqu@$oa 'o pue (L96I 49 | .n,lalq Dupo\tDzo (B\ 5, s!uo8elEJ l?clSoloqdJotu r4:*;:or1e1ereql Jo eruos alElr FEa'DulpouDzo'snpoqt0uc r-.:t u3 i1o4 'eJeue8 e^lJ €l: ?d lere^es pepeles e^eq I sg'Idl^tvxs s's l5 ! Jd) s r{Jl prrrluB luopouoJ aql q luouala ed aql Jo uollJunJ pu? urJod 08 h auimal lch.s Sec.5.51 Examples 81 5.5 EXAMPLES life (Pl. 5.1, Figs. 3, 4). The oral surfacesshow that when one element has a distinctive den- I have selected several Pa elements drawn from ticle triad, for example with one or both of the five genera, Polygnathus, Palmatolepis, outer denticles enlarged, the opposing element Gnathodus, Ozarkodinq, and Icriodus , to lllts- has an equivalent enlarged gap between the trate some of the relationships in the different denticles of the triads, allowing the elements to morphological categories (see Fig. 5.2). intermesh. This intermeshing means that par- ticulate matter passing across the element sur- (a) Ozarkotlina breris (Bischoff and Ziegler face could be very finely mashed, almost 1957) and O. eosteinhornensrls(Watliser 1964), chopped. It is interesting to note that the den- ticles of j uvenile 1criod.us I elemenls are usually pointed, but those of adult elements are nor- The simple blade morphology of Ozarkodina mally rounded. If the function of the elements Pa elements means that there is no oral surface was to cut, one might expect the sharp form to to mesh with the opposing element, and, unless have been retained. the elements functioned as cutting blades, it would have been the lateral faces of the over- zlJ.2/24 (c) lapping elements which were the functional Palmololepsh tenuipunctata Sannemann, E:: :.i: e\ample shows 1955 &u : -_:::r!. trssue(lines) surfaces. Two examples, both fused clusters, ! !-:- -\=:::-1:i are schematlc are shown (Pl. 5.1, Figs. 1, 2). Despite their ': dillerent morphologies (O. brevis has a tall The large Pa elements of Palmatolepis are P.r' - -- : :. crest form - cusp) both show the typical blade tip to cavity good examples of plate morphology, with a flare overlap of this category. There does not large surface area, which is generally smooth. appear to be any difference between the inner The carina of these elements stands far enough fu = r:Lins surface becomes and outer surfaces of the blade elements, that above the plate so that the greater part of the 1ii:: r. but in detail the is between those that were in contact with the plate sudaces cannot have been in direct con- E:: --Lrre complex. Three opposing element and those that were not. tact with the opposing element (P1.5.2, oa : ::--rrrm upper surfaces The Ozarkodino blade-type morphology Figs. 1-4). t -:.:-rre. the hollow, and must have been a very successfulelernent style --- because it remained basically unchanged in L :late form has a large a (d,e) Polygnathus xylus xylus Stauffer 1940; P. direct evolutionary n--::. ]nd the element sur- succession from the Late yerri Stauffer l93t; P. parapetus Druce 1969; r i::.:n or highly irregular. Ordovician to the Carboniferous. Yet Ozar- P. siphonellus Druce 1969 hi5 :: !1pen area along the kodina was also the stock from which many, or \1lth most, of the Silurian to Carboniferous platform :E::-: the lateral mar- The specimens of Polygnathus chosen are style genera were directly or indirectly derived. B-: l- crest form is the assigned to either the plate or hollow mor- ; ri[.: :.-,im with only a small phological groups. P. xylus xylus and P. webbi : \'.'-: 'e mrdline of the ele- show plate morphology (P1.5.1, Figs.5-8), (b) Iciotlus expazsas Branson and Mehl 1938 .t.-::r lateral margins of and P. parapetus and P. siphonellus have a hol- lEi :,-,f\ tiom the opposing low morphology (Pl. 5.3, Figs.3-12). The 1-: arbitrarily defined The Icriodontidae have a very distinctiye oral plate morphologies show very little open space r-= fbrms exist -::dational surface morphology, with some genera having between trough and ridge when the opposing : i-:-:ornIS. a transverse row of three denticles, added in elements are closely appressed. The hollow t ' ::nent morphology be- growth as a triad (Nicoll 1983). By using some forms have a large open area in the central part t1:is .rre not great, but they of the numerous cluster pairs of I elements of the structure, and the carina is usually very g : r indicate some variation obtained ftom the Canning Basin (Nicoll low. Rib ornamentation on the platform oral pr-:;le: rvere processed by 1983), it is possiblein this caseto be sure that surface may be either transverse or longitudi- { ::::..dont animals. the two I elements really did fit together in nal. 'ueruo^oc raddn 'uollsuroc rardeN l€llprsnv uretsed!'ursp8 BuruupJ'fZ/8oEgJd\'lueuelo Uel Jo ulSreursdelie^o fllq8lls luauralelqBu Io uI3rpIlIruroJlEld seuu€. aqt Jo aull eql Jo lasJJoeql pue sluouels aql Jo sdrl louelsod eql Jo uolleredes Jo lunou? eqt eloN ,{aI^ rouatsod '8 IErelel 'sqdE.rSouuoloqd oolals 't (lqBlr) IzlgzJdJ ^ler^ 18€6 pue (UeI) OZLSZ)dJ 'urroJeteld 'lnrd luat|latapeltutsuotar t rallnels gqan nryDu84od-8 '/ sSIl ueruo^ecreddn'uorleturog rerdEN :ellerlsnv uretsed\ Suruu?J 'S/tO8 'gLIgZJd) rueuele olul palllj r(lasol. 'ulseg BJ/d Ual(9 arou pue pesoddo,{llcer|p ueaq e^eq plno^\luetuele lq8u aql eJllul lolralsod -,ra!^ lereleJ'sqd€f !0t61 ^tal^ rS -orr[uoloqd oera]s'S urroJe]€ld peteprsuoc seldu?xa eql Jo lsour esneseg '8urs 'stuauele ed ed,{1-Iurop"ld ur puno} sarSoloqd -seJoJd eql ur palsrss€ lueuJela 3d eql qJlq,?rt -rour luatuala crozoa"l€d elpp[u Jo sadfl ,(q poqtaru aql o1 Jo paurnsuoJ pooJ Jo ad,{l uoururoc erour eql Jo auros palerlsnll e^eq eql ur sJJuJr:JJrpol rJrllrJ palelel eq lsnu {3o I uollJunJ l"rll Jo uorlelueueldul loJ suo|l loqdrorx luetxele ur uoq€rJEA aql ueql 'pooJ -nlos pcrSoloqdJoru 1o e8uer apua € s€,t\ ereql treluts Jo uonse8ur eql ur lsrsse ol se,,r\lueurela ed ueql lseel l€ ro 'eules aql sr serceds eqt Jo uorlJunJ crssq eql l€ql peunss" sI 1rJI lueraJJrpJo Eueuele Pd Jo uorpunJ rrs"q eql JI 'xuAJeqo ro qlnou eqt snorr€A uI eryc ,(q paqsq SINIYUISNOJ Jo Er?d :NOIICNNJ Bd -q31se sluarrnr ralE,{ dq palloJluoc ?{llPnsn INSWS'IS 9'S sr sursrue8ro sno8eqdorcrur uI luerrJelolrr elJrlr€d pooC se^ers,{le^rlraJJa eJ? {oIIJ'tueu -e^our elul fle^4elar qll{\ seJnpnJls e^lssed 'ed,ft sql Jo sluauele fq peuuol ,{lerluesse eq,{stu pu€ uotpeler ro uolpalas -red uorlce e,uloa;;e ,(ue edecsaflsea plnoc derd qlr.Lr pal€rJosse flensn ere sernlJnJls salcrlred poo; eql Jo tsorrr teql uaes pFoxr a{ll-qtool ro qteel eq1 srusrueSro snoSeqd 1r flr,rec qlnour eql Sururlansstl gos ,{q paug -orJlu uI 'ssscord uorlsa8ur eq1 ur ,{ard eql yo -uoJ ssalun'areJJnsJo dq alrsoddoSutut;uoc uorlJnpal arrs]o sJruErlJJureql su IIJ^rse '3tnl ou sre:eqt 'f,8o1oqdrouao oq oql Jo sluelrrola -dEJ ,(eJd r{lIA\ pel?rcoss€ aJe seJnlcnrls e{rJun'pue 'ereJJnsluauala Sursoddoaql qlral a{I-qtool Jo qleel sql rusrue8ro^llBnsn snoSeqdorceu lczluoJ ur aq louuec ruro;1e1dreppoqs stql E UI'paUOrlJUnJ Sluelllele eql a\oq Jo uouel yosacepns eqa (7'I 'sEId'€'9'16) ftt,rec pseq -ardralur eq1 ur luelrodrur sr uorlourlsrp srql eqt .ra,roruropeld raplnoqseql pue aBpIJsulrer 'sno8eqdorceru upql JaqleJ snoSeqdorcru,(1qe q8rq pcrd,$ aql s^\oqspalersnll vsua)opou -qord lsour ueaq eAEq ol puru€ luopouoJ eql suDauryrqg ;o ueutceds eqa uouttuoc f,ra,r JeprsuoJI leql JlJrl atets ol luelrodut st t; 1oue:e fSoloqdrotulsarc eq1;o sa,ltleluesetdog 'se1,!s pcrSoloqdroru ;o eSuzr raplnd qJnur e lueserda-r leql stueruele ed rxrop€ld-uou L96l snorournu pue ur:oyleld Joqto er? ereql lnq pBorxall srs aropou sntDaanlq snpoqtouc g) [9 s 'ras s qcl JeruluB luopouoJ aln q luauolo Bd aql Jo uolpunJ pue urrod 78 ro:.i: animal LCh.5 Pa element function: constraints r::- : =:iorm and numerous ? ::-:::]ts that Iepresent a l: : styles. : :.:::--:phological here that I consider : : r r'ra\e been most prob- :r-: -i:.:: than macrophagous. ,: -: -' :3nr in the interpre- -:E : ::::'trS funCtioned. In a ri:- : :]i teeth or toothlike !:i .....;iated with prey cap - . , -.rics of size reduction IE- -=::::..n proceSS.In miCro- s-. ,- reeth or toothlike r, ...ociated with prey ::-- .:: mav be essentially " : --laii\ely little move- ::-::- : ..ie\es.Foodparticle e: - i=ier currents estab- . - -: ::rts of the mouth oI : -,:-: ::: r:sic function Ofthe rhe inoPcri^n ^f :-i: ' n element morphol- i. : : :o differences in the r^ rhe mefh^..1 hv .r +ar1 in the nrnrcc- ---- :Li : :-: :\amples considered : , Bureau of --PCl. , _ :. :rr1e\ Collection .: I Ptrelement pair. \:r,r. r Fofmation, i: ,.:: P: element pair, - :.. Indiana; Salina : -,:: I element pair, :::lement Iit lnto = : 1:.:ern Australia; j :: -::::o photomicro- ai,rried and more - . .- \r.rfalia; Napier ft - - ia:5 l0 (leit) and the amount of fE: _- ?:riormmarginof J-: - \ airern Australia; 'usruo^ac reddll'uorlsurrod slJrHur8lr^ :erlellsnv ulatsay'A'urspgSuluuEJ !t !€ '6/99€ g)A\ 'dsnceqt ot rouetsodstsetuela ueeaueq de8 etoN aer^ rorretu€ lpret€lrouur (Z ^\au ^{er^ Ierelq ratno lerol€roqe 'sqdefonruoloqd oarels'I (je^roD €ZlgZldJ pu!. Geddn)ZZLSZJ\J rS96I 'uroJ ateld '4ed^\er^ tueruela petrnrsuocar uuerrreuues Dtopundlnlt4 sldapuutlod-i-l s8tJ 99x IIV-Z S aretd anssn pelerTrJ peqseuJelur saJeJJnsIero ot Suldrl Jo sapFcqJrp eql Jo eluos lno petuod qlr^{ uorlJunJlou prp pu" pelf,erry s?r{ (086I) surow f,el,I.uo:) (2761ltorlN'?16I SaJBJJnSl?Jo Jleql qluir Pelueuo"{l[e4ue^ eJaA\ slueul rxoJlspurl) esn raql Jo errIrl aqt te onssn {q -ele sueuressrp eqJ (sg6l IIorrN) Sururuhs pare^oJ eJea\ sluauala ed aql l€ql pelsaSSns s?A\I€UrrU€ eql qcq^\ qSnoJql JoIEA\aqt ulo{ uaeq seq lr srolJ€J asaql Jo II? JoJ lunocJe oI peurclqo salJrusd pooJ-uou uro{ selJru?d PooJ 'eJqrns Jelno eql uo rselt\ Jo eJuapr^a Jo {JeJ a^UJadsoJdUos ol a^ars se pauonJunJ 'sluau 3 sr areqt lErlt puB 'porrad qu{oJ3 eql SurJnp " -ale suaureJsrp eql ]o Surlsrsuoc 'snleredde p:uorlJunJ e^eq lsnu sluJrusla 3ql lErll 'JJ"J aq1 ;o 1.rzd rorralu? agJ '(gg6y 'lzia s33ug) -rns ralno eql uo sr sluauele eql Jo uouerJJB IETUTU€eql Jo uorSeJ p€eq eql ur pua Jorrelus ql,,!\or8 eql leql el? eseq; uorlelerdrelut aql rPau paleJol se/( ernpnrls srrlJ lErxru? I"uorlJunJ uo slursrlsuoJ Jaqlo erP eJeqJ luopouoc eq1 ;o snleredde pezrlereurur eql 'eg S '8lC ur palsaEEnsreuuetu aqt Jo sluetuele rouelsod eql eJe s1uerueleBd aqJ ur 'Jeqlo qree pr?/t\ol pelro^\ ,{1er'qcagaa,teq lsnur sluerxala Pd eql'uo4rsoddo ur Suruorl NOIJVItrUdUflJNI -JunJlrJrll Jo J^rlesrpursr sluauele SurlsleJJalul :NOIIJNNd JNSI,\IS'II Bd eqt Jo ,{Soloqdrour eqt JI 're^e^1ol{ etets ''9 Sunsar eql ur flrurxord esolJ ur aq ol peeu faqt ppo,tn rou 'uollf,€ ur ueq,r.ralelr8rpralut 's11acSuqa:ces-eleqdsoqd paurel 01 sluerrJele eql roJ pasu ou sr araql esnereq -uoc 1eq11e1cod ploJ-urls e ol slueuele urnlel 'slueruele Sursoddo Jo drqsuoqelerralur pcrSoloqdrou.r ;o aer8ep q8rq z ecnpord 1ou 'slusrrraleeql uorlour Jo flqeqord p1no,naSesn adfl srql 'eceld 1oo1 Ieret€lslue^ard Xle^rt.oJle qcrq^i eul'rpc eql Jo,{Soloqdrour ;o pue quror8 ereq,n sptoJ ur{s e,rqcelord e8pu eqt i(q red?d oql Jo euEld eql ol papulser sl lueur -rrede-r -a^our IIV txet eqt ur passnrsrp seddt Ipreue8 eql Jo srled leql 01 paurnlar ,{aq1 ara,r uo4rsod Suqsar e lueulele Bd Jo uolFp elqrssod eqt lo seldu€xa T S BIC ur aJaA\sluaurela eql ueq,{\ ,(pg 'e-rnldec ,(erd roJ pesn pup pelrele eJea\ feql 'SuruollcunJ uotut_Lsod Jle,r sluaurelasluopouoJ Jrll ueq^l tBqr palseS /) (\ -8ns ereq sror{to pup (e9361) uols8uag rl SIXVxvlDT r. lsrl uorlrsoddo ur \l l"/ peuorlJunJ slueurala aql l?ql al€Jrpur leql ,{3o loqdrou lueurele uI suorl?rJE^,,Aoqs 'luauala e4 eql ,{po,{11ensnpuz'stuauale suapuntuoc eqt f,tuo'sdnor8 o,rt eseqt uee.rteq uorpunJ ur ecuareJJrproferu e -ro1sen8re sluauale suep -unluoc eql puE sluaurala sueulacslP eql uaa^\l -eq seruoraJJrp pcrSoloqdroru aql t€qt (986I 1ocr1q) areq,tasye 1no pelurod uaaq s?q lI ',{3oloqd.rotu ur secuareJJlpeqt 1l ureldxe leqt edfl pooy ut secuelo;pp aq feur qseu puPouroqs-e uotEl.tNv 1r 'sarcedsro Breue8 peleler ,{lasoycyo eru e:aq [r'9 tes I qcl IBrrrluB luopouor aql u! luaurela Bd aql Jo uolpunJ puB rrrJod t8 r3limal [ch.s Sec.5.7l Pa element function: interpretation 85 b rasp and shear 1l ll ------,lL \ VU tr€ : -. r : =.rion of Pa element rhe text,^"^ All move- E : : : :.e paper by the ridge a r - :' :--.:r\el! prevenh lateral r.,:--:,:-d pocketthat con- cl::: :: cells. I FL\CTIO\: ^ fE --: :'J:teflor elementsOf qr,--i -> ol the conodont =--. .i:i located near the ::-=: eqion of the animal - r . :nrerior part of the r= :: :he discernens ele- Ei : : :. to SOIIplospectrve $r -:r..d- particlesobtained cr-:- .:rrch the animal was 1::' The discernensele- c: :.: their oral surfaces - E: ::d not function with r=: Ciliated tissue 'r;d. =c-- : :lir. plate form, irL : - :\ l- outer lateral x. : : -'J:p. wCB 356/9, 'snoraJruoqrpJra^r\o.I 'uoDeu oc aSuel{lrng :Ellsj}snv urelsel\ (ursEfJeusdpuog ,zgIr,0g gcg ',{llq8rr qsaur ot lueuele lq8u s^\ollp teqt lueuele Uel Jo ur8rErurelno uo a^oor3 pue s?ulter !ZI Jo rrolleredesJolunoup eloN red luelueleJo rouelus lueuela lqSuJo Isro,II {edtselxoleJo !oI ^ror^ .rrpd ^{ar^!g .(lq3lr) Aer^ Ierelplralno lueuela g3l lelo '6 lueuela Jo louatsod pue {rrcd ^\er^ ,696I ^1er^ 6zLgz)dJ (DeD8ZlSzCaC 'uro;,lloltoq 'otuetuelo pelcn4suorer arnrCI m auo4atimqlDiitl;;-fti:g ;Brg .snoleJruoqjpJ.re^to-l,uoplruloC ,Zllgog e8uuu lrng :erlelsnv uelselt'urspg euedeuog fl}ff uoqrsod leuoqcunJrreql ul pecpld ars slueureleeqt ueqard€lro^o ol ur8reurlq8rr paBleluoeql ol parJrporutrltupotJruirs st tuolxeleqcee ^\ollp !9 Jo uISEur tJol eql lpql eloN luauele lq8u Jo lprrlEl / rrBdluouolo lo mrr^ lueureleqel ^rer^ ,5 .(1qir:)Ierelel Jo rred luelualaJo rolratsod't rredlueuela Jo ror:a1ue l pug ^1er^pral€l'S ^1el^ ^{en Z,L;ZJLJ oJoD9ZlSZJdJ 'rxroJa{olloq 'rled luauele patcnr$uo.or '696I exrlrq snpduDd sn|tDuqKpa-L-E s8tJ ,Elled 'uerddlssrssrll 'uoleuroC elloa :ea{ol',{lunoJ uorEt\l Jo quou,{llpnb pouopueqv SuporpunJ ele,{ s}ueuela ueq,{ lcEluoc ur ueeq e^sq louueJ leql sapr^ec peJs$ slrplnoqs Jo uorlEledJs ,I .(lq8u)lEseq Jo eloN Aror^rouetsod 'Z a\ar^latel 'sqderBorcruroloqdoarats gZLgZJd) pve Ofei nZ,LgZJd) 'turoJ lseD 'rrpd lueuelo peFnrlsuorer 'rS6I p?olxau slsuaropotusntoa llq snpol.ltDug-Z 'l sSrl '99x IIV-€ S eleld sluauale eqt l€ql stseE8nssralsnp pesnJ ur aql Jo qtpll{ ruur 8 I eql Jo ruru 9 0 tnoqe sluaurela qd Jo drqsuorleler aql 'Je^aA,\oH dn 1oo1 faql pessaldd?ata^{ slueuaTaed aql 'uorlJunJ8uDln3 e lsaE8nssaop sluauele 'pd ueqdA'rl]III SZ'0Jo sseJxeur llede pa^orx e^€q eql lualxe eurosol pu? 'qd eqr Jo,{Soloqdroru ol p?g o^eq plno.!\ slueuroleeql Iool EuqlnJ eql 'uorpunJ qlool € e^eq lou prp feql ]?ql € s€ petJs e^eq ol epslq eql Jo eJBFnsraddn .uorlEluorro petsaSSnsa^Eq (Sg6I '/16I IIocrN) sepnls eqt roJ snql elqrssodtsosolr rreql reqlo 'qlaal atejqa eA Jo sed,&etIlos pue urer€ slueualeeql uaq,ruru gz 0lnoq?Jodel slueuala tuopouoJ Jo sarSoloqdrourr?lrrurs -re^o epBlqeSererre ue e^?q [pnls slql q pesn aql uo pespq 'uo[runJ qlool B p€q sluelxele sepedssnqQuS{lod aql Jo slueuele ed eql luopouoclPql papnlcuoJ(6t6I) uossddel suo4elnJl"J3ur,to11o; aql 'ss€dot e^€q ur posneq llIA\ urrr 8 J lnq 'e8e ueurcadsqll^\ rslueurela plno^\ selcrusd eql qcrq^\ uee^ueq pue sanedsuee,qeq peuel el€q IIn\ rllpv^ oql suJursJsrPeql Jo sJlJrlusp JlarJslp srll Jo 3ul 'uorssardruocueuou-lsod i(q pasnec uoqrol -JEds eql rxo{ peuruJalap sr ezrs elJrUed srql -srp euos epnlcur,{zru pue Frrrrue 3ql Jo uors 'txtx I 0 ueql eJoru lou dyqeqord se,rr qcrqa\ -uaurp umurx€rrJe luesaJdarI[1( rrJrr]8 I Jo alJrued pooJ aqt Jo Jeleluelp eql JoJ lunorc€ ol arn8r; eqa '(Eg6I 'lo t" s33lIg) uo1ue.r9ruorl q8noua dluo lnq 'selJrIBd pooJ eql Jo eBEssEd peraloJal ueuroedslsJrJ eql roJ ruur 8 I lnoqe ot r{lluerJrJJnsU€d? a^orx ot ueql roJ seue,rr8 sr prurue luopouoJ eql Jo qlpft\ eql ^{olp fressacau ps 1r 'uorlJunJ Sut11nce erreq ol '(1961 peoxeg pue qocrtr1)tuetuele ?ursoddo uo prlleJ lou Jle sluJrxJla Jql JI 'uede pJ^our eql f,q pedd€lra^oi(lued are,r srr€dEd pue qd e^€qplnoJsluaualesuepunluoceqltsqlos'uors eql qloq ur sluauela leql elsJrpurol uraessral -u?dxe euros roJ q8noue elqlxau ueaq a^€q snql -snlJ luaurole Sursoddoaqt spr€1r\olpelurod lsnu pruru€ eqr Jo fpoq eqa Suruedo 1a11n3 lue(xele qcee Jo ec?Jrnspro eql qlr^\ uorrIssJ -qlnou eql Surpuno-rrns aJeds Jo ruru €'I esla^suElle ur paluJHoaJ:^\ slled luauJle Ed aql IIIJ f,lIS€e pFo^\ sluetxela eql pu? IeuruB pue qd qlog lurureleqd eql Jo puJ rorrJlsod eql Jo luaua^ou qlr^\ pelerJosse anssrl a^rl aql qtr^\ lueuele ed eql Jo epslq aql Jo puo -Jeuuot pue alJsnur eql puz (ur1s) luaun8elur rouelue aq1;o EurddeFa^oauros eq feru ereql 1o sra,{el rnog purruB aql Jo qlphr ,{poq Surlsar lnq 'sluatllela €d aql Jo iorJelus e]e sluaur eql uqll^{ pelepourxoJ? uaaq aAeq PFoc uors -eleqd eql '(1961paorxe; pu€ llocrN '€86I -uedxe Jo aar8ap s1{l l"r{l ,{loIIIun sl tI ruru I Z 'p p s?BIJg,)sluaurela suouraJsrp aql purqaq lnoq? Jo 'releer8 ulru € 0 ls?el le ueeq eABq lsnl pelecol eJe slueuele suepunluoJ eql plno^l lEuru€ eql Jo qlpr^\ aql uorlrsodSuulnJ '1e1n3aq1 ;o Suruadoeql spJEA\ot,{lroualsod e o1 peleredas sluauralo ?d aql qtrl!|I€lurue salcqredpoo; pelorrr sluauele esaql Suueloc lt s 'Jos s qcl IBrrrluB luopouoJ aql q luauala Bd aql Jo uopJunJ puB urroJ 98 r.roimal [Ch. 5 Pa elementfunction: interpretation 87 1: = ::nents separatedto a i ; ::r erfthe animal would : -- :r:n greater, or about : :: :his degreeofexpan- =- .-::tmodated within the :i - : =nimal.Four layersof rE: :_: :nuscleand connec- ra: : ::t movement of the fa.-:r lrould easily fill the ; -: . ::drng the mouth- : .::. .ri rhe animal must t = .- rugh for someexpan- r: _. > could have a : --:a::s-lements are not called On ::- -: . ir is still necessary i-- =: :irliciently to allow ::.-.:: but only enough ri=-:: -!..rf the food particle \ : r: ::lLrrethan 0.1 mm. :: red from the spac- -:: :'. of the discernens --:B :: ::. particles would :: r:.-:ed that conodont r-: :,]:::on. based on the e! r: a!-nodoflt elements j .:--:.are reeth. Other - -:!- :e:: - :::. crest 10rm, 5 : ::.::.-oi \ie\}. Note E- ;::a :i3ments were :--- :- \i:ssrssippian. q ':- -PC:5726 (left) r: :, - ' -;ieral view of r! -r' :.: ieir margin of ilE ; :_ t3 elementsare _': r - -1: Burr Rnnge . ;.r -- :pct5728 (lelt) : -i.' iareralview a: ot separatron_ :c -:._ : -ne\h rightly. .: - -::alrntferous 3q:ns 'luelxele €d Jo self,ls aql Jo f,luelnurseql pelou (9961) 'p o s??rtg use,nleq serSoloruoqpe1se88ns eleq sJaqlo rn :qr dlq€qoJd pue PooJ 'seurur.JJeluopouoJ roJ lJpotu e se lzttuayod pue'(egg61) uoslSueg'(796y)rlsareruez5 .pe1€laJ {s \ur uoDcunJ leqJ (pJ euos sJeJJoleql srusrue8Joyo dno-r8raqloue 'uoucunJ sr (snxorqdure 3'e) elzproqcoleqde3 eql ,{1aso1clsoru eq ol pe^erTeqeJ" sluopouoJeql 4 lalder{J srql ur passncslp 'euoq 8ur,{pa,roenssll snorrtllqJ qcrq,t o1dno-r3aq1uo luepuedepeq L!\ sluerrr +iri rueralJlp aql (t) JO SISTSUOC13ql eJnlJnJlS EUrlnc e,r43a;;e -ele luopouoc :o; palard.talur uorl3unJ eql aJ: $ slueurela suapunluoS ifuer' e seq lnq'esues peldscce ,{lpreue8 eq1 'sno8eqdonrru ere snxorqdruepue sno8eqd ;o ;lrrrJunJ aql teqt tsaSSns ur qleet slr€l elunt aql leql 'ool'raquleued -oJJerxeJ€ sqleuSoleeqcsz '1ua:agrp,{rea e.re + ur sasueraJIlc (q) 'qlnou eql olul Ixeql slJodsuell pue JalE^\ aql sursrueEroEur,l1 ;o sdnorSo,$ aseqlJo slrq?q '(seot uor; salcrlred poo; sernldec qcrq,t aceJins pezr Surpaal aq; sluopouoJ ;o se,rr1z1e-r3u1,r1 1)wersnlr pesnJ pue (€86I -lereuru e uo pedole,rep uralsds SuuaqteS-pooJ alqrssodse peuoDuerx,{puanbar; ueeq eleq ! lcalrlouoJpeq drutrqsuo1 pslellrJ € e^?q l?ql 'sprouuc eql s3 qJns'slusr snxorqdurepue sqleuSoleeqc'sJee{ luecer uI a :-e'rusrueSro eql Jo ql"ap -ue8-ro;o Joqrunu € pa{oolJe^o 'JaAoJ enssq 'ureyqordeql Jo uounlosar € JoJ t?- rE -raqlo qcee o1 dlturt SuqunoJsrpur'(0861) uoslSuag enssltpal€IlrJ laq lsaq eqt sploq ,{lr1ero1peq drurrqsuolu?rc tr ::;n stuauele aq1 (e) ro; lroddns s€ pelras al?q ,(eru luauele oql eqt ruog suarurcadspuorlppe ;o ,{ra,rocar :uotlcunJlueurela fiW (LL6i 11ocrp{q psodord eqt uer{t reql€r aqa dnorEeuo,{ue o1 ,{lesolcool Iprmuetuop ,a3e1s Fr[ .]o] seulTeprn8se ea.res ernlJuls lroddns-eyce1ue1e Jo (916I) surol l -ouoo or{l;o luaruuSrsserno lrruruoco1 rl.-r:tol Jql ',(leurrunsuI ,{ear'uo3,{q uorlsaSSns eql qlr.lr reloc anssn sql le 'lou pue uedo suoqdo -rnodaaq o1 luz 'seuneJ elenbe o1 papuel (9961) '121a s88rrg ,(ye4rt -lJodulrsr lI aJuelquraseJprcry:adns p sr eraql 1rcro1 a3ue: azrs aql JoJ srou aq pFo,{r uorpunJ 3urra11q e para,roc asnecaqlsnf sursrue8ro;o dnor8 Surlsxa aruos Dr...6- snonulluoclill^\ Pue -anssrl eJJ^i sluaruale ;1 alqeqord u:as olur uorlele-rd-re1urrno eJroJ ol ueql 1erxrue r:i t se qrnru se eq ,{eru ppo,r uorlcun; SurdserSe ra,roc enssn tnoqlll\ luopouoJ eqt Jo Iepou luetsrsuoJ{1eure1ur r ueJs a_yqaqa saSuetluoP peuortJunJ lueruala aql JI 'uorlJunJ luauela uE lcnJlsuoJ ol luel:odrur arou sr lr luql I rrs rp!$-plro,1 aql ureldxa uorlelerd-ra1uraql t3eJJef11ee.I3 ppom lueur 1o NOISO'IJNOJ 8'S r;-61 qqa6) tttl 9699 -ele egl Jo ernlcnJls cqeqdsoqd eql Je^o reloc Bu pqe sdep002 s€ Suol anssq luauurured ;o ecuesqe ro acuesard aq1 : ulera: ieur snxotqdue ;o 'sluau !E PllEI aql luopouoJ aql -ale eql Jo uorpunJ qloot e .ro1 tdo prp lnq 'ln3 | € .io] aleplpu?J aAIpEJlle 'dnor8 ,{ue qtr.,\irdlqsuoq€lal alrur;ep e 1se33ns lou lnq peqsuc'uI€A€ 'pue sece;:nsurroyeld lduri :rroseer leleles JoJ lou prp faql'peq drur-rqsuoluerg eql uro4 Sursoddoeql ,{q passeJdeq plnor'r salolu?d ',{Solore perelooer uaurrceds ey8urs e go fEoloqdroru 'uorpe Eurqsn:ceql Jo ,{JuarJrJJaaql ?urs?ertur - Golors{qd uI pelslJlser enssq-Uos eql Jo uoqelerdrelur f,reuluqard Jo llnser e sr lueuele e4 aqt 1o uro;re1d ue-r:uaiap sr'snxorqdrue uo pssuq sluoPouoJ eqt.;o s:rtrut;1eaJqtssod eql ;o lueudola,rep ar{J lI lnc lou plno.t lnq + puE :urJoJ PaJUPApSUE Jo reqrxnu B passnrsrp(9961) Trla s8Eug 'rallprx al€lnoru€d eql qsnJt ro eslnJqplnoa\ proqroleqda3 eqt ol peteler 'esnJo:un eql le arnsodxa sql urgql uaa,tleq lq8nec .rapeuralzlncrged n -ilaulradsar slueuale paleynlsod slr JoUB ereld 1oo1 qlrv'or8 era,t ,(ue Eu11orJo lpql eq o1 3u1o3 sr sluatrrele Fcj&)srp aql ol sanSopuB 1e1cod p1o;-ur1s aqt ol luauele luopouoJ eql eqt Jo uortJe slrtreJJa eql'1uerue1eSursoddo InYZTJEToUIUT IJe[ Selnl Sururnler ;o f,lncrgrp aql sr Jaqtouv stueur 3r{t ts€d fII?Jel€l Sulddgsqcea 'raq1a3o1a,rour 51 ooriuu:eg) snxorqdrue -ele luoPouoJ Jo esoql qll,{l a1q[eduo.rur aru uaql pu€ Jeqlo qJee uro.r;,{e,ne1nd sluaurala lrr: lnJnlu pue selctlred saurds qleuSolaeqc eql ;o qprol8 1o epour pue eql eraq.{r'39's 3rg ur pe1sa88nslEqt ol Jellurls Fexp pu€ seqcleJ l€ql ernl uo4rsodruoc aql leql sr sruelqord aseqt Jo euo sr stueurole?d pu? qd rlloq Jo uollour eql JI :-Jo laaq,{ aql Pu? 'leDelu 'yaporu qleuSoleeqc aql ;o suelqold roferu eqt 'lnr aq 01Pseu plnoa\ pooJ r :qr iurtelnSar e^ers lsllrul Jo euos pessnrsrp ,{lelenbepe 1ou e,req ,(aq; eql Jr elq€uortsenbsr 1r 'ruru Z 0 ot I 0 uztlt q aq1 ue8ro leaq,!\ eqt pu€ 'a8e uzuqrue3 ;o seldures ruor; paJoAoJeJ eJour tou ,{lqeqord '[erd pqualod uo suorl t illr€ltturs Jo seoJP ]eqlo slueurala luopouocolord o1 saurds qleuSoleeqc -JrrlseJ ezrs aql uer'r8 'oqY lnc ,(leAIFeJJa rrrro iu aql 01 uarutceds ;o ,{lueluls pcrSoloqd-roru petqnopun aql lou plnor urSreurraddn aq1;o sace;rns8uqlno I Jrr uollEtuatll8es lueIedde uo ,(pI?Iu pespq 'sqleuSoleeqc pu€ sluopouor 1erluelod eqt pue reqlo qcea lsed peddqs s qJl lBrxlu? luopouoJ aql q luauolo Bd eql Jo uollrunJ puB trrrod t .dmal lch.s References 89 €:rhs. based mainly on apparent segmentation of the trunk of their late bars, would have performed a slightly dif- r:::loeical similarity of specimen to the myotomes of amphioxus. ferent function. h : ttoconodont elements Other areas of similarity are the buccal cirri (e) Variations in the morphology of Pa rc::. of Cambrian age. and the wheel organ. The buccal cirri act as an elementsare probably a reflection of the dif- 1u..::lr discussed some of initial sieve regulating the entry of particulate fering characteristicsof preferredprey species. iot -:: chaetognathmodel. matter, and the wheel organ is a ciliated struc- (f) Food for the conodont animal was pro- :. :har the composition ture that catches and directs a mixture of food bably microplankton, and the animal was an I- \-: ::3 chaetognath spines particles and mucus into the mouth of active predator. A microphagous feeding t --:..:e of conodont ele- amphioxus (Barrington 1965). These struc- strategyhelps to explain the use to which the rG :.rrculty of returning tures lack mineralization but could be elementswere adapted. * :r ihe skin-fold pocket analogues to the discernens and contundens (g) Amphioxus is a possiblemodern relative !i:,:: atter its postulated elements respectively. If conodonts were of the conodont animal. If there is a relation- E r-: .-.i- - related to the Cephalochordata, they represent shipbetween amphioxus and theconodonts, it is E, ::sussed a number of an advanced form; and the present example, clear that the presentlife styleof amphioxusis I -:e conodonts based on amphioxus, is degenerative, that is more much more constrained than that to which Ea::: ol the soft-tissue restricted in physiology, morphology, and conodontsof Cambrianto Triassicage aspired. lr,: preCimenreCOVered ecology. ft--:: bed. They did not For several reasons amphioxus would be an ht =ship with any group, attractive for a modern relative of candidate ACKNOWLEDGEMENTS h-:: iuncrion of the ele- the conodont, Tbe larval stageof some species of amphioxus may retain a pelagic form for as Peter J. Jones(Bureau of Mineral Resources, he:;: of permanent tissue long as 200 days and move over distances of Canberra)critically read the draft manuscript ;t-::-- :rructure of the ele- 8000 krn (Webb 1975). This could help and made numeroushelpful suggestions.SEM f-=-,-: :he interpretation of explain the world-wide synchroneity of cono- photographyand plate preparationwere com- I -:t element functioned dont ranges. The life span of some amphioxus pleted with the invaluableassistance of Arthur r. i]-rnlng function would may be as much as 8 years (Courtney 1975), T. Wilson(B.M.R.). Publication was approved [ were tissue- and with continuous growth this could account by the Director, Australian Bureau of Mineral -.:]ents ft::-:ern rvould be more for the size range found in some conodont Resources, Geology and Geophysics, Can- li;:-: I rended to equate faunas. berra. b :-:_sestion by Conway In summary, the following statements may t-J:e-support structure serve as guidelines for further discussion of Pa nq :r \icoll (1977) that element function: REFERENCES r: ed as support for (a) The elementswere located in near prox- -n EcE 1980). in discounting imity to each other, at least at the time of the Barrington, E. J. W. 1,965.The biology oJ d:: : number of organ- Hemichordata and Protochordata. Oliver and death of the organism, as proved by the Gran- Boyd,Edinburgh, 176 pp. rvr::. rhat have a ciliated ton shrimp bed conodont animal (Briggs er a/. Bengtson,S. 1980.Conodonts: the need for a func- r a-. elopedon a mineral- 1983) and fused clusters(Nicoll and Rexroad tional model. Lethaia 13 320. lr-:=,: food particles from 1986). Bengtson, S. 1983a. A functional model for the E--s :hern into the mouth. (b) Differences in element morphology conodontapparatus. Lethaia 16 38. Bengtson,S. 1983b.The early history of the Cono- I -J3 rurtle lacks teeth in discernens and suggest that the function of the donta. Fo.r.rilsand. Strata 15 5-I9. E .en-. but has a very contundens elements were different. Bnggs,D. E. G., Clarkson,E. N. K., and Aldridge, le-:iie that consists of (c) The different styles of Pa element R. J. 1983. The conodont animal. Lethaia 16 f: lrone. discussedin this chapter had a generally similar 1-14. da- te.g. amphioxus) is function. Conway Morris, S. 1976. A new Cambrian lophophonte from the BurgessShale of British Ez:rns that offers some (d) involved the ingestion of That function Columbia.Palaeontology 19 1,99-222. :..r I conodont affinities. food and probably the crushing of prey. Other Conway Morris, S. 1980. Conodont function: fal- r:-i rhe similarity of the styles of Pa element, such as cones and denticu- laciesof the tooth model. Lethaia 13 107-108. re: erniJauqJJe snleredde rEl"Jls !::rllE lso(u os E,:: pare,\ocal ueaq e^eq f :-:.rrouol Jo ftrroteru eq; \OIIJN(IOdINI I'9 r-=::..r e dq pegoddns serrr ,rz:-- plqrssodsl lI slueurele u:=.-: snte:edde eql leql -5j:1!:1lJ pesnJ IxoJJ eJuap r ::nlJelrqcre srql s,{e.r-re tz : ur pe8ue:te sluatuale ;=::.tado'se,r1eq pesoddo n*.- :qr reql stseS8nseBelq E \rnbaun apt,,ro.tdlou m :--:--\-r qinoqlly snl?Jedd€ :::r:lrqJre eql uo slulerls n:: -i1.ror.te1sodpue ,{Ue !- ::: qur\ eouenbes .reaurl E:': aql Jo uotl€rn8lJuoJ 'ZIZ-6LI puE los'Jetg snsuadrasnpolnl r; ::uasa:d eJ" sluatuele '9e 'uopuoj lo {Dlros lD) lqel Ircpouo. -t3o1oo7aq olsodtu{S satDp)oq)ololdi( spe) eqlJo uorysoduloJ lueueleqlnw €86I S U'lloJIN r:*lrr>.uE peluaur {o PUB{unJl seueJJef S d lI pue uolSuureg iA f E :ul 'g77-217 7 sctst4doaCpuo [8opaD u'llDusnvto (r::'-: aqt sf,eldsrp 'utsuoc 'snxorqduEJo uorlnqllslp aql gL6I g f 'qqe,16 purnot AWg ellellsnV urelsel\'urseg SuruueJ | ::-:-lg uopueJg eql uorJ '018-908 9S ,{Solotuoapdto loutnol eqt tuo4 qsg procsruoeeledueruo^ec reddn 'sluopouocolord u€ ur sesnlerpdd€ :\t::Lr lBurru€luoPouoS v ueuquiEJ Suoue pesruSocer luopouoJ ,1.6I S U lloJIN seurdsSurdsej8 qleuSolaeqJ Z86I H 'ils^\euezs 'E9-I€ ItI Ddod lDlrads ISYUISSV 'ZLl 691 'EnL 18 uoda\ apg-uadg DJlDlaV lo f,ntcog loctSo|oag [3o1oozoa1o4 {a^!ns pxlSopaD s4ois palun 196I 'wals[S luopouo):(pel sepoqd l H cuI eu€luol^l uDttquDJ aql uo unrsodu.t[S ltiuor]Du)aiuJpuo 'auolseurl qJInC reeg gql ruo4 slelurueSuueeq -ras aq! rct adDd I]oqs :( pe) rol^eJ A n -tuopouoJ €16I nd H 'llocs pue 'C lA 'uollew ruI se8elquesselernlEu luopouor ueorlopr6 'vnL-6zL pue tSolouoaDpd 3urreq1e3-poo; r-i.-g g a (I'qtlursd tr\ lseIIJee ueuqueJ Jo uoqelerdrelur crSoyorq Ll ulsrueq.eu -oeled 136I H'rls^\Eruezspue g 1 'rlsrede; e se sntereddeluopouoJ eql i/6 L{ 'ruorlspurl 's€6-626 '8ZZ eI o|olla'I EIJaut ) plluoporopued tl [SolotuoaloJ Euerpul ruorJ $al -ele luopouoJ eql Jo uouJunC 086I '1 'uossddaf lo launo[ .ILI_egI -snlJluopouoc uEunls pesnl 696I v J {.ollod ZI Dtoqla.I y'sI] '.ursuocsrla 'I9-6i'xessns'lalseq.lqC 'uortJunJ lueruele luopouoJ 6161 1 'uossddel 'poo^uoH srllg stuopouoJ lo ,Go1o1qoao1o,1 'tzrlql n olDqta'Iswto! IUEluopouoJ v :(pe) aBpr:p1y f d :uI euerpul ureqlrou -relrursur pueed,{1 ai lapoaput p 1osasqendde luop tI|o.I] s.relsnlcluopouoc uerrnls Jo uorleuurexo -ouoJ ur tsellle8uerrelueuielA I/6I -I 'uossddef -au 186I g I p€orxeu pu€ s l{ IIorrN '6y1-Lgy tr7I3o (qdluSoaSoaDpd ' 1p 1-gg1 6 stcs,(tldoagpuo tSoloag uotlo4snylo -loiaoaolDd'tSolotDtulpoaoloJ' prunot AWg Erl€IsnVuratseiA'urseg 8uruueJ 'Iepour FuorlJunJ A\eu € :sa8elquresseluop erlt Jo ueruo,\ec reddn eql uory (1961 're13er7 -ouoJ 816I S J V',{estueupue f A'sBuIqclIH pue goqcsrg) sr^atq DutporltDzopue 0i6I 'reJ 'ttz-tlz LE uopuollo &aDoSlD Bqooz -Jnels snlfr snltx snq1ou8f,1oIsetceds tuopouoc aWIo Dlsodruts satDptol.lJolotd:( spe) seuegel aqtlo uortrsoduro.lueueleqln6 986I S U'floclN 'S d )I pue uolSuureg ,lA I g :uI tunlDpaJuol 'EIZ-L6I L sctslqdoag puo [?o1oag DuotsorqiuDtg Jo suorlelndod ueeucjrelrpel^l uDllDIsnVJo lounof AWg erlertsnv uJelsed\ pue ees qlroN;o e:n1cn:1s-e?epue sdqsuoq 'urseg Suruu€Jeqt Jo u€ruo^e6 :eddn eq1tuo:; -ele.rernleredruel eql El,6I II V ,1\',{eulrnoJ s qJl IeruluB luopouoJ aql ul luauala Bd eql Jo uollJunJ puB rrrrod 06 lqimal lch.5 - F.:r :: ot the canning Basin, - =\!7 Joumal of Australian lry':.:: - 197-213. ari ::nent composition of the [r, - .--:I: us .r11as ryfus Stauf- rr,:': .; breris (Bischoff and r --:- LFper Devonian of the re-: .\ustralia. BMR Journal gt. ::: Geophvscis 9 133 14'7. k---:=i. C B. 1987. Re- ti..-. :.rnodont clusters frcm t R. J Aldridge (ed.): tf-:+. : 'nrs. Ellis Horwood, A conodont animal from the lower Silurian of F-\=: Srlurian conodont clus- Wisconsin,USA, and the apparatusarchitecture of J, --.;: of Paleontology 43 panderodontidconodonts bj.l;ski. H 1981. Paleo- E :: Cambrian and earliest M. P. Smith,D. E. G. Briggs,and R. J. Aldridge re :.:lral assemblages.In: .: J -.- Pdpers for the Sec- !-: .::,nr on the Cambrian t: i:-i-_.J Geological Survey l--:: 169-172. - ABSTRACT taxa of that age.The evidencefrom older rocks CL=:--ognath grasping spines l:-r:tan protoconodonts. A conodont animal of the genus Ponderodus is, in contrast, scanty,and until Mikulic e/a/. &':. 56 506-810. from the Brandon Bridge of Waukesha, Wis- (1985a,b) reported a Parulerodusassemblage E :r::rtrurion of amphioxus. consin, displays the anterior portion of a seg- from the Llandovery of Waukesha,Wisconsin, rs::: -!nd R. P. S. Jeffe es i, mented trunk and an assemblage in which the not a single Lower Palaeozoic euconodont E. -rpo5id of the Zoologi- -9-212. elements are preserved in opposed pairs. bedding plane assemblagewas known. How- rr -r5- I The configuration of the elements, which are in ever, fused clustersof coniform elementsare linear sequence with the cusps pointing adaxi- occasionallyencountered, and representatives ally and posteriorly, provides several con- of severalgenera have been described,includ- straints on the architecture of the Panderodus ing Belodina (Barnes 1967, Nowlan 1979), apparatus. Although the single specimen can- Besselodus (Aldridge 7982\, Cordylodus not provide unequivocal evidence, the assem- (Repetski 1.980), Drepanoislodus (Smith blage suggeststhat the apparatus comprised two L985), Parapanderodas(Smith 7985), Pro- opposed halves, operating bilaterally with the conodontus(Repetski 1980), and Scolopodus elernents arranged in planar or arched linear (An et al. 1983), as well as Panderodus.Of arrays. This architecture is consistent with evi- these. clusters oI Panderodus are the most dence from fused clusters, which also indicate common and have been describedtiom many that the apparatus comprised seven pairs of areas, including Indiana (Pollock 1969), elements. It is possible that each half-apparatus Podolia (Dzik and Drygant 1986), and North was supported by a contiguous basal structure. China (An et al. 1983\. Unfortunately, clusters provide a less reli able basis for restoring skeletal architecture than bedding plane assemblages,as the 6.1 INTRODUCTION apparatusesare rarely complete and it may The majority of conodont natural assemblages also be difficult to assessthe extent of any have been recovered from Upper Palaeozoic post-mortem disorganization. Two of the strata, so most attempts at reconstructrng known clustersare, however, more complete aDparatus architecture have concentrated on than the maioritv. That of Besselodusarcticus ',{-rleruru,{sJo srx€ autps aql aJeqs '(CZ 3g 'BS86I) /r rarltnllyi ,{q paterlsnllrlsrrJ sged gos paaraserd puz snleredde eq1 (u) uelurreds 9x 'uosrpEl 'ursuoisry'AJo ,{trsre^ruO'IIlnesnIAI l9:1-. PP aBPuPtV) suelu ,{Soloa0'pZ/I00}iAn:Uel ot rooelup'ursuorsraA',{tuno] .eJ€rl eql l;i rorl?lueuSes ped€qs-A cqsalnPrlArrorJ lrrurrup]uopouo) snpt)puld | 9 8tJ .E pue eql pauorlsod sr a8elqruasse^poq eq1 (r) FlrrS€s eql ol lBrurou ]o t€ p tE ::'J"punoq eqJ uolPes :(e9g6y p G-:r arotu uaeq e^?q plnoJ cqn41ry) ler8alur are ,(eql teqt 1se33nsacuep F f::Jl Jqt Suop uotlelod -rla Jo seurlJo raqunu B lnq '(2 9 'I g SBIC) e8elqruesse aql se JEJ se pualxe lou seop FC :i" salruos ue^as ls?el EeJe pezrrcreuru aq; ,{1qeco1eures aqt uorJ r-:ru 8ur,{1-rea.o'dar81:ep rE :ql ur pue eltlederong q- J: luepl^e aI€ lelllrue 'IUnrl Jo uorlntuourBespuE snlursddpJo uoDrsod8ur^\oqs :spip^\dn rouetu€,E//l00t1\\O .tBu_rruD r,: --rtueulSas Jo seJBJI iuopouoi snpoDpu,.J Io Surmejp epronlerarup:)-Z-9 3U 'qun:1 peredel e :,.:iins raluJo] eql Jo lIX fq 01 suorl?luelJo luaJeJ ar: ,.-eurlads uotuPJC pue pocerlso prdocrlrpredel e ur elrlederong se r';--1 rqireq 1elot eql ]o rs:: :??q eqt pulqeq Sut,tlo: pas{lEu" leql ot relrurls Iereuru alrq^\ Jo ra^EI urr{l E sE pe,rreserd e:e sged eqJ punoJ x- qinoqqe 'pue 'atuJ Z Z Uos se,r rDuetsodeq1 iqels e e8paaq1 uo Ea:i esJql Jo euotr1 (9961 lou l:ed 1o pe,rreserd sr fpoq eq1 ped :orraque eq1 ,{p6 F:= Prslel Ut pe,trasard 1o E= uoluBrc eql Jo {uul .IVhIINV .urur rE r s lsBalle peqteeJ VHSS)OVM flHr .{O SJUVd rdOS Z'9 r :-:o (1uo (q pelueser E :-f:\ aql leql slseSSns :uo ieq ur8reru re1-roqs s-- ::-qire.r1sere s11er'r,tpoq '\986I'lo ra eiprrplv p:= iq patecunrtsr ut8rerq 'Eg6I lo 1as33r:g) pupltors 'uoluerC Jo sno : ::ft $ tunrulxeur e sulelle -JeJruoqJEJ eql uro{ sleulue ueeceqleuS,{yod -:\: rr ! :lunrl aql lutod srql pa,ueserd-ra11sqeqt qlt,r suosrredruoc,{-reu pt- :qr sluaserdal lueuraln ur!q -rtrrlardslrurad'pe^rJsJrd,{;:oodtraq;e'st:ed r=T :as ) uorlJ€dujoJ l"llue^ UosJo aruas:.rdeqt 'Jror-urJqlrnl sasntereddz E :'-1 SV SSOTJP IUII] g'I uuoJruoJ Jo uorlBrolsar lernlJelrqcre eql JoI q. ,nre:edde 3r{t pulqeq etep ot eJuepr^e elq?qer lsou eql seprto:d 1rse rc:-u arE slred qos aql 'acuegodurr:zpcr ed Jo 'rele^\oq 'sl (q'eg86I -a:;j: \poq aql sllullap Pue 'p p tr:rl4ryg) Bqsolneid Jo etrrrrolop snoerel :c: 3ql ]o tuelxe eql slues 13re e8pr.rg uopuerg aql uror; e8elqruesse r-I\--:=-l?ql'eeJB sIqI 'xIIlEIu aueld Suppeq snporapuad eql luepr^e d:: :ou tnq 'qlEauaq puelxe st a8elquasse aql;o uorlezrue8rosrpuralJoru a::je:ong a1rq,t eql 'qe1s -lsod aruos q8noqllz 'Surssnu tuetuela euo .:ielqruasse g aql ot qcled ,{1uoe,req 01 stueas Erlopod Jo ,{Je^opuel'I eql E!--rr spualxe Jatlal pesl€r ruo-q (9961) 1ue8f:q pue lrz(,(q paqucsep .r r:zp e 'olPq eqt uqIA\ Jelsnla snpolapurd eql ,{\or aql Jo pua 'eBPlquessE euo 1elueurJlJ rxJoJrpolsro ue Surpn;:ursluaLu or :ql qtoq sedolsr'ue -ale pasnJ,{11era1e1yo serJasE qtrll'sntzJedde rF ijsrppar leraue8 eql -JIEq errlue ue f,lqeqord sr 'puepeero quoN i :,ed 'o1equoqf,nper (ur) Jo uerJr\opro rle-1 aql uorJ '7961 e8pr.rply " 9 g qJl VSn'ulsuoJsrl\ Jo u"rJnlls Jal{ol aql uorJ IBrI|luBluopouoJ v Z6 Fr{I [€ ""S fSA lch.6 Sec.6.3l Development of the sp€ciesconcept in Panderodus 93 -in. (iii) a reduction halo, pale grey in contrastto 6.3 DE\'ELOPMENT OF TIIE SPECIES the general reddish colour of the rock, CONCEPf IN PANDERODUS envelopes both the soft parts and the assemblage. Prior to 1966 the elementsof Panderoduswete Within the halo, a dark grey area of slightly describedin form taxonomy, with distinctive raised relief extends from the fluoraDatitic morphological types given separate specific patch to the assemblage.At the edge of the names.Bergstrdm and Sweet(1966) produced slab, the white fluorapatite can be seen to the first multielement reconstruction rvhen extend beneath,but not beyond,the dark grey they recognized that two element types, matrix. This area, therefore, probably repre- broad-basedforms referred to P. compressus sentsthe extent of the underlyingfluorapatite and narrow-basedcostate forms referred to and delimits the body trace. If this is so, then P. gracilis, belonged to a single species.This the soft parts are narrowest immediately two-element concept was maintained by sev- behind the apparatus where the trunk is eral subsequentworkers (Cooper 1.975,1976, 1.3 mm across.As the animal is in dorso- Barnes 1977, Barnesetal. 1979), who pro- ventral compaction(see section 6.6) this meas- posed different designationsfor the element urement representsthe width of the fossil at types (Fig.6.3). Cooper(7975, L976) rccog- this point. The trunk widens posteriorly and nized that a more extensivesuite of element attainsa maximumwidth of 3.4 mm before one types could be distinguished within the margin is truncatedby the edgeof the slab.The narrow-based costate category, but Barrick body walls are straight; extrapolation of the (1977) was the first to describe them. He shorter margin beyond the edge of the slab attemptedto demonstratehomologies between suggeststhat the width of the trunk (rep- these elements and those of the symmetry resented by only one margin) may have transition series of ramiform apparatuses, reachedat least 5 mm. This contrastswith the applying the notational scheme devised by trunk of the Granton animals, which are all Sweet and Schrinlaub(1975). He considered preserved in lateral aspect (Aldridge eral. that the broad-basedelement was an M ele- 1986). None of thesespecimens is higher than ment, but gave no reason for rejecting the 2.2 mm, and, although they show some nar- possibilitythat it representeda P element (cf. rowing behindthe headthis neverexceeds 50% Cooper1981). of the total height. Although the Waukesha Sweet (1979), while recognizing that the and Granton specimensare compactedin dif- narrow-based, costate forms could be sub- ferent orientationsto bedding,the dimensions divided,did not considerthe notationalscheme of the former suggest a larger and more applied to ramiform apparatusesto be appro- taperedtrunk. a&r\ ..:! oi Pdndercah6 conodont priate for Panderodru.Instead, he deviseda set _F$rrds; showing posrtion Traces of segmentationin the Waukesha of purely descriptiveterms (Figs 6.3,6.4) for -_'::I !ej-:atrarion ol h unk. animal are evident in the exposed area of all the elements distinguished by earlier fluorapatite and in the immediately adjacent, workers and also recognized an additional In- The mineralized area dark grey,overlying matrix (Figs. 6.1, 6.2).Ar twisted element, which he termed 'tortiform'. r ::r as the assemblage least seven somites are delimited, and extra- Thus, he proposed a quinquemembratecom- I a lumber of lines of evi- polation along the trunk suggeststhat there position for the apparatus. Although these tter are integral (Mikulic could havebeen more than 20 in the preserved elementsmay not be comparablewith thoseof section.The boundariesare straight and near ramiform apparatuses,it is conceivablethat a : fo F\):itloned at the end of normal to the sagittal axis, in contrast to the set of homologiescould be recognizedwithin V-shapeds€gmentation of the Granton speci- coniform genera, and, in an attempt to mil preserved soft parts mens (Aldridge et al. 7986). produce a more widely applicable scheme, r arl. of symmetry. Armstrong (in press) has applied a modified pu€ al?Jqrxeurrq ott\l puB snlDiso)lun '(186I) saureg pu" ua{c€r]rtr pu€ (186I) +::!. ladolorloq PalseSSns 'sad,& d J ,tq pu" uosu€Jg E!\ iq pesn ISJIJ,{Soloultu 3l"rquerurJl eql f,q perJld,q auo 'suorlrsod seurPg PUe uelano51 'lqal l purlsrp ed,t eql ot peltdde i::1 saddl luaurale ualas -uroc snleredde aerql parllluapl (I86I) snw$orrun d 'seroads (txloJll?nJle) eJour | :-.-i:irq puE uaur seulBg puE uEl,roN slueurelJ sE^\ lde3uoc el"rqurelxltl 'eldulrs V {lz(I 'snpotapuod 'p p [--:: naql uI pu? 'esPJ eql alelsos l?Jr]laurur,{sepue'(ruroylcerS) elersoc ol \6L6I) uorsle^ p::<:-lSns erlopod rrJo{ sr?J4 '(uuoJrssardluor) paseq-epli pepnlcul oq^\ saurBg Jo uon€lou III ed,{J aql Jo q: :=--rn1alelduoc-leeu eqJ sr\lureddgsnpoopuod eql Jo Euauralaeql roJ sroqtnEsnoue^ ,{q pasn,(Solourlurel € 9 8rc 1 :;.. or dn euroq el€q ol r+-- qJEa '3d,{loqdroru luPJ =--: rnl e se ]r Eurlerdrelut uJ_lolrlJol r.r---* I1Eur lueserd lueurele u-' .:rrads auos ur %I [Iuo a:: qs-noqlle'lueluel3 ,rl, rllJol ts sg J ouloa {rJroj!?lol Fh:---. 'parrEdun up papnlJur a:::a;-oloruoq aq plnoJ [lE r';:: qrea ul sedfl lueurela L!JOlliDncJo rllroJrloncJD r::: -:ldurexa rol '(€86I) :::: ur lueserd eq r:::leJ lEql pelse88nsa^€q^€uJ uJloJrlrurlsD [::- :l]qt OIUI UOISIAIPqnS3 urJoJlllf,or6 rE ':?lu asaqt leqt pelse88ns d Jo} rleuJur As ulroJrlrLurS rrqJtals ulalaq -E:- - -r-s ssord ul 9861 c::: :..::Jrlcse6-y 9 3rg 6uoJlsuJlV JC >1rz lu06,( B 0 sauroE ? uDli oN .r- urJolrlJol ulrolrflol urJolrxoldurts urrolrlDn?rD uJlolrlrursD alolsoc rliroJllruls 9/6rSL6L urrolrlrLlrs Jadooo rP!e(l It'9 tes 9 qJl VSn 'ulsuoJsl t Jo uBlJnUSraraol eqf urorJFurluE luopouoJY n6 di. tSA [ch.6 Sec.6.3l Devefopment of the speci€s concept in Panderodus 95 similiform osimilitorm orcuqtifofm iorliform folciform s milifor m c\ osrmiliform orcuotiform L''''.--4 -___/ folc itor m ---=::t/ toriiform a 0 a t I Pondefodus !nicostotus (Bronson & Mehl,1933) lmm A rrnstrong Fi8. 6.4-Descriptive termsapplied to elementsolthe Pandercdusapparatus by Sweet(1979) andfoUowed rn pfess herein.Sketches show inner lateral face(upper row), outer lateralface (middte row) andtransverse section (lower row) of eachelement type. Symmetricol suggestedthat thesemay provide the basisfor ramiform apparatuses,which they illustrated a subdivisioninto three genera.Other authors by application of the notational scheme have suggestedthat rather more elementtypes devisedby Jeppsson(1971). They considered may be present in each species. Jeppsson the tr element to be paired and to be sub- (1983), for example, identified eight to ren symmetrical in most individuals. From their element types in eachapparatus, although not clusters,Dzik and Drygant (1986) were ableto all could be homologizedbetween species. He distinguish two distinct element types within included an unpaired, completelysymmetrical each of Sweet's similiform and asimiliform 'tr' element, although it had a frequency of categories.The similiform types were termed only 1oloin some species.In order to have this 'pl'and'tr', with the pl havingthe more pro- element presentin all individuals,rather than clinedtip. A similarcharacteristic differentiated interpreting it as a juvenile, extreme,or aber- the 'sp' and 'oz' elementsof asimiliform mor- rant morphotype, each individual would have phology, with the oz element being proclined n;:' i_.j :Pparatus, to have borne up to one hundred elements. to straight and the sp element proclined to The near-completefused cluster of P. unicos- erect. Size differences were also recognized h-:.=d (compressiform), tqfi6 from Podolia suggestedthat this was not betweenthe elements. I =tr asvmmetrical costate the case,and in their descriptionof the speci- We havefound the termsproposed by Sweet rE-i \owlan and Barnes men Dzik and Drygant (1986) recognized (1979)to be the mostapplicable to collections lF ::!tncl apparatus com- sevenelement types. They employed the ter- of isolatedelements, and employ them here in liE: I'v the trimembrate minologyfirst usedby Sweet(1979), but also our discussionof the apparatus structure of Te r:membrate types, and suggestedhomologies with the elements of Panderodus. r:- : aql dsnc paullco:d 'ue^r8 aJ€ sjaqujnu asoqa\ 'lueq8urlloN ,(lrsre^run ,{3olo}uoeEledortlw aq} suoDrellor eql ul {luEroduel suaulreds llv 820/1619 ;r--- :'i? aseq Suoye slrqrqxe Jo lruIl Jo 'lue(]lele urolrclpJ s,{er^ pratPl ratno pue rouul 'J 'H luelllala uuoJluo} lo IEre}El reuul Jo Zg0/26lS ^\eI^ r :.: ..1 pesodxaro pelJes 'O 6Z0lL6L9 'luelr-Iale uuo]llltuls Jo s^\el^ IPlelel lelno pu? reuul 'C'a 0t0/26L9 'luerxela uroJIIIUIse pue relno'g'V =-:- \rpul slred Jo s^\er^l?ral?l reuur pue rerno'C'J It0/2619 'lueuele ruroJllen.l?Jo s^\el^Ierelel reuul 1o 11eus reqselnx/d pet?losl-S nrl 09x IIe Jo aBpug uopuErg aqr wor! sn,oBorlun s poDpuDd Jo slueuele 9 c -- ued lBrluec aql uI 'rorJel ,- - :ue 0I) sluerrralaalqP :-:-:-?rI lalleus Jo Jr"d v :-- :.>:1r eql pue 'snl€leddE -ir leurou reeu aJe :.-: - -; rrsr{l pu€ seseqrreql ,{8o1oqd [- -+ 9 9s8rg) L. :::qr pue ',{lyeco,rtnbeun : sluauele f1uo - -: i '(ss Bts) L::- -. uorl ,{letluenbes r;:: : aqt uorldr:rsep ;o l : :-i1\ sr ef,eJIsJalB[ euo : i .'-.:zd ls?el l€ ere lsou --:.:i]llp sI sed,{l 'i lueurals c- . :rrr-Iatsodturod sdsnc . -l ::::\iuell ele SlUeIuelS .^1 .-:;:edde er{l Jo aurlprtx r: :-::Ieur eql qlllA 'SJled r-:. :r.11ur slue(xele aqJ, ", }tIOtrSOIVd VIJSf,)INYM rc s 1r\ llvddv sHJ s'9 'lelJel"ur qJns uI ;' : : \lajEr pJnoJsluelrlele L' : _ ?r: \as uor] slueuela :+ : ur de1:e,ro alaldruoc :: : ::inurs ,{Iaso1cesaql ut snJrlsoJrun ur ezruzo otDurlJullod Durpol.rzzp'(peorxeg pue :- : -e crteueSoluoleql eq eql qll.[{ II3^\ trrJoJUoJ d 11ocr51) 'suerurcads pue i -. >--r:uraleuuolllllllsE PUP -ca-re,r sedf,1luarrrela aqJ 1 1 dluo o.tpt)Ll outpo7azo '(peorxeg 1loct51) t:: :. jJ puE azISur uorl€IJe^ ,{q paluasarder snJoAJnJ J qtrl 'sntolso)tun sn1uad snpo1ng '(-res111e6)4a8anll 2snpolng '2' pue uosuerg) snlzl !- -- :-rlaruelePalPlosl eql uI aq o1 sercads tu?urtxoP eql s^\oqs (€€6I 'ds ZoulporDJuae'(1qa11 ]" . .-c \q apErrlsuolsl^IPqns lqel pue uosuerg) Irnossll Jo uoll?urloC -soctnb1qo snpoltsdoq :sorcads 3ur,lro1lo; aql :: : : qPun ueeq a^€q eld e8prrqureg eqt uo{ prreleru ed,{1odo1 qllt sepnlJur pu€ esJeAIp ,{lelEtopotu sr paJeaoJaJ 'i0tIizz:vItz eseql Jo uosu€druoJ €unEJpat?losl aql lo o/oZL uorpellor luopouoc aql ,{qroN O U rO tr..\:-r arE aseqJ (S 9 alnlrlsuoJsnpuapuDd Jo sueut:ads 111 aq1 fq passaco:duaeq s€q,{trlPJolEqselneid aql g L: -:-: ruroJrl.rolpue 'uJoJ 'sJelsnlJpJsnJ se Pa,uasalda.lu 1e e8prrg uopuerg eql urorl aldrues 3{ t V ::- ixlOJllIuJIs'lulOJIJleJ slueutala snponpuzd eql Jo euroS ds snponstl tsntDlsorlun :;: rr5 iq senads snpotap 1oA pu? snponpupd '(sepoqu) gccru{ NO(NVUS flHr hlou'{ :::-I JO SUOIpnJISUOCaI sntD^.tnrat snpo.tapuDd'(peorxe11 pue 11ocrp) SJNSWU'IUINOOONOC (Igrvlosl t'9 h e qJl VSn 'ulsuoJslM Jo uEtrnlts ra^{ol aql llto{ IBIIr!uBluopouor V 96 "qr rin. USA [ch.6 Sec.6.5l The apparatus of th€ WaukeshaPa nderodus anirnal 97 s.'. Panderodus recurvafi^ reconstructions of Late Ordovician Pan- JP-- unicostatus, - and, Wal- derodus speciesby Sweet (1979), comprising :o: -:te Panderodus elements falciform, similiform, asimiliform, arcuati- kei clusters. form, and tortiform elements (Figs 6.3, 6.4, rc ':: Panderodus constitute 6.5). These are present in the ratio I r=:::a- Comparison of these 2: I.4 :2.2:1 : 0.3. rE-- r lrom the Bainbridge We havebeen unable to recognizethe subtle iH-::: (Branson and Mehl subdivisionsmade by Dzik and Drygant (1986) &{finant species to be P. in the isolatedelements from Waukesha,as all ?- -::la atus represented by variation in sizeand proclination of similiform l T1: element types we rec- and asimiliformelements is gradational.It may rr-. conform well with the be that ontogeneticand intraspecificvariation in these closely similar element types causes complete overlap in collections of disjunct elementsfrom severalindividuals, and that the elementscould rarely, if ever, be differentiated in such material. 1mm 6.5 THE APPARATUSOF THE Fig. 6.6-Conodont assemblagein the Panderodwunicos- tdtrs animal, UW4001/7a (part); cameralucidadrawing. WAUKESHA PANDERODUS ANIMAL The elementsin the Waukeshaanimal lie in uppermost,with no costavisible to the anterior pairs, with the members opposed aqoss the of the furrow. The element is probably midline of the apparatus.The long axesof the asimiliform, as a similiform element would elementsare transverseto the trunk, and the havea costatefurrowed face.Elements 1 and 2 cusps point posteriorly. Identification of the are a well-exposedpair and are in contact elementtypes is difficult, asseveral are broken, along most of their length,unlike the falciform most are at least partially obscured,and only pair which are in contact only at the points of one lateral face is visible in all cases.For ease maximumrecurvature. The furrowed facesof 1 of description the elements are numbered and 2 are uppermost,and no costaeare visible. sequentially from anterior to posterior The recurvatureis greater than shown by ele- (Fig.6.6). ment 3. and the basesare shorter.The distance Only elements 8 and 9 can be identified from the anterior margin to the posterior mar- unequivocally,and theseare of falciform mor- gin of eachelement appears to be greaterthan phology (Figs 6.6, 6.7). The upper edges of that in element3, althoughthe latter is incom- their basesand their proximal anterior margins pletely exposed.The combination of a broad are near normal to the midline of the acostatefurrowed face and a relatively short apparatus,and the cuspsare proclinedto erect. base suggeststhat 1 and 2 are arcuatiform A pair of smaller,fragmentary, and unidentifi- elements. able elements(10 and 11) lies to their pos- The elements of the apparatus in the terior. Waukeshaanimal are similar to the majority of IEE ri \\'aukesha,all x60. In the central part of the assemblageonly those in the collection of isolated elements Er r:: inner lateralviews of small parts of individual elements are pre- from the same locality, indicating that the i':c. ::menr. 51911029c, servedor exposed.To the anterior, element 3 specimenbelongs to P. unicostntus. I tFz\ of falciform element, a1r:.+ Lnit. University of exhibits a long baseand the proximal part of a If we are to use the Waukeshaassemblage proclined cusp. The furrowed, outer, face is as a basisfor restorins the architectureof the sr 'aJnlJelrqcr€ eql le^el?qna 'snl€J€dde JOUeISOq plno^\ cL :::::.1i Jq aql l€ql saleJlpul 'seues J€eull oa\] uI ueur eql 13 asoql usql paJeds ,{lasolc erolu arE Fj r ij ?r1u3^ rv euEld^eql Suop peceds'slueruela a8elquresseaq1 rolrotu€ eqt le sluauelg (3) s 9 8IC -rJedseql Jo srxe eqt ;o -:::qrod,\H-6 '(9 pu otluoc) tovalsod eql ;o uorlern8t;uoc pa,rresard aql 'paePUI g 6 1 rue8.{rq e 4rzq -n ol rorJetu" tuo:1 tou sI lI 'azIS . .' tti 'a3e1qLu:sseeqsJ{nplA aLltarnpord lou plnoJ leuotleper8 -l D Surppeq o1 a13ueqStq € le ernltellqJJ? IelpEJ tueurala ur uorl€rrE^ sr aJeql q8noqllv (J) .^q e.1oSuruatteg pue asdello:;o s:ss:rold leturou 's BdlalunoJ rtaql;o sut8reruPszq puelxa snleredde l1 6' 'oJuaH 'srxE {un! eql qll^\ UBIJeplno,!\ sJeqlo eqt o1 ,{lelzrurxo:ddz (e) 1nq 'uorlrsoddo ur lsor ol euroJ plnoJ sluaurala -Jleq qJea uI slueruale eqt Jo sdn dsnc aql ,U 'laslJo puE etuos 'pelll ele/ peoq aql 11 pasodrutredns teq^\auos et" 'Je^e^\oq '0I JrJ,r stueuJlJ aql qJrq^r ur aSelqur:sse g slueualg aullpnu eqt ol 1a11e:zd.{lq8nor ue ecnpord plnoa,r eueld Surppaq eqt ol sunr snlercdde-11eI{qJea uI slueurale aql Jo 'Iunlt eql ol lsedser ol umerp au11y (p) lsrrjrou pauelleu aJnlJalIqJJe Ielpe] asJa^su?rl spueleurrxord aql tJauuor a1z i :ouuBJ OJITuI SnlEJedde '(tsourraddnsa3EJ'pe,t'ol V eurlprru eql ssorJe pasoddo stl€d tuaurala Jo ol E '---'Jop a'I) Sulppaq sarros reeuq e acnpord lou plnoa 'uollPluauo -JnJ 'Jelno eql eleq lred eql uo slueurele eft !rq.= r-,aualleu {unJl Pue ,tue ur'rauuoJ aqt;o SurueleJg'(3 9 3rg) srxe yp) patuerro ,(1-reluts a;e sruauele llv (J) :!: :-l1o d€Jle^oasolt ro 8uI 'e8elqruasse ror:a1sod tunll Jql ol esle^suellro 1a;12-redlJLlllJ s:xP eql Jo Pue r!.2: ]ino t\ srql sESutPPaq ol Suoy :reqt qtr.t pe8uelte aq plnm slueruela eql spr€,\\ol all slueulele ruloJIJIeJ eqJ (q) ueeq e^Bq ot flelqun ',{Foue1 r:'-:-r 3ql'arnlcallqJJe lelp€l qll^\ snle:zdde ue u1 q1r,t '(Ot S 'O S '8 9 SAIC)paqcre ro aueld z ut reqlta -sod pue ,{11erxepeSurlurod sdsnc E :-, C \.r \ louelu€ 's,{eue :eaur1 Sulsoddo ur ro (:lduexa:o; torretsod ol lor.lJluB uorj a.ruenbes.reautl i: U:i -:lrrll IPllue^-oslop eql 3u1q ur pa8uerre srred ur sluauala eql (3) E : :i?d lexe 8uo[ Jleql 'sferdruel;o csrp lero eq1;o f,laruru,(s -rncco eq E-: i:::qrod{H-8 9 3rJ -ruasar) lerpe: ueeq e^eq plnoc luerua8uerre :lunoJce olul ue{e] lsnlu eqJ slueuele eqt uorlrsoddo uI llnsal teql sdrqsuorleyar lueuJala-relul eql Jo salnl l,a:-? ;o plno^\ uorlsluarlo slql ur selnlJallqJJE lueJeJJIP -EeJ lero^as erE elaql 'snle,redde snpoDpuod Jo raqrunu e ;o 3utua11e1gpeua]l?U ,{11er1ue,r 'secpJrelno pen\oIln} rlaqr tr€ldslp stua(ele -osrop uJeq seq lEurluE:ql leqt tue:eddest tt IIE lpqt aloN lspr xatel Jo qderSor.ru uo:1ca1eSutuulcs pup 'slrcd Uos eql ]o eutlplu 3ql qll!\ s?plculoc peralters-trecl :(uudralunoc) qf/I00t,4[n'lEturue ]rrt4 eueld srqa Sulppeq aql ol IslrJrou st,{:1eruw,{s -sotun snponpuDd aql q e3?lquessE luopouoJ-/ 9 3!l Ieraletrq Jo eueld eqt t"qt e8elqruess€ aql Jo eullpru aql ssolce srted-1uaruele10 sreqtuaul eql ;o uoqrsoddo eql tuo{ luepl^e sI tI '1:potu {ue uo slure.llsuoJ;er:,"es sasodtut sluaurale Jo luerua8uerre aql 'sselaqleuoN 'sasnl€rBdd" tuopouoJna urloJluot JoJ e3uep -ra.a a1qeredruoc ,{1uo eql sepr,rord uerurceds eqse{ner!\ aql sasnle-redde ueaceqleuB -,{Jod;o a.rntr:trqJle Jql lJnllsuoJer ot a8e snoreJruoqreJ Jo sa8elquesse eueld SuIPPeqJo p pesn (net) 'p p 'a8ptrply 'eurnlo,r srqt^lauel Jo t rald€qC u1 aueld Surppeq e oluo snlereddz lzuorsuerutp-earql leut8t.to eql 1o 3ul -uJnpUpup asdellor;o llnseJeql st :Selqtuesse EqsJlnEA\Jql ur sluJurelJJo tuaureSuelle:q1 snoovsoNvd .{o suoJJsJIHcuv sorvllvddv 9'9 [9'9 ces 9'rtJl VSn 'u!suoJs!!\ Jo uBlJn[S re^rol aql luo{ lErulu? tuopouor V 86 rdE. LSA [ch.6 Sec.6.61 Apparatus architecture of Panderodus 99 i {RCIIITECTURE OF {: =re nts in the Waukesha s- -.i collapseand flatten- ft.-:;-;imensional apparatus rk:: Ir Chapter 4 of this zt: a:- r 1987) used a variety G: G:-- re: r =gesof Carboniferous I =: lrchitecture of poly- - rr=\=s. The Waukesha t :r- !rnl\ cornparable evi- i :-:lnodont apparatuses. :--- ::!ament of elements r=:r:Ii on any model. E -- i opposition of the *-r:--: acrossthe midline of k ::: plane of bilateral | : - ::; bedding.This plane E*::; oi rhe soft parts,and fu has been dorso- anterior -rnal - F::::ning of a number of fe: ::- :his orientation would D r:: the elements. The anterior B I ::-.; oeen radial (resem- Fig. 6.8-Hypothetical radial architecture for the Pandercdus apparatus. A, with elements arranged with (r. -,:: e.raldisc of lampreys, their long axes parallel to the trunk axis, anterior vrew. B, the assemblage that would be produced by direct i linear arrays, dorso-ventral flattening of A C, wlth elements arranged with their long axes transverse to the trunk axis, --::,--rsing anterior view D, the assemblage that would be produced by direct dorso-ventral flattening of C. t:::: r Fiss 6.8, 6.9, 6.10). r*.-: :ciial architecture, the unlikely to have been oriented at a high angle The alignment of the elements (constraints d : .r-:rsed with their long to bedding as this would have resulted in stack- and e above) suggeststhat they were all situ- I cr::.nsrerse to the trunk ing or close overlap of the elements. If the head ated more or less equidistant from the midline. t-J i of the former, in any and trunk are flattened in the same odentation There is no evidence of a horizontal radial l:: t:L-\ducea linear series to bedding (i.e. dorso-ventrally), then the arrangement. The position of the elements acrossthe midline. A F1-: apparatus in life cannot have been vertical with indicates that they were set in rows parallel to rL::.-rure fl attened normal respect to the trunk. the axis of the trunk (Figs 6.9, 6.10), although ;i::; nould produce an tr: rhe elements were E ::ad *'ere tilted, some -"* rest in opposition, but Rn0 -:-.r--: :he trunk axis. Hence, ll'l^A.'u P '41 V .E li u I fc----:pseand flatteningof a Elll ll tr z: = iish angle to bedding Erll frilt r tte \\ aukesha assemblage. rli: configuration of the lf !Ulldl il 1Jn rtu:. rhe axis of the speci- Fig. 6.9 Hypothetlcal lmea. architecture for the Panderodus apparatus, with the basesofthe elements in a : =res. indicates that the plane A, ventral or dorsal view. B, anterior view; elements illustrated in withdrawn position, to occlude they would be rotated together m the direction shown by the affows. The Waukesha assemblage is E: rhe architecture, is preserved in the occluded position. | &1,I aJuepr^e lseq eql asoql ol JeIIurs ,(11ercr;radns'ptluoporapued e pesoddo o,r1 pasudruoc sruotsoJlun'd lo 'uonePe$ 'oulpopg lo stuarnelapesnJ ,{lleretel Jo sJelsnlc snle-redde eql l"ql stseSSnspurru" Eqselned\ | { or Jeadds lou seop 1I peqrrcsep(6461) uel/iroNpue (L96I) seuftg eql ur eSelqruessE er{l ',{JeIIIurns uI s:(nale ]o eJuenbas aleld 'stueuralaEutsoddo eerql o1 dn ( sere; -re1norraql pai(eldsrp qurrt Jeqlo aql Jo 'JoIJel )pnuPJ e^\ qEnoqllv ;o sdq dsncaqt qlr,n reqlaEolslueruela pesodel esoql a qrrr 'lsouJaddn seJ?JJeuur rleql qll^{ Ertrurs are (6 pue 3 sluaur -xn[ ,{1pra1e1aarql sessessod(9961) qstedeg qull euo Jo sluJrlalJ eql Jo uolle^l3sard aql ut rela uuoJrJPJ raErel eql ,tq pequcsep snpo1(pto3 Jo relsnlf, 'Polres llnser pJnor qcre ue yo 3urua11egleJlua^-osrop ? ar€ nuauela urloJq€nJlEe -erd f,1-roodarour q8noql 'pe8uerft flJBIIuIs 1ceJr61uorlJaJrp Eurce; lualstsuoc e olul de P qnE \\ aql uI Jollelu€ eql y 'rorrelsod ro Jorralu?orll Jerllrele 'snpoop peqcre uB Jo sluaurela aql ?urrq o1 parmber ;q rsaireJ eqt pue lolJqr1Cd -u"d erlsa)lnel(\sq1 q1r,t ,{8o1euzfq 'luaruale aq oqe ,{eru 3ur11r11o aarEap e t€ql etou 'mlBfeoo€ |Frq eql ul rapro alelncrua8aql qllt 'eJnlJelrrpJ?fteuqe slno^pJ o1 luau4rad sr 11) snleredde reueld z Jo aJnxeg Ellrrlsar:oc slueuele aql lueuru8qeasolr ar{I (7961 e8puply) pua euo prlual-osrop fq pecnpord eq plnona lceJJe ttEql paleFlsod,(aq1 e3e1q te tuornele (ru:og11uopolsro)alelncruaE e q1,t rEltt.ursv Jaqlo aql uo esoql ueql laqtaSot eqt rr iq papr,rord eleP 'seuesur slueurole(ru-ropluopocelslp) alelsoc flesolc eroru leJ plno^\ qtft eqt Jo qrull I Dq s! (986I) lue8,ftq pue ,(1e.ra1e1xrs sasrrdnrocrprq 'snJrl D snpops auo uo slueurele eqt',{IJouelup pelll peeq eqa t:uatJ Jo acuanbes -sag, Jo sl aldurexe eleldrum lsolr eql eql qlh{ peuallPu eJe,t\'Je^e,t\oq paqtE "{?rl€ mzzrue3rostp rualrour-lsod ',{1e-rs1e1pesn; are snleredde-;pq e Jo slueu ue ;1 aueld Surppeq aql ot aAIlBIer 1eurue srql FFe uaaq sBq Jelsnp -ale eqt qcq^\ uI PeqHJsapueeq e^?q slusru eql ]o p?eq eqt Jo uoD?tuerJo aql IEII^\ rel lO pue {rzq q1t,nr.aerEe e,11 -ele luopouocne urJoFuosJo srelsnlJ [?Je^es -leur ou 'uJe €d srql ecnpord lou ppo,n {erre :ie lo acuanbes eq1 ratle reeuq '-reueld 'pec€ds-,{Fe^e uy Eurualleg !!lun aq plnolA lnq 'pel8ulur SUSISN'IJ Surrnp peaq eql Jo apnlql€ eqt 1ceger tsntu lEq o\r eql luo{ slueuela (ssn.{ nou'{ flSNuoIAs l,'9 uerurceds pqselned\ eql ut uou"Jn8lluoc eql !e r,-r,)s-roPueql a?elqurasse ueqt 'tu"tsuoJ ela.r Surcedseqt 'pueq raqlo eql urpord pynoc etnlcalrqJJe uo']I aJII ur tuelsuof, lou se,t Surceds eqt leql 5o iurualeg 1ere1e'I Iellxep 'srelsnlJ pasnJ alecrpur ,{eru lI 'ueuJrJedsal8urs e 3o srseq aql apls r41p aEelqurasseeqt Jo Jo fpnts eql uog (9961) 1ue3,ftq pue uo eJuepquoJ qlr,r pesn aq louuet aJnlPaJsrql 1o ecuenbes aql 'aJolu > zq fq pacnpep 1eq1,(lequesse st lueru 'JeAe^loH rorrelsod aql In esoql uer{l pepaorJ -a1e qa: pened Sutcur,tuoc e 8ur -e8ue-rrepaqore uV 'A\aIAIeJelBI ur spt€,ir{ eJorrJ er? JIeq JorJelue aql uI esoql eJuls irl-) uuoJl'ls] aql ,(po q1t,tr -u,{op Jo sprer*dn pexeg ro paqcre sfe:re 'a8elqurassp aqt ur stuerrrele aql Jo SuIJEds rr1 lno1 ,t'oqs sluaueJe eql aq1 pue peceds ,(para slueurele eql qturr (q) aqr ,(q parnporturar€ stut€rlsuor,"ylju""r" l6irq pue lrzc teql arnl Jo 'Jouetsod eql ueql pec?ds I pnoqrle 'sluatual3 Jo s,t\ol ,{[JsolJarolu sluauele :or.r:1ue aqt qlrm (e) Jo r€lJuaeq e^"q {eur sa\orluatuelo aui.oioo rrr+L\a pa:rp f,ue 8utrca,ra: ::aq1res,(erre :eauq eqJ pelIIJ uaeq a^eq plnoJ lnq 'lpluozuoq 'ipoergru8ts sreJJrp lelsnlr se peSuerrepue 'f,1ere1e1qSuperado'se,rpq ueeq eABqtou deru snle:edde aq1 ;o eueld aql r :ules eqt aJ€J 'eldur?xe lc _rosaJeJ leral€l Jelno eqJ 'uorsnlJaoJo uollcellp moqss^\orrE Fe-\\ eql ut Pa,rresqo sdrqs 'A\er^rouelue 'B r(llellue^ro,{ €srop reqlre pus sple,{rurpelcerp sdp lueurole:,{er^ Ierelel 'V qcl€ ue uo slueurelaaqi ses?qaql qll^r snpo,lvpuDdeqt rc! atntrelrqcrplEaurl 3tg s .$oqs slue(Ilale eql pue Jo 'snlEledde letDeqtod,{g 61 9 '!a\e,$orl 'sr (9961) tue8d:q g v ry euolsoJlun d Jo JalsnlJ I uuuad o1 slueruala /r\e; ool '1696I) IJollod ,(q PeProcar I to slelsnls lsrrJ eql 'Ima3] l'r6-6I) uel,top f,q PaPnlc rala eqr ;o Suuapro luereJJIP qr lo qJ€e 'ra^ea\oH 'aloqB [8 9 'cas g qJl ysn 'ulsuoJsld\ Jo uBlrnIIS Jaraoloql |IIo{ Ieu4uB tuopouor V Edn. LSA [ch.6 Sec.6.8l Basalsupport in coniform euconodontelements 101 above.However, each of their clustersshows a ments in coniform apparatusesis provided by different orderingof the elements,and, ascon- the clustersof Besselodusand Panderodus.The cluded by Nowlan (1979), they are probably half-apparatusconstituting the Sesr€lod.us clus- faecal. ter compdsesseven elements (Aldridge 1982). The first clusters of Pand.erodus were In the PodotianPanderodus, Dzik andDrygant recordedby Pollock (1969), but thesecontain (1986) recorded six paired elements and a too few elementsto permit interpretation.The seventh, the tortiform ('hi'), for which the clusterof P. unicostqtusdescribed by Dzik and opposing member is missing. It is therefore B Drygant(1986) is, however,almost complete, likely that both these genera possessed lrfu r-L=sof the elementson and the elementsshow some of the relation- apparatusescomprising seven pairs of ele- r rE:-:l) - B- anterior vrew, ships observedin the Waukeshaassemblage. ments. There is nothing in the Waukesha The outer lateral facesof all the elements,for assernblageto refute this conclusion. example, face the same way. The Podolian Ilaierallr'. and arranged as cluster differs significantly.however. in not r revealingany direct evidencefor two opposed dor elements more closely rows of elements,although this is the architec- 6.8 BASAL SUPPORTIN CONIFORM h :r:r:rerior, or ture that Dzik and Drygant (1986) derived. EUCONODONT ELEMENTS :urs erenly spaced and the The elementsshow four principal orientations, Severalauthors have suggestedthat conodont c iexed upwards or down- with only the falciform ('ne') elementsretain- elementswere mountedon a supportingstruc- I ll€E. An arched arrange- ing a convincingpaired relationship.Further- ture (e.g. Kirk L929, Stewartand Sweet1956, irlr t-hat deduced by Dzik more, the sequenceof elementson the sinistral Schwab 1965, Lindstrom and Ziegler 7977, (19S5) from the study of side of the assemblagediffers from that on the Jeppsson 1979). A cartilaginous framework dextral.Lateral flattening of a linear or arched was envisaged by Lindstrdm and Ziegler architecture could produce a more irregular (1971), who suggestedthat the shrunken assemblagethan dorso-ventralcollapse, with appearanceof some basalfillings may be due EO\I FTiSED elementsfrom the two half-apparatusesinter- to the post-mortem contraction of a weakly mingled, but would be unlikely to significantly mineralized cartilage-like tissue. Jeppsson alter the sequenceof elementson either side. (1979) comparedthe grossmorphology of the f a-oirbrm euconodontele- We agreewith Dzik and Drygant (1986) that crown of severalconodont elementswith the kritred in which the ele- this cluster has been affected by a degree of shapesof fish teeth, and pointed out that pike are fused laterally. f,rr:.3ms post-mortemdisorganization. teeth attachedto a basalsupport possess flared tte erample is of Bes- The sequenceof elementsdeduced by Dzik bases,similar to thosedisplayed by many coni- *l-h comprisessix laterally and Drygant (1986) is not in agreementwith form conodont elements, including those of :ii.-'rm) elementsin series, the data provided by the Waukesha assem- Panfurodus. Unsupported teeth, in contrast, (c*rtrdontif orm) element at blage.They postulatedthat the sizegradient of haveparallel-sided bases. If conodontelements B1v!: ). The closealignment the elements corresponded to the original were supported,the contactbetween mineral- fir*;rure. with thegeniculate order in the apparatus,with the smallestat the ized basal filling and unmineralized support E rrth the WaukeshaPan- posteriorand the largest,falciform, elementsat may have been transitional,and the degreeof G alterior or posterior. A the anterior. In the Waukeshaspecimen large mineralizationmay have varied betweentaxa. I tbough more poorly pre- ?arcuatiformelements are at the anterior, but Jeppsson (1979) extended this model to I Cordylodus described by the larger falciform elements (Fig. 6.6, ele- ramiform and pectiniform elernents,conclud- Itxsess€sthree laterally jux- ments 8 and 9) are situatedtowards the pos- inB that their morphologicalstyles are also in tsetier with the cusptips of terior. Although we cannotdetermine the com- better agreementwith attachmentto a skeletal L elements. plete sequenceof elementsin the assemblage, framework rather than directly to muscles.The \o*1an (1979)described it does not appear to be one of simple size apparatusof lcriod.us,which containsnumer- lfrsed elementsof Belodina, - gradation. ous small coniform elements (Nicoll 1982), rficially similar to those 4re The best evidencefor the number of ele- lends some support to this contention. Nicoll D'!-rd ur) V H'Suolsrurv -uz7 ur pesruSocal sesnleJedd€-JleqSursoddo -rAopJOul purrou sr ueql raleerSqJnur sr aZIs 'std €€ eql su?f,"4soreleq 1o sa1eld I€to aql ro spD eql lnq 'luaurelepnpr^ryu u? eq ot peJaprsuoJ led :-iu.rJs (lce4sqe qsltSug -uoz,{urorled ;o JSrp Flo aql a{rl selnlJnJls ueaq (1eleueB setTtlun srupouSoaUrtv rf)z1 a.t:., ruaJo[pDaqj puo outqJ Jo eJueprle ou eprlord saSelqurasseluopouocna '(Of '39'tgOt urorts?ragpu€ raddepl) snorod riL : s6l p p 'Elde:'/.reJ-':Jv 'S, €9 'xassns rrrroJruoJ (Ig6I Jar^uef) ernl€lnJsnrx arou sr lnq 'enssq auos pseq pcrd,$ 1o 1eq1 11 ..na sruopouo)Io t8o eyqeredruoc dq pale-redo ueaq aleq osp,{eru 01 JsIrIIIrSSr eJnpnrlsoJcru eqJ Jorrelue eql $j: a 1)J:uI sesnterpdde suer"rlsorataq Jo seleld pto eql (996I te sseJoJda{ll-8uu$ I?Jel€lp qll\{ r?q paaoq € E- i.{ snoteJruoqnJ iIg6I Jo ueplp1 rer^u€f) spquozfuo4ed pup Jo ur.roJJql ursl ll sJlJlluapJql sBlqSrJLl JruBS !t-= :[l t86I C U',{qroN (I86I rar^uef'€961 uos,neq) sprouu^u aqt ur aql pue 8uo[ Ixrll sr,{poq pseq I -d l|qrlws' f u'eSPupIv lnoq" Jo I lnoqe E-1'T pueltocs'qSrnqurpa lueserd aJe seJnlcnJls A\e[ 'pezll?taumn lnq slql '(t86I urorls8rag puD raddel)) dpoq + =:-: eJuapr^3.A\eu-sluop 'elll?redo r(gera1e1q r"lrurrs '(SgOf lEs?qlelluBlsqns e puEensstl u,trojc;o sdtrls ,{q { .:i6t d Itr'qlltus pu€ '1o p e?prtply) sel€ruerr errrotsoqleuS-erd pue pelJeuuoJselcrluep IIlroJIuoJ Pe^\oq Jo saues -D f C s;8ug't U'e8puptv sluopouoc uaa^{laq drqsuorlelar palelnlsod aqt € s? padola^epsr u^{orJ aql snqwuSoaq)iv uI {3o1o1uoao7o4 :;-! it SZ qll/{r sr snleredde snponpuod '3urqr; .::--:e Yr 'pue.l uolSurqser^d luelsrsuoJ etlt pseq cqqnreqdse pu€ euoJ Ipseqpal€ { =orl stuSlueleluoPouoa JOJpalsnJlsuoJeJ e^?q err arnpalrqJr? eql -uruel 3 Jo pasoduroJserpoq psPq pedeqs-A rF=\-- \ 'a86I I U 'eSpuplv 'JJrl ur peuelleU ,(ll€relul Jre,$ a8ftl uo pelunour er? su,!\oJc ruloJrurl gJrq^\ su€eJ€qleuE[1od eqt uJo{ uor]Jes-ssorJ qclqm ur snpoaloJ sueurbeds peJn ssSNsus.{su -ced Jo ur pejeJJrp a^eq ,(BrrrsnpoJapurJd |eql sal?Jrpur -?tl QSOy) 'p p pue (9161) oqualel ^o1lsreg tr aJII ur Surueueg Jo eueld e slcagar Eurppeq -so14l'eEpuuopoaloJ eql Jo sa^nElueseJdJJul ol uonetuarJo srql JI 'loeds€ IeJluel-osJop 1zseqer{l Jo luerudole,repelrsuolxo aqt sr 'uus v rl^ [q Pepr^ ur pauaDEUsr ueurceds eql lpr{l selucrpur e8eyq ^poquorluelle JeqlJnJJo ,(r{lJox'aJnl"eJ Jgr{louv rr :i'idBriotoqd pue 'uosuDl -uress€ ?qselnp1N eql Jo fulerutu,{s pellesald ' 0 SAt Elnt4) sqru€rqous"le !E--: :re\ sSuh\€lp aurTeql eq1 'sueeceqleu8flod uotupJC aql se a1e3uo1e pue srurepoceldur pue (6961 peetspH '/96I JE-: trlpuapuod uo lue8,{tc s€ lou sE^\ snponpuDd l€qt lsaE8ns Fpoq eql uosrueq) sueo€JlsoJeleqlseru€e eql ur punoJ E= iq reded aqt Jo fdoc Jo suorsueurrp aq1 1nq ',{8o1oqd:our enssq-Uos sr 'a8elqrec perlclec -re1nqo13'eJuercedde b -r.'rd JoJ 'a\esJpl!! 'secue prlrplep ]o JJuJpr^r Ieuorlrppe elllrl sapr^o:d .lelrursJo enssrlV Jtunlo^srgt jo 7 ratdutl.; -f=c taet rc r{u3q1 osF uaurJads pqsa{n?d\ eql uIsl"dUos aqlJo uorlB ul (€'Z Id '186I) r{s,r?ruezs [.q snpoytptoS €qo:qqzg eqt uo4 lerra]pru -,r.rese-rdrood eqJ seleru"JJ ssel,tef 1o dno:3 ur pJle.llsnllrJr? sJ.lnlJnJts r?lrrrJrg '(7961 p ia ,seuu"r{teJ d '-.uBtuo lS € s€ €luopouoJ aqt pepreSe.r (986D p D ,rolsreg) snpoap) pue (.EL6I 'p n seu:l-g,) I -..truu3){ t d r(I puP eSprrply 'uolueJo (xoJJ sleurup luopouoJ snporapuod Surpnlcur'ex€l uenr^oplo sltr:$alJ luopouoJ pel"Iosr snoJqruoqiBJ Jrll ur paAJJsald ssrnlJnlls IeJaAasJo s8urllq leseq eqt ur paproJeJueaq pJ-iJns'ie^rns leJrSoloeC snle.redde pue enssrt-Uos eql Jo srseq aql uO elerl seJnlonrlscrllnraqdg pertnber sr qro,ta DU -{ uosrPBI -UtSUOsSrd\ reqrn; qcnu q8noqlle 'uoddns pseq Jo eJue IIINTd.{Y UOJ SNOIIVJI'IdWI 6'9 " r-\:U\ iSoloeo'pqd1se16 -lsrxeaql Jo ecuepr,reepr,rord,{eru sluopouoJna I't. EJrurJadseqse{n"A\ etLL f,1rea;o salnqr.41epcrSololsrq euos 'slueruele :E :--r \pE roJ InJelerS ar? a^\ pJzuelnJsnul?npr^rpur ueJlJnoJ Jo uoDerado i le:ri-oloec srouu 'crTnlrtrAl 'srugou8oaqcty ur se Jr.ll plno^\ uErll uorsnlJJoolur sasnleredde ?\-t-1 elorq Eqse{nert\ eql uo ,ror e ul pa8ue.r-resllun uJoJluot elaJtslp rrJorJ -]lpq o^r1aql SurSurrqroJ rusrueqcauluercgJe r :-:relder{r sIqI S0IS/EUC paJnpord eq p1noc snponpuod pue snpopssag arou e apr,rordppo,n goddns pseq'snpuap f,q pecueurl ,{q u,noqs ad,Q eq1 srelsnlc 1zq1 ,{yelrtun -u"d esoql w qJnssluelxele uroJruoJ roc I f'U :I N ;o Io , | :uopouoc uo {roa\ JnO tl tleJ lnq ',{ll[Iqrssod sql peraprsuoc (ZL6 d '1JOO '7961) ruons8reg pue raddely '1uerue1e snon8nuoc ruroJ aruos sNI,\isc(g.IA\oN){cv -dns 1es€q Jo lnoqlrll pnprlrpur ue o1 lualerrnbe sr .alJrluap, qf,€a uorlsunJplnor ,{3q1,roq aBesnuao1 llnJrJJrp eq leqt Pu€ 'perrnJf,o snleredde-;pq e 1o ernpnJls plnoa lr 'esEJeql srsrql ;1 sasnlereddeJer{lo ur 8urlroddns eql Jo uorlezrl€reuru alelduot luerlrelarrrJoJru€r e ;o ssaco.rdeql uo elorluap rruJe 1en3ur1aqt ol ocuelq qolqA{ ur eyduexe arer e sluasatdat snq|ou a13urse q1r,r snoSoyouroqeq tq8rru slueuela 4 tnpolassag pue snpotap -SoaqctV l€ql alqrssod sr lJ sluopouoo uetJ Iecruoc esaql Jo qcee leql pe1se88ns(7961) e'qcl VSn 'u!suoJs!l\ Jo UBIJnIS Jaaol eql lxo{ IBrxluE luopouor V z0t LSA [ch.6 References 103 -n. I s :nssible that Archeog- derodus and Besselodusbear most resem- early Siluriancarbonate platform of North Green- i a ::re example in which blance to the lingual apparatusof myxinoids. la(rd. GrOnlands Geologiske Undersqgelse B ulle- Ea:.-n of the supporting tin. Barnes. C. R. 1967. A ouestionablenatural cono- rl]ca-arus occurred, and that dont assemblagefrom Middle Ordovician lime- Gqrr 3lent to an individual ACKNOWLEDGEMENTS stone,Ottawa, Canada. Journal o f Paleontolo gy 4l r Bergstrdm (1984, 1557-1560. -J I r-. possibility, but felt it Our work on conodont palaeobiology is Barnes,C. R. 1977. Ordovicianconodonts from the tr-: .ri rhe type shown by financed by N.E.R.C. Research Grant Ship Point and Bad Cache Rapids Formations, Melville Peninsula, southeastern Dist ct of could be produced GR3/5105. This chapteris related to a project Fra;rklin. Geological Survey of Canada Bulletin h=--:,lus ::nits arranged in a row on the Waukeshabiota coordinatedby Dr Don 269 99-719. Mikulic, Illinois Geological Survey, to whom Barnes,C. R., Kennedy,D. J., McCracken,A. D., -- we are grateful for adviceand encouragement. Nowlan, G. S. and Tarrant, G. A. 1979. The The Waukeshaspecimen was loaned by Dr K. structure and evolution of Ordovician conodont apparatrses.Lethaia 12 125 1,51. Westphal, Geology Museum, University of !L FOR .{FFINITY Barnes,C. R., Sass,D. 8., andPoplawski, M. L. S. Wisconsin-Madison. Dr Rod Norby, Illinois 1973. Conodont ultrastructure:The family Pan- : r-:i-tissue and apparatus Geological Survey, supplied the collection of derodontidae.Life SciencesContributions Royal :ri r the Carboniferous isolated conodont elements from Waukesha, Ontario Museum 90 36 pp. t A.-:r Granton, Aldridge and Dr D. J. Kennedy, Brock University, Barrick, J. E. 1977.Multielement simple-cone cono- *: ihe Conodonta as a St Catharines, Ontario, provided topotype donts ftom the Clarita Formation (Silurian), Arbuckle Mountains, Oklahoma. Geologica et re-:res. The poor preserv- material from the Bainbridge of Missouri. We Palae ontolo pic a ll 47-68. t-:re \\'aukeshaspecimen also thank Dr Jerzy Dzik, Academy of Sci- Barskov,I.S.,Moskalenko, T. A., andStarostina, L. lcq=] evidence of detailed ences,Warsaw, tor providing a prepublication P. 1982.New evidencefor the vertebratelature b:r- :ur rhe dimensionsof copy of the paper by himself and Dr D. M. of the conodontophorids. Paleontologicheskii b: P;nderodus was not as Drygznt on Panderodasclusters from Podolia. Zhurnal, 1982(1), 82-9O. Bergstrdm, S. M. and Sweet, W. C. 1966. Cono- Doh,gnathaceans.The The line drawingswere drafted by Mrs J. Wil- donts ftom the Lexington Limestone (Middle n-..:i .-- lhe Waukesha assem- kinson, and photographicassistance was pro- Ordovician) of Kentucky and its Lateral equiv- trfu .recimen is flattened in vided by Mr A. Swift. alentsin Ohio and Ir'diar'a.Bulletin of American E- Ir rhis orientation to Paleontology50 2'11 441- Branson,E. of flattening in life it B. and Mehl, M. G. 1933.Conodonts llE:< from the Bainbddge (Silu an) of Missouri. Uni- bE";:d- mav have differed in REFERENCES versityof Missouri Stuclies8 39-52. * r-rlssnathaceans which Briggs,D. E. G., Clarkson,E. N. K., andAld dge, *r n life. Aldridge,R. J.7982. A fusedcluster of coniform R. J. 1983. The conodont a ma Lethaia 16 lI. ia\ e reconstructed for conodont elementsfrom the late Ordovician of t-1.4. Washington Land, westem North Greenland. is consistent with Cooper,B. J. 1975.Multielement conodonts ftom tiE-=rus Palaeontology25 425-430. the Brassfield Limestone (Silurian) of southern between conodonts Aldridge,R. J.,Briggs, D. E. c., Clarkson,E. N. K., Ohio. Journal of Paleontology49 984-1008. -nripE .-:aniates (Aldridge et al. and Smith, M. P. 1986. The affinities of cono- Cooper, B. J. 1976. Multielement conodontsfrom :-larerally operative, donts-new evidencefrom the Carboniferousof the St Clair Limestone (Silurian) of southern ,FE :tructures are present Edinburgh, Scotland.Lethaia 19 279-291. Illrn.ors.Jownal of Paleontology 50 205 21,'7. Aldridge,R. J., Smith,M. P., Briggs,D. E. hrr.a 1963, lunvier 1981) G., and Cooper, B. J. 1981. Early Ordovician conodonts Norby, R. D. 1987.The architectureand function from the Horn Valley Siltstone,Central Australia. , t Janrier 1981, Yalden of Carboniferous polygnathacean conodont Palaeontolo gy 24 147-1 83. p]-eles of heterostracans apparatuses.In: R. J. Aldridge, (ed.): Palaeobiol- Dawson,J. A. 1963,The oral cavity,the'jaws'and !r LlrErated by comparable ogy of Conodonts. Ellis Horwood, Chichester, the homy teeth ofMyxine glutinosa.In: A, Brodal rtr 1981). Coniform Sussex,63-75. and R. Fange (eds): The biology of }|{yxirle An, Taixiang, et al. 1983. The conodontsof north Universitetsforlaget. Iese-: provide no evidence Oslo, 231 255. China and the adjacentregions (i\ Chinese,with Denison, R. H. 1967. Ordovician vertebratesfrom b !-=l disc of petromyzon- Englishabstract), Scielce Press of China,223 pp., westem United States. Fieldiana, Geology 16 of heterostracans. The 33 ols. 731-1,92. -esran-r=s recognised,in Pan- Armstrong, H. A. (in press).Conodonts from the Dzik, J. and Drygant, D. 1986. The apparatusof tFEtI] Jru€8Jo uD uo JncJo fmuorq lo sPuDlllY slwld 'rnlf,lPJ pu"'spuoqdoqlrToc Et 'sisnllour ur suaqs x u! euoq apnlJur saldu€xg ln DnpoJd eql sr 'suelslt {:lfiu€ Suoure punoJ sr l€ql Ft! anssrl prsq ,{uv uror; Jo PUr{ '00€-16Z tS f,laDosuoau l7 aq to lournol 'urseg 8uruueJ eql Jo ueluo^ec reddn eqt pue nostJerg snsu'dxa snpouol I'1, pctgo1oo7 ,{l{qdouou etuolsopdc .loJ eJuep lqew luopovo) -\olrlooouJNr -r^e sE srrjsrueqceujSuIpeaC 986I id C 'ueplel aq11ouoqrsodruoc luaualerllnl^tr Z86I S lI'lloclN '/t-9€'xessns'relseqrlqJ ,tz-ztz 6zz 'poo,A\.roHsrlll 'stuopouo) lo t8qolqoaolo{ USSS Zr?rN ttutapoly Kpolqoq releeqrtqduv :(pe) e8puplrT :uJ stueuela luopouocne {slnIJI eql yo slrsodepuel.r^opro aql uI spurr &nor iq pel?uurop i( etol f N pue luopouocered 'luopouocolotd Jo suosued puoqdotuopouorenblun 916I V I'o{uelelsol l 's8-8/ Fiau(norq u^\ou) uorJ lue -rrjoJ Iprnpnrls ,{Jpuluillerd /86I H 'DIs^{eIuBzS II€ g sauasuopuo1 . Dq SILL %EZu?ql sselalnl 69-1y 6 nc8olauoaDpdn o4Sopag' t t 6I Io &aoos lo{ov aqtlo suotltosuotl lDltqdosollqd ::cr :.noq8norqlrncco ue8el lqew pue losvetfl snpolnotsnueb eql Jo sluop 'VSn 'ulsuoJslr{dJo ueInFS Jer'ro-I eql ulo{ pue perrese:d dlpuoqdecxe rreu q986I suqg salpung euluml -ouo3 S/6I d H 'qnEluoqcs S lA ']ee,,ils elorq V f Jo 's8-91 'Jropuessenlxpue g q 's38trg' 'c[n{I ptraleu E reo 9z 'g 5 C tr degor alqrssod) salpryS[.3opa5 [.t1sta^1unSuno1 uD13t]g eoul.l' 'LIL_gIL -:ltal rl padeqs ,{1snotre,r. -ord tueurtuoJpql urelse^\eql Jo .{qderSqe.qsorq 8ZZaruat)s uTotqperPoq-uosuPunIS v P986I I uoueururexg srusrueSto pup sluopouoJuerJl^opro etP'I 6t6I C i['lee^{S 'J.ropuessanD{pue '9 g g 's33ug'9 (I 'clln{lw tlln 'NEI_19 Itb\an qr 1peul"lqo suollses 'ELZ-r9Z 6ZE pue opouol f,aatngpnSo\oag tuPpunoq ar? sllnseJ aql sulEls OEtSoloiuoalod!o lrr.rflof orqo IeJtuec-lse^\ uuallng lo Ern speq euoq u€Iuo^ec elppll^l eql luo4 Jruelsds eqt pue ,{qder8rlerlsouoJqJueunls pue ilrsuatu aqt ol pre8er Ie4UeCJo I sluopouoJ gsbl ) ,4^'rae,{\sPue'v I'ue^\r}s {pee-uerct,rop.tgelel ol JJUoJaJJIlelJeds peul"ls eJeA\ rf?ttleeq qll/t\ 's1d77 'dd V9g 'uto43ut11ol11lo t11aaa qll^\ 'reqeno 'puelsl rlsoclluv 'uoDeu oC deg t te pauortJos pu" 'uIJJEJBd -?un '|slsal\Oqd ptqs?nndun pueluearo quoN slllg eql Jo pu€ ,{qder8rterlsorqluop urelseepue lseg 1o ,{Soyotuoaepdluopouoc (uztc -ouoJ 186I^SolocaoeFd )I J'seuleg pue (I V'uelcerJJl^l +-iqap'paxrJ'pezll?reutlxep 'tt-b g DJrSolouoaDI,d rrpog 'sEo9 ruor1 -r^opro) uerlcoratqd\-uplxeql S86I d htr'q_UIuS IqeW '€65-06S /tto p oJlSopag pacBluopoiepued elllutEJ.reqn onapoapurg, serceds 6€ 4,tu -piuoal'd Io loutno[ sluopotJoJUPIcI^oplO elp erg l ualuoposoJ up ue8unqtnsrelun e[elnl rrrJoFuDf,ed 'I^l E:l: luopouoc -prlr{Jruos Jo JrnlJnJlsorJrl^lsq6l A\ ')I 'qe^lqJs -{nrtsuraC IL6I M'r.Ferz puE 'rlIolspur-I '602-L0z 'gL6-6n6 SSf,8o1uuoa1o1 lo loutnoJ JJVUJStrV pue 9EJlDtsuosapung uaqcnSoloag np ua8unpuoqqv 'selerqeue^ pue sluopouo. ol suonPlel sll 'sel€ls pelrun urelse^{ eql ujo4 srelsnlc luop snqJquSoaqJrvIssoJ uer.r^oplo eJppIIl clletu -ouoc pesnJueDl^opro,{lreg 086I g f 'qslede)I -8rueaq1 7961 I^ S'Iuojls8ragpue g'reddepl 'st6-626 '96n-E6n pj ,(3o1oluoa1o7 euerpul tuor] s.re] BI g sauas 'nuaos lo loutnof uoJr.tatuv atold '!J lo loutnol puneJ rq-el f I puE sneqPc 'I -snlctuopouor ueunlls pesnd 696I v J'IJollod ,{reurruqerde-opuoloJ Jo qsl}uelcr^oplo 'nge-IZE Z t3o1oo7rcJ anlty 1€reua8uI sel€.rq qlr/\1 pJlPrJossesluopouoJ 626-l u s 'lrl) ) DilapoapulH -eue^ re^\ol Io senssllpleq eql uo SII€tueJqll^\ '99 gl ototts puo sl?ssoJslqtsnoql e^qeto^ Io 'uotele{sopu3 aql :I SqCuelqotusPleIssoJ puP -ord euos :selpntseuoc elduls €86I -l'uossddet 'ILI-egl a'I surepoJ€ld Jo sepnls JISololsIH 'lS6I L'8].yo ZI otDl enssll 'uolpuru Flelu uos '6n-I 6Ze uyalryg DpDuoJJo lueuela luopouoJ 6/6I 'I 'uossddef f,aung pc8oloag ceqeno rpueFl qsoctluv 'uoll 'tzI-IDI n u!o41a7swJo! -eturoc l€erne^ aql tuo{ sluopouoJuelar^oplo .regtursur pue eddl z1apoap tHlas.sntereddeluoP '1 et€'I 186I U 'C 'seureg pue S C 'u€l1doN -ouoc ug tuetus8uerrelueulelg 1161 'uossddal 'glz-trz vl-61 '69r-rzr odoL opouoSlo {a^tns pttsopaD 'ulPFPrC Jo 1 l3o1otuoa\o4atotqauaAlo lounol ,sueqleu8e, lJulsrc 'puplsl ereursellg 'uoneujlod ulelunol^J ltsso]Jo e.ueJlJludtseql ol ecuereJelPIJeos ql!Y' q.uttqj ruory uot8urqtg outpo1ag snua? 'eleruerJ eql 1o {ue8o1,(qdeq.1, 186I 'd 'rel^uef "q luopouoceql Jo srelsnlcpesnC 6L6I S C 'uet,t\oN |ZI-L\IE TJtoasavanss\ paUDlD) seTetqele^ .EIZ-L6I L srtstqdoa{)puo [.gopaC tserlEe eql uI enssBpel]IcteJ 696I 'B "I'pee]sl"H 'sluoPouoaprluopor?pueq uDtloqsnvIo lournot AWg e{Pl1snv ulalse r11I-t€,I 6I DtDqDT 9'qJl YSn 'ulsuorslll Jo uElrnfls ra,|Aolaql urorJ |Bu4tr? luopouoJ Y n0r ETin. LSA lch.6 i itlR Journal of Austrulian le('::: j 7 197-213. l- F--f clusters of the conodont E::-::ron from the Thumb :llesmere Island, District --.br-,:- Srrr-ey of Canada Paper d 3.:-res. C. R. 1981. Late &rr:. ::.rm the Vaureal Forma- fu,- .}rebec. Geological Survey . -r:t : -{9. il:.:.=::; studies of placoderms Soft tissuematrix of decalcifiedpectiniform elements ici::l:. Ir The endoskeleton, tE :=:l lissues of lower verte- of Hindeodella confluens (Conodonta, Silurian) -.r-'..,. ''.'r Zoologi 2 32I 354. l- : -=: Silurian conodont clus- L. E. Fahrcus and G. E. Fihraus-van Ree .- ; -.,1r[ of Paleontolog! 43 L 1- , Ordovlcian fused cono- r -:: qestern United States. r C-:: a.::tchen Bund.esanstall35 ABSTRACT cellular componentof the tissue;that is, it is 5- \1:-:rirructure of some Mid- Pectiniform conodontelements of the Silurian producedand extrudedby the cells (for exam- Journal of Paleontol- ples, see Nancollas 1982). Although there is -;':j:.:'-> speciesHindeodella confluens Branson and [rer..:-\\'hirerockian (Ordovi- Mehl from Gogs, Gotland, Sweden, were controversyabout the actualinitiation and pro- fre :-:..log\ of East and eastern demineralized,fixed, dehydrated,embedded in cessof mineralization,it appearsto be gener- | :- t- PhD thesis, Uni- ally agreedthat cellsmight inducethe mineral- -:)ished paraffin, and sectionedat 6 pm. The sections L!- --!r pp.. 22 pls. were stainedwith haemalumand eosin. With ization processbut are not themselvescalcified lS-*==:-\\'. C 1956. Conodonts (Nancollas1982, Westbroek and De Jong Er :::an bone beds regard to the intensity and localizationof the of central 1983). However, in this contribution we pre- frn: ,': irnal of Paleontology 30 stains the results are indistinguishablefrom sectionsobtained with tissuefrom freshlykilled liminarily report on the first experimentinvolv- L;:= l:oolician conodonts and organisms.Examination of the sectionsreveals ing decalcificationof conodont elements and ::rrern f r:e Midcontinent Pro- variously shaped cells, bundles of fibrous subsequenthistological staining of the freed tr: tt ersity Geology Studies - material (possible collagen), collagen, and soft tissue.This experimenthas shown that the mineralizedtissue of conodontsuneouivocallv lc,:..-=r-:b. H. P. 1975. Cono- lumina. Bundles of fibrous material and col- j .girJ includescells. r -' Branson and Mehl, lagenoccur throughoutthe sections,but consti- I a::.o4tologica 9 41-59. tute lessthan 25olo.This is considerablydiffer- li structural com- ent from known biomineralizedtissue which is rc:L1r.iont.-:-rminary paraconodont and :r= in: R. J. Ald dge (ed.): totally dominatedby collagen. 7.2 MATERIAL AND METHODS f C:,::ionts. Ellis Horwood, j--:-. r- -: Pectiniform Pa elementsof Hindeodella con- F- mechanisms as evi- (referrcd by many authots to Ozar- 7.I INTRODUCTION fluens -:ilingmonophyly. Zoological kodina, see Fihraus 1969. pl. I. fig. I for -\=,- .r.ociety84 291-300. -:. Any kind of hard tissueincorporating minerals illustration of a typical specimen)collected at that is found amonganimals, plants, or protoc- Gogs,Gotland, Sweden,were decalcifiedindi- tistans, is the product of biomineralization. vidually in a commercial solution 'Cal-Ex II' Examplesinclude bone in vertebrates,external (Fisher Scientific Company, Fair Lawn, New shells in molluscs, coccoliths rn coc- Jersey), which includes a decalcifer (trich- colithophorids,and calciumoxalate deposits in loroaceticacid), a fixing agent(formaldehyde), plants.All kinds of biomineralizationappear to a mordant, and a surfuctant.Initial attemptsat occur on an organic matrix that is an extra- using decalcifier and fixing agent separately 'o€sx uorlerrJru8el^l3uurcls ursoepuP unpureEH (^lorrE 33je[) uaSello.pu€ (.&oIIpllErus) IEuelEur (ue8€Ioc a) snorqgSul,,rtoqs xr!?tu enss!-tlostuauele uroluqced Jo lrElaO-' 3rg '0€9x uoqerqruSen Suluretsursoe pue tunleulepH sller e3.relpue (^{orr?IFus) tueruSrdlq paulltnol€upc p 8ur^\oqsxrrterx enssr}-tJos tueurele urroJrurped Jo IrEteC-t 8rd '0€9x uoll?crJruSel^lSurur€ts ursoa puE txntErxaEH(,{oxE SuoDlplrelEul snorqrJ pu€ '(^1ore uedo) Il$ e8rel'(^{ox? eBrEI)IEurr paqruerq '(,,,ro:-rzlerus) Suruq1ec tEg qtr^ IEuece 8ur^\oqsxrrleur enssrt-lJos lueuele uxoJrupredJo lreleo-Z 3rg '0€lx uoppcrJIuBEIl SururEtsulsoa pue tunlelrlepH pultunl pup 'eruetsqnsElnllec-lrlxa 'sller ]o spurl luelaJllp Suuloqs xglErn anssq-uostueruale ur.roJlurlred Jo uotces Paluorrouo-l 3u I / eleld ereql aslueururPluoJ qlr,{r SulFep e1l\ aJe -srp,(lasredsreqlEJ sr qcrq^\'leuelEuJ snoJqrJ :sr sql aII[ ,(pn1se ur uonsanb snorAqo eql aql ralJnu 11eursrire,r qlr.{\ sllec pe1z8uo1a '(896I'p ta J€uzield)spIJB oulu? elqepelep ,{q psuq ,tpuenbar; ere 'peqcuerq eq o1 reedde ureler etr l ort s3 plo s3 sluopouoJ lslll u.toqs seurrleuos qJrql\ 'slpueJ Suol aql IJoelJnu a^Bq sarpnls rell]Bg urPueJ sprJe ourlrJs punoJ ot t€U dq pepunouns ele qclq^{ relcnu Jo rrIroJ eql ur spnpord uouep?J?ep Jlaql punor PuorsesJo qllt' ruseldotf,c peuElsun ,{Iuo pu? 'qleap uodn eulu uoqs p ulqtl^\ at€Ja ,{qecrlcerd o1 paurels-f,11ear'req o1 s:uadde -alursrp'uollBzrlersurr.uorqjo slJnpojd JJqlo leqn ,{q peu{tno ,{l1zqred seurlletuos er" pup auoq ur xrJlerrJeql EuluJoJ 'slrJqrJueSPlloJ euruny padeqs ,{pzln8ar:r eqa tuaur8rd ryep pue seJnlJnJls IIer l3rll pel"Slnurord ,{lIEJeueS ,{q peulllno eJe eurrnl eseql 1o auog'ruseld ueaq seq lI 'sulels IsJrSolo[q qlra\ lsPeJ ol -o14ccrlrqdoursoa ,{ll€a^\ pue relcnu pele8uole ,{11pqedecs1r o1 p-re8erq1t,ll 1ce1ut,(lleJlureqJol qlrA\ sllal ,{q peuq flqloours ere Eurunl punor -srq uBurer ueJ plo €I l00t anssll leql lceJ aqJ sIEuEr 3uo1 se paluase:der ro 'pedeqs aq1 sr ,(pnls srql Jo llnseJ elqEIJ"IrraJ lsotu eql ,{1-reln8e-u'punoJ Jeqtre ere (urr/ g7-g ruor; uouces-ssorJ paleurlse ur Surflel) eulunl NOISSNJSI( CNV NOIIYJTUdUSJNI''I, aql surlllnl puE 'lBrreleur snoJqg'sllal Jo lsrs -uoc suoDres aqt adocsorcnu 1q31 eqt .Iepun 'srusrue8ro pa111 ,{qse4 uorJ anssrl qlIA '(I |td.) paurelqo suoDJesuory aJqeqstn8u ttsrpu I arz ruseydol,{ccrlqdoursoe ou Jo ssal Pue lloalJnu suopces eq11o Surur?lsaql Jo slpser eql 'surels .r:3.re1:qt fueu a,req oslu sllal eseql sllal oqt Jo uorlezrl"Jol puu ,{1rsua1uraql ol pJeAaJ Jo ralJnu aqt u€qt rell?rxs ,{llJurtslp al? qclq^\ qlld|lueuala luopouoJ auo uro{ pernses ralcnu pele8uola ol punoJ rlllt\ sllal eft eleql onprseJ enssq uos aql uorJ peuElqo suo4e^ sflec e8ftl eseqt o1 .{lrunxord asoyc u1 pale8 -resqo uo paseq ere suo4dr,rcsepSur,r.roqo; aqa Jo 'Jeln8ue :elqeus^,{:a,r st slec -uole 'punoJ 0'1, eseql Io odeqs eqa useldoft clltqdoutsoa sJ,'lnssu ,{13uor1spue rJoalcnu IIErrrs [ueul q]I^\ slleJ relleus puE 'ru/02-SI ralJurprp lsa8uol 1o 'utsoe pue urd g-€ Jo Ielcnu qltm s11ece8-rel ,{la^Ileler are eJeql :azrspue edeqs ur ,{re,r slac :eqlg pu? urnl"uaeq qlIA\ paur?ls eJaa\ suoll'es 'sllal snonurs,{JlBrJ aqt pu€ (urr/ 9) pauortoes se,t\ anssn pappequa -ue3'pele8uoyeqtr^\ pelenosse eq o1 sreadde eq; 'urgered ur peppeque pue palerp,{qap pu€ 'p3{J?d ,{lesuep pue cqrqdoseq ,{13uo-4s sB/i\anssn luopouoJ aql 'luauele arll Jo azls aql sr lerJJleursnolqrJ reqlg rnoloJ uI asot aled uo Surpuadap setnuru 0Z-gI ,{leleurrxordde pue lue8urr;errq sr lr Jo errJos ralJnu lc€l ol lool qJIq^ 'uolrexu Pue uorlnlosstP8utmo11o3 sreedd€ 'ses€c lsour ur 'pu? 'selpunq se srncJo 'ernsodxe uodn,{laletpaurur tsorule paler8elut 'Sursourols?ue-uou sr 'slleJ eql Suorue pos:ed -srp enssq pee4 eqt se 'lnJsseJcnsun ala^\ [t L 'res , qCl suanufuo? qppoopulH lo qualllep uJoJlullJed pegplBcop Jo xlJlBur anssll lJoS 90I n.i:.i.ua confluens [Ch. . Sec.7.4l Interpretation and discussion 107 : :: : -i non-anastomosing. _ : most cases,appears : :r is birefringent and - -- : :: fibrous material is : :ensely packed, and :,: :: .\ilh elongated,gen- / - .-,:: and size:there are r. - ruclei of 3-5 pm and :- r_: pm, and smaller - -.leoli and strongly : - - Tre shape of these - : .-:- ar. rOuud,or elOn- : :. :o these large cells : : irr elongated nuclei . .-- : rhan the nuclei of l-.-. -;.ls also have many . - -. nophilic cytoplasm - tit .t ' \ . I'J\ \\D DISCUSSION : : -:.-: of this study is the . -.ll can remain his- - : ' ::f,rd to its capability : -. .iains. It has been 14t '.. r'. - 1: cell structuresand qPl -: : : -.: natrix rn bone and - -- ::ralization, disinte- : -: -:on death, and only : :-::s in the form of -: :.rlier studies have 'tS - . ,. :ld as,170Ma retain ,: -. ? etzneret el. 1968). ::: - i a 5tudy like this is: i - There -..nraminants? ;,. _- - : :rr kinds of cells, : rl r 130 -rr! lsmall arrow), Haenalum ' and -- : i :nt (small ' arrow) t{! s -,r:: al (smallarrow) _ :-rll + rql qtr,!\ drqsuorleler ere eJeqJ ue8Elloc eq ol peJeplsuoc $'Ieltnu lEql sernleeJ Aluo aql lueserder €unnnl aqJ 6e: grr-1 osnlloru;o sed,{1 '8ur.raqlee,r ot Jo {3pl pu€ 'ecueSuuyerrq 'ernpruls str Jo Fef lrl llelnJrueul pue sluoP asn€req 'enssrl snorqrJ eql Jo e(uos lseel lY -qns Jeleu snql pue pesodxe flerree-qns releu mzla: rqeuaEol,{qd lcerrp 'sluerJlnu Jo s1o1Surpueruep anssn elll3e ,{lqeqord era,r fpnls slll ul pesn stueluele [.6i rnqll,/|\ pu" eqeleld drer' e sarldurt uor1ezlJelncsel altsuelxe eql luopouoJ eqr ',(1prlqJ parrncco e^eq JIIrt\ Ier '3'e) uoqezl ! Isrqrsert\ 'luarrJsle luopouoJ 3ql Jo enssq pazlJelnJss^ -raleru crueSro ;o uoqeperSep ptxJeql ou snql srseq eqt uo p paseq) I u.soq Jo aql Jo uoDeluesaJdar prrldo eql sr ,lallPtu :(LL6I p urelsdg 1o lro^\ aql uo 'sr:-rlno(x pue spodoqcelq elrqar, l€ql salecrpur f,pnls rno'unlleqlopue J.08-09 ueql reqAII se.rnle:adruel o1 pesodxe slql uI pesn ; n-n ezqe.rautuorq cDrlede ;o edfl e tuasarder ,{lqeqord aql Jeleu aral\ feql rpql saloqs ,{pn1s ;-uoure punoJ "ulunl Jql '^lpuoJJS lE]ra leql 1o auros 3uru11sllat pele8uola eqa palrodar sluJurel: luopouoJ eql Jo lnoloJ 'JJo I c-3 sr ipnls srql ul paJel ueaq lou s€q lueurala ;o ed,{1 stql ruo-r; lnJ ere/t\ sellddns Su4roddns-aJll eqt etuo :drr aq1 stuatuela luop sprol Jo uolleurrxexa WgS Pue 'a3.re1elmb are sarnpnrls leuJslur eql uo aJuengul [EuJalxe lllla-4 alnlJnIlS xIJlsIu neql fpnls sql ur pasn s]ueuale €d oql :stueuele ssslur pellnsoj a^Eq plnoqs qJlq,\\lJuJ E'Juoq 'qtoat uI pue al€q sluaurele er" 'laureue ;o ezs pue ed,{1aql qt1,l\selrel splo^ Jo azls eql ueql flrso:od ssel ,(lqeJeplsuoJ eqt qtlt\ FidBrr. leuorsecco teql [laIII etlnb osl? sI lI Pe^resqo erea splol luopouor 'sJopeJ eaJql dq Paureldxa aq iq sr ll 'euoq lel palecol dqeseq eJour,{pnls Jno uI s"elel{?t\ ,(eru sluaurele"(llsrrd luopouoc PIo €ntr 00t Jno Jo ,{! "lreuruJop pu€ *1c-: 1o Surlsrsuocd1a3re1 'dsnc eq1;o dq aq1 ol esolJ pa^tesqo uaaq a^€q -srureqJolsrqp€lur ,{11er1red1sea1 le eql ens a,!\ E uB qtr^\ J?lnllac-uou spro^ qcns WgS aql Jepun 'o r :pe^resqo araA\ -srl uos Jo uorlerl..reserdeql lBql e^ellaq D{ -rr .,$oul f,puasard a,n (pururnD spro^ aql lue(xele eql uo slnoqeareqA\ 'xrJleur lueuele luopouoc peIJIcleJep f\i lou sr enssq luopouoJ ol anp eq ,{eu (y961) rc13ar7 pue trtorlspul'I eql slueserder f,1enpz lsrralelx Jno leq1 8utf,es -selEproqc r.aH puE 'sJsnl dq sproa.-ro; palroder 1uq1pue sueurceds rno ur egesf,lqeuosear IaeJ eJoJereql ei1ysarSoloqd adfl u:epout 4:r-:q atelncrgeur Suoure ur ?ururnl eql Jo :rrs eql u::,\uJq ,(:ued:rrsrp -.roruolsrqrulluls q1^\ ansstg;o q reql uorlBzll€Jeulluolq eqJ pezrI€lntss^ ,(1q8tq se?t\enssll luoPouos € euleu ol eIqE uaaq seq uroql\ Jo euou 's1sr8o tcr]ue{ ol lsasols snql eql leqt pepnlsuor eloJelaql sr lI 'uolx(xot lolsrq snoJeunu ol suolpes Jno u,{oqs e^eq '(186I rsrl\ rc17ar7 pue ,{:ea ere eurunl Jo suor}J:s anbrlqo ssal ro a,,\A'uorlrppe ul uoqzreda:d 1ect8o1olstq 1o 3 uI lF :rllnotue IOUIIUSslsIOd eJor[ pu€ esre^Sue4 'ssoqces PalJl'le3ep Jno uI purl f,ue :oy pesn aJoJeq re^eu ^roleloqel ErLr'alll?d3-alPuoqr?t Jo 'pue 'sueturcedspeleerlun eql uI uourutoo f,ta,r '(esod-rnd raqlo ,{ue Jo) suoqeJederd 1ect8olol pue a-!c sluauale luopouoc eJ? spro^ qcns ororrJ Jo urti 0€ Jo releur€Ip -srq loJ pasn {lsnor,lerd ra,,r.euluaurdrnbe + roJ lunoJre (sepcaloru e aa.eq,(eur fpnls rno roJ pesn sluaurele luop lEuelsrrr qlr,{r 3p€ur uaaq elzq suorlendard I relnlec-er1xe raqlo -ouoc repcrged eql q splo^ eqt :peqcuetq eq llv'suonces rno Jo IEJoAeSuI slocru pessoJc r t{Enl pezlFreuruolq Jo ol reedde sos€Jeuros uI qslqa\ 'slasse^ Jo suoq repun aouaSuu;arq ellclsJ pcrd,{1 q8norqr 1Lr- iq patuasardal sI suou -ces esrelsueJl lueserde.r ,(lqeqord lsou sPIoA palselep'enssu eql ul uoll€zllereulur peAlos g rnoqe iluo flpuorgodor4 :seql '([86] :a13ar7 pue urollspul'I) tu/ | -slpun Jo slu"uureJ erz (,$lqrssod aql lnoq? pue) slueunueluoJ qlh\ r ruopouoJ eql tBql sl uoll€ JOJelauerp unurxBrx pal€lse qll^\'ullr s 0-I 0 pauro,r fprelrac e,t (r cuoul {ue pue sluaruala rrJorJazrs ur Surdre,r'spro^ rcF?aJJr o1 reln8ar Suqeep lou eJ? aA\ l€ql Surptcep ro; suosear E F:UDIeJep eql uea^\laq se dn aroqs ssrnlculs 6seql (WAS) adocsorcrru ,{reurud rno f,111qrssodpumas eqt o1 pre8er $ I€ql ssells ol qsl{r ert\ uorlcala Suruueoseql repuf) '(af,{A'd 'fset qtldyueurrreds pelee4un eql uo palJesqo -Jns-\l urapoul ur peztuSo n17ery pue uorspul-I aes) ,ra11eru e1rq,t, e-re,n srusrue8ro Jaqlo ro 8ur:oq 1o suoqec , irauapr uE 8ur,{1durrsural sB 01 peraJal ueeq e^€q seJnlculs eserlJ -rpur ou 'elouuequnJ 'pu€ 'lueurele paIJIs tq:-.erd le :eJa-rd aal 1eq1 'seJnpnrls enbedo e1rq,n sz edoosorctru lq8q -l?cep eql;o qt8ua1 aql Jo sprlql-o^u ragr i1:ea1c'salSoloqdroru eql Jepun pe^Jasqo aq ue3 seJnleeJpelsllecuEJ -rxordde perdncco e^?q plno/{ ,(pnts sltIl^lalElu JoJ Eqf -slleJ ,eurfqcueseru, se ro salJrsa^ full 'sruro; punodruoc ur sasEqrleql peuoqras enssq;o ecard eql epcs Jo asnetaq u pue 's11ac:e3re1 dllcurlstp Suo1epuz sap[uep pue dsnc eql ut ,(pepc4red ,{ldurrs 1no pelnr eq uDf, ,$1gq1ssodlsrq eql 'Surssacord !(qe urcuaJun aJe ei[ 'luaulele eql Jo seere ur"uac ur 'JeAea\oI{ Jno SurrnP PecnPorlut 'slleJ Surf,dncco 1oc u1 raqure 1q311f,1ensn are sluaurala paJell€ slu€ururetuoo (7) pue 'spror Suusrxe ; ps:adsrelur (relcnu aErel -ufl luauala luopouoc lJ€lur aql ul saJnl sursrue8roraqloro Surroq luoJeu (I) :slueuru€l m qrrn ua8elloc;o selpunq -cnIls u^\ou{ ql,t\ peq0uepr ,{ll'anp aq u€c -uoJ roJ seJJnos elqrssod o,tn1 eq o1 readdz anssll / qC] suanvfuoct4lapoapulH lo quauale uroJ:lupJad paUlJFJap Jo xFlBur lJoS 80I H"llq confluens [Ch. 7 References 109 ki:ed with known struc- bundlesof collagenwith smallfibroblasts (with cautious,since studiesof cellular mineralized d ,--ioodont element. Un- large nuclei) interspersedbetween the larger tissueof early chordateshave exclusivelybeen : L.uaIlY light amber in col- cells. based on the morphologies of cavities (pre- rrt:ia areas of the element, We are uncertain about the nature of the sumedcell lacunae)as seenin thin sectionsof Fq' :rd denticles and along distinctlylarger cells,and refer to them simply armour, scales,and teeth. open spacesin those Fod tbrms, tiny vesicles or as'mesenchyma' cells, There are alsoother cell of our sectionsthat are poorly decalcifiedshow r(=n b€ observed under the morphologies,clearly identifiable in Plate 7.1, that cavities in the undissolvedtissue do not r E re opaque structures. that we prefer at present not to describe in necessarilycorrespond to cell outlines. Until bre been referred to as terms implying an identity with cell types rec- stainedtissue sections of early Palaeozoicver- E Lrndsrom and Ziegler ognizedin modern tissue. tebrate tissue (e.g. ostracoderms)have been He: the scanning electron We wish to stressthat the major difference producedwe do not rule out the possibilitythat Itt-le structures show up as betweenthe decalcifiedmatrix of the conodont conodont elements may be the result of an rrlri-s. rarying in size from elementsand any known type of biomineraliz- early type of vertebrate biomineralization. ttsi:o maximum diameter of ation is that the conodontmatdx includescells. Neither, of course,do we rule out the possibil- u Ziegler 1981). These Proportionallyonly about 20-2570 of our sec- ity that the conodont elements represent an + -=Dresent transverse sec- tions is representedby collagen.In other types entirely unique approachto biomineralization. E r some casesappear to of biomineralized tissue collagen and some We welcome ideas and suggestionstiom n-= in rhe particular cono- other extra-cellular products (macro- readersof this preliminary report. a rt: our study may have a molecules)account for the entire matrix. r (a. anore. Such voids are Conodont elementscompositionally consist E [Eueated specimens, and, of carbonate-apatite,which sometimesincor- 6-cs. transverse and more poratesminor amountsof fluorine (Lindstrtim ACKNOWLEDGEMENT ni.ls of lumina are very and Zieglet 1981). Mineralogically they are L.E.F. wishes to acknowledge continued finan- rB:..:e concluded that the thus closest to francolite, a product of cial support from the Natural Sciences and rs r:sih vascularized. The biomineralization that is known to occur Engineering Research Council of Canada for !r -.:e size of the lumina in among inarticulate brachiopods,certain mol- the the Palaeobiology of conodonts. I tl;r reported for voids by luscs.and chordates.However. the decalcified study of 4k: r 1981) may be due to conodonttissue is not bone or dental tissueas ec{eaent the voids (lumina) we presently know it: Bone is either totally . m.ier the SEM such voids non-cellular with an extra-cellular matrix REFERENCES ld-- to the tip of the cusp, largelyconsisting of collagenfibrils or, in cellu- Er more basally located lar bone, it is dominatedby bundlesof collagen Epstein,A. G., Epstein,J. B., and Harris,L. D,, l- k is also quite likely that with the occasional'trapped' osteocyte. Dentine 1977. Conodont color alteration-an index to rh cirh the type and size of and enamel, in teeth, are totally different in organic metamorphism. US Geological Survey fuens used in this study Prolbssional Paper 995 27 pp. their matrix structurefrom the decalcifiedcono- [email protected]. E.. 1969.Conodont zones in the Ludlo- | 5E\l eramination of voids dont elements. The type of tissue encoun- vian of Gotland and a corelation with Great I eiement has not been tered in this study is also in no way similar to Bitalla. Sveriges Geologiska Undersdkning, ser. qga.redcells lining some of that found among extant invertebrates with C, Nr. 63933 pp., 2 pls. ttr represent a type of apatitic biomineralizationsuch as inarticulate Lindstrdm, M. and Ziegler, W. 1981. Surface rEdr indicates that 'white micro-omamentationand observationson inter- brachiopodsand molluscs. A. (ed.): f/eati,re bJ representation of the nal composition.In: R. Robison On the basisof known types of biomineral- on InvertebratePaleontology, Pa W, Supplement of the conodont element. ization (e.g. Westbroekand De Jong 1983, 2, Conodonta,Geological Society of America and rl.rtation implies a very Watabe and Wilbur 1976) we exclude any University of Kansas Press,Lawrence, Kansas, dins lots of nutdents. direct phylogeneticrelationship between cono- w41-W52. I rbe fibrous tissue, because Nancollas,G. H. (ed.) 1982. Biological Minetaliz' donts and inarticulate brachiopodsor known ation and Demineralization.SprirgerVerlag Ber- tsrerrngence, and lack of typesof mollusc.With regardto a phylogenetic lin, Heidelberg,New York, 415 pp. d t.r be collagen. There are relationshiDwith the chordateswe are more Pietzner,H., Vahl, J., Werner, H., and Ziegler, W. lle {oqs fllereue8 asaql x -:2.\ralq IeclqderBrl"Jts I geuodar aq teql selcods r i:gr 1 rddrsstssq4reddn :.ioryq5 8ur[1:e,ro aql Dl noauo eql Jo $un?J rglp snoncrdsuocpelou Dr'.id uErrrauv qlloN lL--I aqr txo+ sluopouos b :*.:I:ea eqt Jo euo uI -\OIJJOOOUJNI I'8 Brprl urorJpeurJap uaaq F aql 1? suon hs -Lr€punoq=- q euneg qtr.{rasoql s IE(r4 uolllsu€Jl eqJ uoll + : \rssJJSoJdJo ecuePl^e asenbas P ur{lr^\ slnf,co q1 -uonras-dn serJlsJa^tp r q3r e qll^\ peJnpoJlul sI + !-l _IolP^relul ue urql[A\ -t- parzldar ere (I L6I) EI :o J eunsc Jo sluop F lEq euoqEl)lo Pu? qeln 1 >i: ut Sutldures pe1rc1aq rcYurssv Fcf I\ ') 'uoldulqlg 1 lI I JO UeICIAopJO tEqr ldnrqe uv 'ddzts 'ssejd purlo.rEJqlnos Jo dlsje^run uuold puo 'puefug/uopuol'vsn/uotsog'pue oH/rqrerp sattuqatatut aql u1uottozlluaulla lo stustuoqcaut -roc ',{ueduoJ Surqqyqn4 lapre[ q uoltol aqJ gL6I ( spo) 'l,l ) 'rnq[,{\ pue 'N 'eq€te.i[ -nlunJ)D pJau 7on3o1oqpuo uolJonlDraulutorg ',2-8I std '791-911 971 octt|dot8 '€86I (spe) '16 g '3uo1 e(I puE d '{eorqlsalA -oruoaopd .ueluopouoCrap erSoloqdrou-or1q,,g lBrqunloJ 'dd I9t'EulloleJ qtnos pun SunzlesueuurBsnzuaqJsruteqJ rnz 'g96I , qC ] suan[uoc DllapoaryrH Io $uauele urJoJlulFad paUIJIBrep Jo xlJtBu ansstl lJoS 0 I I H.Ua confluens [Ch. z Err:-a. {61 pp. De .t.rne.E. W. (eds.) 1983. c; ::ologrcal metal accumu- | Company, Dor- bc:r-::shing LS \. London/England, An abrupt changein conodontfaunas in the Lower Ordovician of the Midcontinent Province R. L. Ethington, K. M. Engel, and K. L. Elliott ABSTRACT developed in the two formations. Subse- quently, these two faunal assemblageshave Detailed samplingin the Lower Ordovician of been shown to be widely distributed in Utah and Oklahoma has shown that the cono- shallow-/warm-waterdeposits in North Ameri- donts of Fauna C of Ethington and Clark ca, and componentsof them have been rec- (1971) are replacedby those of Fauna D ognized from the Siberian Platform as well within an interval ofless than a metre. FaunaD (Moskalenko1967, Abaimova 1975). They is introduced with a few coniform speciesand are not present, however, in the thoroughly diversifiesup-section. The faunal replacement studiedcoeval rocks of deep-/cool-waterorigin occurs within a sequenceof strata that gives in Baltoscandia (Lindstrdm 1955, Sergeeva evidenceof progressiveshoaling during deposi- 7966. Yiira 7974\. tion. The transitionfrom rockswith FaunaC to By 1970 th€ geographicaland stratigraphical those with Fauna D is suggestiveof the rela- distributions of these conodonts had been tions at the boundariesof biomeresthat have established sufficiently to permit Ethington been defined from trilobite occurrcnces. and Clark (1971) to considerthem as charac- teristic of two of the five distinct faunas that they reported to exist in homotaxial sequence 8.1 INTRODUCTION in the Lower Orodvician of North America. FaunasC and D, as they designatedthese two In one of the earliest systematicstudies of faunal assemblages,have become standard conodontsfrom the Lower Ordovician of the frames of biostratigraphicalreference in sub- North American Platform, Furnish (1938) sequentdiscussions of Lower Ordovicianrocks noted conspicuous differences between the in which they occur. Typical speciesof Faunas faunas of the Oneota Formation and those of C and D are illustrated on Plate 8.1. the overlying Shakopee Formation in the The top of the stratigraphicalrange of Fauna Upper MississippiValley region. He found few C wasreported by Ethington and Clark (1982) speciesthat he reported to persistthrough this to coincide with the top of the House Forma- stratigraphicalinterval, and observedthat even tion in western Utah. They found the lower these generally show differencesas they are 33.5 m of the overlying Fillmore Formation to 'dot.rolaquf0 !uor}]es 'uoFPlrrrogIlrH efzuetrl/{ // /t?^\q3rH'oglx 'seuof pue o5r\rq slsuauoslDpsnpoloauo Iz BrJ dot ur 'uorteurroC arzue)ow'uoqcas /tsaq3rH !0/x .qs:|urll.tuosuoiq snpoxoj Rrf ^\olaq 6 IIrH ,a OZ 'dot Arolaq ,06x trr 8 I 'uoqeuuoJ IIH erzue)rl l 'uo4ras {eelJ lalpueqJ 'qstl.ljni slsuapauo,snporv,-61 3q luoqErnroC ruortces .OU" ,qslnrJnl .RrJ e^oqEur S t laarC IooJ l""rr::fi*U, slsuato! ;nportuorv,_.Bl ool lrr 'uoFetnrog erzuatrl ruoqces {EAqSrH.061x .qslutnc snsuapsntnuD4o^olJ-Ll Brf ^\oleq e IIrH J // 'es?qe^oqe !00IX ,(qsru:ng) u60 'uoteurrod {aarC IooJ'uoqces laerJ rolpueql s4zazli snpoqrunv gI ArJ 'opProloJ luorlces .gzIX ,qsrulnl q.ln5 unosql 'uorlEurol nolrupl l ua\ss1q,snpo17Dd,-gl.3q 'dol tU .uoqpuuol ^\oleq 9., llIH elzuo)rw 'uopres leerJ relpueqJ'00IX 'uol8urqtApuE rlstedex slsuanotlltDusnpossoA lI St_! 'o01 iuor}ces ,{oteq rlj 6 'uor}prxroJIIrH erzuetctlt Z/ ,{e^\q8rH,ogx 'rallnyt lllunlralt DurpouosottJ-tl Aq 'dor .0S . ,t\oleqrrl S / 'uoqpruroJIIIH arzueycw 'uorlcaslZ ,{€^iq8rH I X iepuzd sryDInBD snpolelo)-Zl At:!t 'dol trl SgZ 'uoqprll.foC ,0tIX^\olaq .uel,roN {eerC looJ 'uor}res Z/ pue srle]tlpl aDu'tp snpotacoy,l 11 8r4 'esEqe^oqe rrl 6 0 'uorteruroJ^p^{q8rHlaelC IooJ 'uollras tZ {p^lq8rH,ogIX '.ds snpo|oau)i 8t9 'aspq .uorpes ,ggx ,.ds 61 e^oqe ur 6 0 'uoqeurol {aarJ IooJ lZ ,{s^\q8rH mpotoaugi-6.Brg ,uorlres .08IX 'aseqa^oqe ut 6 0 'uorteulol Ieerc IooJ // ,{E^\q8rH , ds snpopauoz_g -arl .0'x .qsrurnj tu € 0 uorterrrroJIIrH erzue)rtrl uortres learJ relpuEql snt0ulrltnsnpoqtuDrvj. L 3r.4 ruoqpuroC ur 6 IIrH erzue)rw 'uoqres lZ,{€^\qirH,00Ix ,qsFrnJ sltDvguaut Rrt ^\oleq'asEq ,09x ,ruprtspurl ,snpo$to,,g e,roqEur €01 'uoqsuuog ap€lqpur) 'uollras /t ,{t?^rq8rH xat snpodo\ocg-g 3q 'dol u g€Z ^{olaq 'uoqErrrrocIaorJ rooJ 'uosces ,, trosugrsj sryarDtDdsnpon4xnz-t-?tt ^"^:jjfool ,{oleq..'91-1 tu'.-(ryT.t:: uoq ,(lqaw 9€z -erruodleerJ IooJ'uor]ces //,{earq8rH'99x pu€ uosuetg) saD)udupDnbsnltorotd[tD-t Zlt 'dot urgtz'uoqprrtroCleerJ IooJ'uortres ZZ,{B^\qBrH,0/x^\olaq ,qsrulnCqsrrrlsrox?M Dutpoqr4n-Z -Brl 'uortpruro{leerJ Suuds ,(tqel lseA ]o eseqe^oq€ rrj /Z 'uortres // ,{e,{q8rH:00I x pue uos'd€rg)suD pp snporotld,t1-l .Bq 'J EUneCJO stuelllelA IUOpOUOC lerrd,{t}o ere IZ-l I 'l '9 sarnBrJpue 'J eunEJJo uorl]uqxa3ur^\ollo] pernpor}ur $ leqt euneJdtrsla^rp-^rol .S-l eqt tuasarder0I 8 saln8rJ'CEunrC Io slueuale tuopouoc le.rdrq alp.ilsnlltll satnBtJ-I g eleld ueeq aa€q pu? '(896I opuBs S e) ,(1uoye,rey eql Jo sluoPouoJ leqlo eql uaal\laq slsrxe aJu?ssrsuuocale l€ peuodal ueaq a^eq o pu? ,{1ruqp cqeue8o1,(qd asolJ ou leqt pepnlcuoc J SsUneCSUrUrelUOC $IJOJ 'AJeql{aSlE 'Uelel deql lesnoll aql;o dol eq11e serrepunoq Fun€J -Jepun uoeq p"q SuqduIes peceds ,{lesolc qf,rq^\ pue l"uollEullol eql ssoJJe lsrslJd 'qsrurnd uI (286I Dlsfaded) sexal uralse,tr ur JeqtouE sryotlns ,mpodolocg, pue sn1allotod 'g 'seneds pu? 'lrclaP uI perPnts p?q {telc puP uol o,tq ,{po leql pe^arTaq IrEIJ pu€ uofurqlg -8uqlg reql gsln urelsel\ ur euo eql 'suorlJas 'eJotrrllrC aql ur rar{8lq se8u€r leJrqdBr8rlBrls o,n1 ,{Iuo uo pes?q s"A\ uoqezrl"Jaua8 srql 3uo1 a,req puz lr eloq€ luepunq€ flSursearcur 'acul^ord luaurluocprw eql Jo uercr^opro aruoceq,{eqt tnq 'le^ratu srqt uI punoJ tou ra^\o'I eql urqlr^\ suoqeleJJoc rxJrJ fuaA loJ are,,A(116I Ir€lJ puB uofurqlg 1o strutolunu ,{lrunlroddo ue sraJJouorssaldxa Iecrqdp$rleJls -toc'g :) rrrortspurl xat snpodolocg pue esoq^\ euo puP 'sluopouoJ ;o frolsrq,(Fee '(1qe1r1pue uosue:g) snlalond snponqrng aq1 ur seSueqc punoJord lsour eql Jo auo s€ '(1qe6 pue uosupJg) snlotlldo.rponb snuotol pe,l\erasea sluopouoc pel€leJun Aq J Jo -d{19 se qcns C eun"C crts alJeJeqc aq "Un?C Jo lueuacelde-r ldnrq? eql 'eJoru1rC rea\ol eql ur ol pereprsuoJ p€q JarlJBe,{aql l"ql serceds1ec asoql PUEuoueurroc esnoH eql Jo lftd reddn -rd,(a ',{lrsra,lrparol Jo sluopouoJ esJ?dsureluoc II'8 res 8 qcj aJu!^oJd luaupuoJpltr{ eql Jo uelJ!^opro rora(rl aql q sBunBJluopouoJ u! aEuBqJ zII Hc:inent Province [Ch. 8 Sec.8.1l Introduction 113 ;; -- a.lrmation and those = replacement E::: :-.lbrupt tnodonts was viewed : : :rund changesin the :r: tis. and one whose t:rers an Opportunity :."=:-: -: \\ ithin the Lower d:: :::nent Province. .=. ': :ased on only two r.-:.:1 Urah that Ething- : .:-:red in detail, and - =-.. r Repetski 1982) in :.:ng had been '.. under- ;r. , :,-.:taining Faunas C :r :: a reconnaissanCe -:: :,: - -jir. and have been ! t u Ia\al fue le selcsJoqll 'uorl"JsprsuoJrepun sPeqJo ecuenDes roou IerAnII" eql ql€euaq sJegoo€slppuE eurlc ! F: \lasqo lou e^?q ad\ eq1 ;o se-rnsodxealaldruoc apr,rord uor8ar -u. s aqt ;o qurl urar{lnos eql uo spuacsep lnq 'esnoH aql uI u"ql prJ€ srql ur uorleleEe,rpue 1os Jo aJuasqe 'qlnos aql o1 adols eq1 uo qSlq st suollEuroJ rx[IIloJ aJolrl eJ? spod prauaS aqt pue roog uo, uec aq1 ;o adols aql uae/r$eq pBluoJ aql 'e8uEl eql Jo qfual FFIFn lO sluaIIJS?IJ uro,t daalsaqa 'qlnour s1rreau da11z,rurcru aq1 sutol aql ol I?trIJou ,(1a1eurxo-rddesI sIxE esoq/r\ r!f,fq !s In seuolsulelo ed,{l leql &€lnqu Surpuarl ,(Felsa,tquou eql uI eurlcu,(s r'roleqs ? uI suowllrJoJ aJoIulI-I IIE Io ssau{cql eql seoP st '(286I) I-teICpue uo13uq1gJo pue (I96I) pu" asnoH eqt sesodxa uor8ar pe11ne;-1co1q frn :el€rauolSuoJ elqqed azlurH Jo g uorlJes ;o f,1rurcr,rpreua8 aql '?upueJl qtnos-quou srqa e8ue11 asnog 'sedols ' lf,i lJuanberJ eql ur srqf,rq^\ 'uo[Jas paJnseeur rng 'a8uereqt go aqr;o adolsSurcz;-lsa,t 'da:ls aql ut a:e (7361) rr} sl€rls ,{llIS seeJaq^\ ecsJeql slcessrpuof,uBJ SurpueJl ,(lJalse/i\qlnos {reIC pu" uofurqlg f,q paldues suo[ces eql lll:rErl] [€1l4ap JO 1Ue1uO3 e (ey8ue:pen6'u u 9I qetn 'ur?g oIIJ :,^A€IlI 4 ::r{1 spaq esoql auols 'SZT.I 'tE 'ras 'tili[N) erag ',(e1pn e1n1 uolfBtol uolltos (e) Fn :qr Jo teql uor; f,puec Jo apISlSBe 3ql u/' op spuelxalEql p?oj pa^old s\p uotteruloC arourlld -ulun 3ql Jo lsBe url 8 0 lnoqE pue ,(e.'irq8rq HVrn 'YSUV XSSI Z'8 E rqr Jo ,{3o1oq111aq1 eqt Jo qlnos txl 9 61 ,(leleuruorddesl(pnls JoJ :r le ualut Peldu?s eql pepeles,fiqecol aqJ (I'8'8tg eas)gg-9 de.tr rnllrJ-F pazlpleua8 y uotl -q8rg 'g n puo,{eqqurl urequou aq1uo uteSe r+>\O aql UI s" UOUIUIOJ Sursu 'srx? puqcu,(s eql reeu f,a1p6 elna yo 'sarolloJlEql uorssnJslpeql ul pele rrs:qr ur saJ€ldIeJaAas lP -np^a puE pauodeJ aft suollce[Iocasaql tllo+ pcFl -uqt qll^\ sel€JeuolS psJeAOCeJSIUOpOUOa aql UOISSaJCnSq31n spnbas eql ur spaq eql eql uorlEutJoJulaql qll.r uostredruor 'qEln rueNe^\'EeJexaqJ uorlEurroJo,Iourlllc ie,{ol pue urorJ m erlslrJ P ol dn 1tiual Pue esnoH .reddnJo uorlcospaJnssaur Jo uolleco'l-I 8 3lg roJ ElEp ulPlqo ol Jepro ur €laql sallllesol ,lq Io sasual uI Ueqs {celg o1r{}l3 s?uneJoarl es€qlJoI IeAJeluId:epunoq d r.our:addn sqt q8norql aql q8norql perce oc e6 '(9961 punolrl) p.r.rjs at? suol]€ulllI"l eruoq€l{O ureqlnos ur dnorg ellcnqrv eql b1: re.n pue apnlqdure ,t'o1 ul JnJcoosp (I puE J s?unedJo crls.IJelceJ€qt Enq qsrr{rorq ,{eldstp spaq sluopouoc'palsllu€lsqns aJeat sailuerlncco 'G$ lrep runtparu ere i(1ec asoql Jr'O EuneC roJ {1.re1ncgrud's1rclap ;1 rJols eql ur slu€4ue-al Surppu;o Jo paquosep,{eql l?ql secuelmJo rF (ur ! I ot dn) euolsarutl e^rsnloxafgunlntu,{1reau aq1 Suqn;er raqlta F preuraqe se8pel eseql Jo uorluelureql qlra palqulesseara,$ (286I) ;o prr? JuolsuleJ8 euq ;o {r?lJ pu? uofuqla ,(q pel.afloc esoql uBt{l ; sarorspnru aurl dll?dlcuud saldues pacuds(lasolo erou pue reErel qeln | !rr$ ol ulql Jo slsrsuoc uralsea ur le,rre1urf.repunoq O-C eql ssoJo? (seunEJ rF '1eualur paldules eqt uI 1ru1epur peldues ear luopouoc esal{lJo suollnqulsrpplqderElletsolg eqr d$r?lc oI iqder8$Brlsoqt!'I (q) 'qelll uJelse,{ur le^relul PcrlIJJ aql uI ,{ra,rocerrood yo lcBJIueu? eq ppoc pepodar r p:r ruauuuord dq donlno faql ler{l c eunuc Jo ecuereeddeqpEunrolloy E :-Uozuoq paldue5 aroru (I Euneg Jo uoBrnPoJlul ueppns eql l"ql os I IaiOI eql pue euolsaur-I 'erorxlld JeA{o[ruor; seldues e,rtlcnpordaq1 ut f-ll dol aqt sapnlcq leql sluopouosJo setuePunqea\ol punoJIJEIS r r'f ; e q8noJql sp,r:elul pue uolEulqlg lueurerBlderIEun?J ^Je^ ldnJqB u" Fellor araa saldueg lured petep lou plno^\ l3q1uoqnloser l?crqd?r84eJls i eqr iq peryeu sr'uadeals -orq,r,ro1apv'o.rd sgoder aseql 1o 1e 1(3661 r(986I F ]!-t loog eql aleq^\ PalsJol qsrurng) suortcosalrsoduroc a^lo^ul lo Pms"eur eq] Io es€q 3qI uey,no51)qe,r:e1u1 peceds dlpeorq l€ pepelloJ [z'8 res 8 'qJ] aru!^oJd luaulluoJpll4 arll Jo uBltl^oprg Ja^rtrl eql u! sBunPJluopouor u; a8ueq3 711 bdnent Province [Ch. 8 Sec.8.2l Ibex area, Utah 115 The baseof the measuredsection, which is CHEBT NOOULES locatedwhere the floor of the valley beginsto FLAT PEBBLECONGLOMERATES 800 steepen,is marked by the letters KME in red paint. Sampleswere collectedat 0.61 m (2 ft) intervals through a 24.4 m (80 ft) succession that includesthe top 11.3m of the House Limestone and the lower 13.1 m of the Fill- more. Sampledhorizons are indicated on the outcrop by prominent red numbers. (b) Lithostratigraphy In the sampledinteryal, the House Limestone consists of thin to thick beds of limestone, principally lime mudstonesbut with somebeds of lime grainstone and of lime wackestone. These ledges alternate with intervals of silty limestone (up to 1.5 m) that weather to form re-entrantsin the slope. The limestonestypi- cally are medium dark grey, but the silty inter- beds display brownish hues. Ripple marks of low amplitudeand wavelengthas well as cross- laminations are sporadically distributed througb this uppermostpart of the formation. Black chert in lensesof the order of 5 cm thick Grri:::: of upper House Fa_i. l:€\-cllon area, western Utah and up to a metre in length occur in 5-10o/oof the beds in the sequence.Flat-pebble con- glomerateswith thin, discoidalintraclasts occur at severalplaces in the sectionbut are not as Fig. 8.2-Columnar section at Ibex; sampled horizons r r:e svnclinal axis, rising mdicated by tick-marks to right of sectron; section cali- common as in the overlying Fillmore Forma- brated in feet (above) and metres (below). E :rb beyondU.S. High- tion. A generalizedcolumnar sectionthrough ,. 8-1 The locality selected the sampledinterval is shown in Fig. 8.2. ElEl! 19.5km southofthe The lithology of the sampled part of the The general increasein frequency of occur- 0,! im east of the unim- Fillmore Formation does not differ signifi- rence of flarpebble conglomerates accom- ha< down the eastside of cantlyfrom that of the underlyingHouse Lime- panied by an increasein abundanceof grain- f\\ 1 -1, Sec.31, T225, stone. Those beds that have a relatively low stonesat the expenseof mudstonessuggests a hl 1j min.Quadrangle) a content of detrital material protrude as ledges progressivelyshallowing depositionalregime. x ,'zlvon dissectsthe face whereas silty strata form re-entrants and The section was examined carefully for the :esu;ed section,which is in slopes.The frequency of occuffence of flat- presenceof stratigraphicaldiscontinuities which 'o: SecdonB of Hintze pebble conglomeratesincreases up-section as might account for the sudden termination of ;t,--'nand Clark (1982), is does the thicknessof individual units of this the range of the older fauna. No obvious dis- tending f tributary that type. Grainstonesin which the grainsconsist of cordancewas found, and corrosron zones or t E3r its mouth. The steep worn fragments of tdlobites and of brachio- hard grounds were not detected in petro- -:oor r and the general pods are more common in this sequence graphicexamination of hand specimens.A dis- I regetation in this arid than in the House. continuity may be present in one of the re- qr{ete exposures of the We have not observedan abrupt changein entrantsin the lower Fillmore, but if so it is a de: consideration. lithofaciesat any level in the sampledinterval. feature of low relief. Lehi Hintze. who has FS parns€eu eql uqlla\ eql s? eA\ s€ 'euolseurl esno11raddn aq1 {JaAare eJou rClse^{ol pue esno11lsoruraddn Q _ro uo4stuaunJop eql urqlh gtoq'(eaoz sqpdolqaldoto4) 3 auoT aql uI punoJ a^{ slua(Ilalaeql eraq papnls h+Jo:d eroJaq pauruexg Jo IIe pue (euo7 outnsk1dtu(5) g euoT srq;o &ll?Jol eql Jo quou elrrr e lnoq€ sr rlJrq^\ Er !-ruopouoc JoJ paulelqo ged reddn eql sessedruocueuorlJes peJnspeu J uorlJesJreql lB alourllrl Jql Jo lredsrql uo.r1 t -Ei.rgrpp€uJ slrssoJeseqt rno selrqolrl Jo uounqrJtsrp pcrqderE sluaurela,te;e fyuo pagodar(9 elq?I'286I) :3$ aJea\ f,aql qcrqal ur -rl?Jls eql uo ,{preurrd peseqsauoz Fun"J Jo {rplJ pu" uofulqlg 'uoq"urroJ aql Jo as€q tr roJ -illn]eJm paururexe uolssaJJnse olur EJlexaql eql ul (uerJl^opJo eql e^oq€ 0J S'0I) lII Z'€ pelcellor sea qrrq?t\ rqf aroullC eqt ur (euoz slpprtr^J) elrzlreno B)lerng eql qluauaq Jrd, 'qsrurnc snloulxunsnpoqtuDJvL 'qsrcJniI rll A 2uoz;o lred rearol slcor uerrhopro aql pephrpqns(IS6I) ezturH ntoln8uortl ,snpolsro,'uol8uqtA pue qs1ade11 slsuano|tuDrusr?possou'qsruJnC uaFsoq ;npol sllssoJraqlo (p, -lrd, 'qsruJnCsnlouDun snpoL[uoov 'qsrurnl snpauq snpoqluocy) sraqruaurlu€ururop aql FE :-z (r\oqe) teal ur sr 'erotIIllc lnq 'aJourllg aql ur puno] lou ere,r uonercoss€ t F-r:;:GrrerlS-€ 8 8rC Je/i\olaql ur ur 9'0 u?ql arour ou Jo lB^reluru? Ieunq eql ;o slueuodruocuouurocun f1e,rq urqlr,t aceldel.(eqt qJIq^\C eun€desrellp pue -eler eql (g'S EIS aas) ,(1snov'a:dpagoder lu"punq€ erolx eql Jo requeur 3 Jo lu?puecsep se,(repunoq elorullrJ-esnoH eql qlra\ eprJuroJ snorlqo ue sr€une; ,&rsra,rrp-,lr'o1srql ut serceds tou seop C ?uneCJo e8uer eql ;o 1ru41roddn 3ql Jo euoN '(lJ s'8€) ru le evorlap v eql l?ql eraq peluesard€l€p i eu eqI oe ^{oqs ol asola snpo)V ;o sarcads'II € Jo lueuale S 'esnoH ue srlEql raqlouePue erourllc Jql ul 0J 9 tt) eq1 ur sefoloql1l errr"s eqt qty( pendruoJ ru 9 0I 1" srya11otod'gqlt.,r ,{Jrluaprdle^rtel se aJotullrd aq1 ur serEoloqlrllp ur pepef]eJ -uel e,rt.leql uaurcadse pepler,(suorlJalloc lue srsluopouoJ Jo JJuppunqe ur uollJnpaJ IeJJuea r'9 'uorlelardralur l -sard aq; l€ql arJoJurerslJnsaJ eql 'sselsqlre^eN seuolspnluaurrl lxo+ au?J rng (rapue4 sntDqap snporv 'sryalond seuo uaJreq o,{u eql pu? seldruesluepunqe I snponq)ng's'ntortldorponb snuotoldtlg) q ls?al eql puE'euolsurer8 eru11yo sa8paluror; + euneg Jo JllsuelssreqsperaprsuoJ ,teqt sluop pelJeJIoJJr:,r sl€^relurqloq ur se;duesluep -ouoc eql Jo uorlJnpoJlurJo le^el aql A{olaq -unq€lsorx aql'slusruele gI ueqlle/'\eJ ?ur^eq eune; flrsra,lrp-anolB qI^\ Ie^Joluru€ Jo eJue seydurzs11 q1r.rpue (97 aldues) 9gg 1o runur -serd aqt palelnlsod deql ,(puenbesqngeroru -lxer[ s rllrr\ slus(uelers sea\eJorrrllrd rea\ol -III-r Iessq eqt urorJ,{re,roler asredspeq,{aqt uror; salduresur pla aEera,reeql 'ls€JluoJuI 1eq1pa8pel,nouqce ,{oql tnq'(1161) rye13pue 'Euerrrele0S u?ql raaal pacnpord (Sf 'tt 'gf = 95 uofurqlg fq O ?unedJo lueuodruoce pereprs '8'g'€'I) seldruesuer'ss ,{1uo pue'stuaruale z:! -uoc s€a\srrrroJ eseql Jo auoN (zg6l) IJ?13 649 sen (91 aldures)plerf tuntrrtx€ur:eldures pue uofurqlg 1o 9'ds snpopuot( puE 'rxolls red sluerueletSI Jo e8ert^e u? paple auols ag- -put'l snpo,oauool raJJr J^r leql setJadsuroJ -eurl esnoH eqt Jo dot eqt g8norql pepalloo -Iuooprqp ue 'snpolstouodatq3o sarcedse are seldruzg dlperyeur parru^ ecuppunq€eqt tnq - uorlelndod aqt Jo sJeqr[eurEuurncco ,(puanb 'Eueruele 7966 pacnpord seldruesa,rrpnpord -JrJ lsoureqJ'qleJueq Jsoql ueql dlrsJ3^rp 6€ eql uerrcq ere,rn(ru 1 11 l" Pue ur I9'0 1€ ra,no1.(eldsrp uopres pernseeu rno yo ged esoqt) saldureseql ]o o.lr1,{1uo fpessecord selr reddn aql ur sluopouo3eql 'acuepunqeJessel saydruusaqt Jo qJBaruorJ lcor ;o ure.r8olqy cJluoqrppe uI uozrJoql"ql asoql txol] ^\oleq lueJeJJrpr(lloq,n eIe ero(xlld eq] Jo sa4arrr ,{qdBr8pBrlsolqluopouoJ (t) € IESPqaql a^oqs pere^oceJsluopouoJ arlJ .dpnls Jerlreeaql 'uoqceseql Jo lred erourpg ra,{ol-asnoH ur palJetepunuaeq pup a,rarse g8norql pessed raddn aq1 ur dlnuro;uocun JoJ eJueprla e,rzq deru,{aql lpql os 'uo4Jes eql ul Je^\ol pcrs,{qdpepodar 1ou seq '(9461 'I96I) qeln rrJo+ esoql ol lseJluoJ ur Jepuels puE II"us uJelsel[ ur serpnls puor8er elrsuelxe euop lz 8 ras g q3] aru;,ror4luoulluoJpltr l eql Jo u?lJl^opJg Ja^rsl aql uI seuneJluopouoJ u; aEueq3 911 tntinent Province [Ch. 8 Sec.8.2l Ibex area, Utah r77 E :(rntrast to those from a0 r- ir rhat they may have 21.1 r=; been undetected "rd in 4.":red above the basal 3 2 f,-:: are wholly different z 21,3 E k: :rorizon. In addition to b ,-onodonts in the upper : 6 rEJ display lower E :E:teath.-ction The most fre- E of the population d lE-::'ers : stodus, an albid coni- E : ::::r ro Oneotodus Lind- 's: 15.2 E9 fu. rl. 6 of Ethington and 5 '- : c: :hese forms was con- .-;.EEi; P r --: Fauna D by Ethington iE " k --:er acknowledged that 't2,2 ror:1 tiom the basal Fill- E / I:d\ postulated the pres- r:-: a low-diversity fauna a:.t.luction of the cono- E ,'faracteristic of Fauna t:.;tlicarus, Eucharodus i :..idnts Pander). Our ; 20_ I I =r.rpretation. The pres- : i bi . .p€cimen that we ten- 6 t E. carqllelus at 10.5 m r*--: :nd another that is an ; cie-< of :{codas close to ; 3o- r '-::.-i ft). None of the - lre-:n fauna is an obvious T *e : erfthe more abundant J J r::;h rhey replace within rc --ran 0.6 m in the lower Fig. 8.3-Stratigraphtcal rangesof conodonts,trilobites, and brachiopodsin the lbex section;vertical scale is in feet(above) and metres (below) Trilobitezones of Ross(1951) and Hintze (1951) are shown ln rrght cotumn, ri+i rhe Ordovicianrocks lower part of Zone D (Leiostegium-Kainella quence,the rangesof the trilobites shown on :t= Quartzite (Middle Zone) in the Fillmore. The entire sequencewas Fig. 8.3 are comprehensive. lxr area into a succession examined carefully for trilobites, and all ledges Trilobites recoveredfrom the Symphysurina tc :nmarily on the strati- in which they were seen were collected for Zone include Bellefontia chamberlaini Clark, ir of tdlobites. Our these fossils. In addition, each of the samples HystricurusBenalatus Ross, Hystricurus para- b:nrasses the upper part obtained for conodonts was carefully split and BenalatusRoss, Symphysurinaglobqcapitella *-,:.ina Zone) andall of examined before processing in order to add to Hintze, and Clelandia utqhensis Ross. The pe--:-. Zone), both within the documentation of trilobite distribution overlying thin (3.28 m) ParaplethopeltisZone -iae:rone. as well as the within the measured section. As a conse- contains Paraplethopeltis genocurva Hintze, 'eete F Eo{ sraqlo aql :uerEq le^el aql l? {earc Iooc aql Jo es?qeql psulJap tJeiFusqJ eql uo+ seldlu"s oq,$ (€86I) ulls pue (696I) uleH paarofloJ xaql eql uI asoql ql!^\ sa8uer lecrqderSrlerls I iJ.a\oJar roJ passaJoJds€^\ o.r\ '1c?luocaql l? ur eSueqcpunog Jr:ql pu€ sluopouoJs1r 1o uosrredruoc:o1auo E rtro-g {Jor }o ur"JSoIDI v -ord e ;o acuasqBer{l ^EoloqII uI lceluot lsuollElrrJo] aleudorddeuz sr snql ecuenbassIqI O euneC eql uo peJlualur €'81 Jo 1e,r:alutue q8no:ql sureluoo uorl?uuoc rlaarC 1oo3 8ur,{pe,to irldpBrrsolq luopouof, (q) peldrues se,n suoqces o1{l eql Jo qt?g eql l€ql pu? uo4arrrJoc IIIH erzuextr\ eql 'perpnlssea Pl€JlsJo uolssaJJns 1Odo1 eqt t3 rmJo J eunEdJo sluopouocl€ql 'sluop rrunuoJ f,lqeqordsr 1eer3 elaldruoc ,(le4uasse ue pue 'sllun earsseJer s,troqs,{Fealc UodeJ eql 'sseler{ua^eN I !-lBl aql uaa,tlaq Punor8 aqt Jo Sulldru"s pa^{olp uortJes Jo eull eql -ouoc eql Jo luaul"aJl truouoxel lualsISuoJUI 'sedop lrrf 3q1os 'uoucas ,/ de,$ uro{ slasJJolroqg dsserSelpapun spaq ue sluasardpue uoqelueunrop pcrqde:84er1s 'surclunory Endo'J ruopouoc o,t$ eIrrES raqlo seereqA\'aprs,filunoJ aq1 ssorcuflpeer -olq s{r"I reded sq1 ep1cnqrv eql aJues ?upuorslp elqzreduroc e pe,trolloJeq upc leql sernsodxesnonuquoJ ur uercrlopro Je,tro'Ieqt Jo u?d Jo ,{pnts palrodar Frs FJereJ,firuuoJuoJun se i(llueunuordlcalord ol urel{l Jo elxos asn€c -sreuuoJer€ ruo{ sluopouoJ (9961) ELl:s ql ur luaureceldar ecuenbesaq1 urqlrrir speq luereJJlp ,{q padeldstp Punontr 'qeln urJls:,{r uI ;e,r:e1urpaldures 'r{see ; *i:;'8ns drqsuoqelarsrql Surreqleemol sasuodserslqeue,r eq1 eql qlIA\ aarleleJJotare lBql e1?4s sePnlJur g ruopouor re8uno,{e qcrqm sr Eurpaloc pue Surrnseaurleql os 'seJnsodxe euroqsplo uraqlnos Jo surBlunoll sllrlcl1!\ tF u,r ur p1'7) 1a,ra11seto1 qloq le anuaS ere sadols eql pue 'f,ldeels pu€ eplcnqrv eq1 ur dnorg eplrnqrv eqJ, |IIE J Bunec Jo eJuauncco drp slcor eql'sraua\o aql ruoq peurctqo eq p ::.Ti s1 punorEpreq s1q1 lsnru [pn1s -ro;uorsslured pup 'seqcueJpeu^ro f,qdpJ8llBrlsorllll pnB suollBtol uopras (B) Dr \z Y.iFuenbesqns ateFns fyale,rud uo eJ? suoqJes r{log '€Inoq?l{O brlno ralpueqC aql uralsaaqlnos sul"lunow elrq3[l\ aql 1aa:3 Jo VIAIOHVTYO slg{tooJ aql ur sI (a13uerpuen6 utur o F\J aql Jo assqeql e^oqe Jo 7/1 NUtrHJnOS'dnouc srvJnfiuv 0'8 E!ar+ pal3elloc uarutceds , €uroqpHo 'rnds spr€qc1l 'leZIU 'N?I ;&-6orrad Suunp pezruEocel '?I ces 'tlInAN''/IAN :€86I rltrs) uoqces 'sumluno] eplcnqJv t[r1-\rp PaurJeP,(lftelc Y {eerc relpusr{c eql J 's:lrloo pu€ selqoleuroJls uJelsa.irreql ur qcu")I uzurdeq3 eq1 uo 'se8uerraEuol qcnur eleq satcads url.To eqt fq papr,rord pelerol sr (e13uerpen6'ulrrr ZII L eruoq?plo et€lnorueur eq1 pue (gocp16) usua?nquoq rJ l,]of aql Jo Ued Je.&\ol 'stleC reurnl 'AId 'SZI 'ZI'tes 't/LAN stqtnuoN ts"rluoc uI '(* € t or 4 9 tuo{) p .-uunp Eur,,,ro1pqspanurl :ggOtttps) uoqtes ,, ,{e,tq341eql (t'8'8IC peprrlser eroru uele sr ( oJpid) llaqduoc E -ruaurpes Euqelnruncce ees)1e1ep ur porpnlsueaq p"q seuneJ ouuqdolutg pue 'uI 9't ol , € ulo{ uorlJes ELi. {q pe^oual eJe^\ elrqopl nJql Jsn?JJqpup ^p€eJIB :.Insodxa.1o sseu aq1ur a8uer palrrull E seq redoo3 pu€ qtlrln d -r:rear8 rale l€ql os etrrll -alaldurocpue ,{llgqrssacceyo esn?3eqpapalas 'otDisol ntu s-tqitouoN lJeluoc l?uorlPurroJ ls::6ord Jo uoneclpur ue 'suorpes oa\l le fuepunoq {eer] IooJ-IIrH aqt uo paJluer uonJJs eql ;o u-r9 q8norqt 4 m tqiraq Surseercurq1r,n erzue)cl4l aqt ssorJ" saldurespapelloc e,16 luepunqealrnb are sarcadsalepcqreur sesreql\ q : \rleler sauolsurerE aru11 'esnoH el{l uI ueql elolullld eql uI Je,{ol qtnul paleFlsod sr sercedselelmru? aql acuepunqv uolcas E:r-1tn eqt l?ql 'etuoqel{o Jo parns?Jur rww:6 pprolad pue seuols uJeqlnosur suoqeuroJrlearJ IooJ rei\ol pue IIIH elzua) Jno ul luePunqetou lnq luesald s:,.:ns peldures eql uqtlrl -cl^l reddn Jo suollcas parnseeu ]o uoqetol-t 8-3rc are spodorqcerqelelnJqftul pu€ etepcltrv bpqrrl i:erurrd eql 'p,rre1urlsaqEtq sq1 ruo{ mtnJuis[tlolod ^{oleq te-g L'I'9'9 n'L € le ua{?l lo pua sntn 4sKH1o sarcadspue'B8urytq'X 'lJ 'lloJlerf6 rylPpY ll .li\oleq pu€ e^oqe o aulDy slsr.rr,?oJluout wn8asola1 r aqr qino:ql sle etur (lJ I) pere^ocal alA le^a[ sql a^oq€ peule]qo pue Y eJe^\ sJeqrxnu lnq 'eJolxllrJ eql IIc lJ€luoJ l€uorleuloJ ..';ij;i:,,R'*"' lsapou '.trll?rol qJee le uel?l aJelA Jo ru t t lse^\oleql ur punoJerall\ satrqolrt oN """,.1,i::,"*" 'Ia lf -suorDas aql Jo seuolseurrl 'DoJlEN\ rsguqlq'X o aunt pue 'ezlulH F puB sallloo snoacqls 'Pues owndsocun ouunstqdu,t(.9'ssod snlopua8 Jo rrorlcnpoJlur ldnrqe 1o sntnculs(g'azlur]il. olcatouagwladoqla|duto4 'qC] aJulaord lueupuoJplw aql uElJ!^opJO Jo^rsl aql u! sBunBJluopouoJ ur aAuBqJ 8II [€'8 "as 8 Jo Lrtfuent Province [Ch. 8 Sec.8.3l Arbuckle Group, soulhernOklahoma L1.9 of abrupt introduction of abundant quartz 1004 specimens.In contast, the Highway 77 sand,siliceous oolites, and stromatolitesin the Sectionhad 10 barren samplesand produced limestonesof the sections.Thirty-one samples only 537 specimensfrom the productiveones. were taken at each locality, beginning at the The different frequenciesof occurrencein the formational contact and continuing at 0.3 m two correlative sectionsmay be the result of (1 ft) intervals through the next 3.1 m (10 ft) their respective paleoenvironmentalsettings. above and below it. Additional sampleswere Stitt (1983,p.5-6) suggestedthat at the time takenat 3.7,4.6,6.1,7 .6, and9.1 m aboveand of deposition of the Lower Ordovician rocks below. The primary lithologies encountered the Chandler Creek area was deener and/or Er-::d secttonsof upper Mc- within the sampled successionare lime mud- quieter than the Arbuckle Mountains area hi a.3ek ibrmationsin iouthern lEi-':::a stones and peloidal grainstones.Stitt (1983) owing to their positionsrelative to the axis of postulated that the increasing abundanceof the SouthernOklahoma Aulacogen. lime grainstonesrelative to lime mudstones Conodonts are less abundant in the basal ;rG< acrossthe McKenzie with increasingheight in the McKenzie Hill is Cool Creek (l = 19 elernents/sampleat Chan- trrlldat-l at two sections, an indication of progressiveshallowing over dler Creek, 9.5 elements/sampleat Highway eribility and complete- time so that ever greater proportions of mud 77) than in the upper beds of the McKenzie d becausetheir trilobite were removed by winnowing from the Hill (- : 57.5 elements/sampleat Chandler lcEn srudiedin detail (see accumulating sediment. Evidence for con- Creek, 47.7 elementsper sample at Highway -- Er Section(Stitt 1983; tinued shallowing during deposition of the 77). As is the casein the Ibex area,the reduc- ES- R1E- Turner Falls, lower part of the Cool Creek Formation is tion in frequency occurs at the formational i- Quadrangle)is located provided by the occurrenee .of digitate boundary. Conodontsof Fauna C are present R-aach in the western shomatolitesand oolites. throughout the sampledpart of the McKenzie r. Tbe Chandler Creek A clearly defined discontinuity surfacewas Hill at the Chandler Creek Section (Fig.8.5) ;N-\r 1 4. NWl/4, Sec.14, recognizedduring petrographicstudy of a hlnd and somemembers of this associationcontinue ldr $pu1, Oklahoma 7 specimen collected between 2.1 and 2.4 ir through the lower 2.1 m of the overlying Cool $e r is in the foothills of abovethe baseof the Cool Creek Formationin Creek. In contrast,Fauna C is restrictedto the lcams of southwestern the Chandler Creek outcrop, and this subtle McKenzie Hill at Highway 77, as reported by Ectrons are on privately surfacesubsequently was found in the outqop. Mbund (1968). This seemingdiscrepancy be- ipe:anssionfor study must This hardground is just above the highest tweelr the lithological sequenceand the strati- h rr*rerS. The rocks dip occurrenceof Fauna C and slightly below the graphicaldistribution of the conodontsat the bge-- are gentle at both lowest level (2.74n in the Cool Creek) at two sectionsmay reflect the way in which the Eunne and collectingis which a youngerconodont fauna is introduced. formational boundary is identified. The influx resDonsesto weathering This relationship suggeststhat abrupt faunal of quartz sand into the basin that is used to I bds *'ithin the sequence replacement in this section is the result of define the baseof the Cool Creek probably did b r'roiect prominently as unconformity. Careful scrutinyfailed to detect not reachthe two sections,which are approxi- rs rhat can be followed a cornparable discontinuity separating the mately 120 km apart,at the sametime, and the mu-r-side, whereasother same two conodont populations at the High- formational boundarymust be diachronous. dooes. Short offsetsfrom way 77 Section,so the presenceof the hard- The occurrencesof FaunaC in the Arbuckle dlos ed 53pp1iot o1 ,h" ground between the two faunas at Chandler and Wichita Mountainsare dominatedin abun- I aa essentiallycomplete Creek is probably fortuitous. dance and in stratigraphical distribution by Es srudied. the samespecies that are dominant at Ibex. As r s.ctrotrs was sampled (b) Conodont biostratigraphy is the casein westernUtah, Loxodus bransoni, d 1!-3 m centredon the 'Acodus'oneotensis, and Cordylodus angulatus In the absenceof a pro- A kilogram of rock from each of the samples are present but rare, and they occur sporadi- do$- at the contact, we was processedfor recoveryof conodonts.Five cally amongthe samples.Species that are pres- t) and Stitt (1983) who samplesfrom the ChandlerCreek Sectionwere ent in Oklahomabut only rarely representedor b Cool Creek at the level barren; the others from this localitv produced absentin the Utah section are Clavohamulus c: ::]?JOl Ueql JleqsSuDeJ 'uorl ;o aruos palrodar (896I) puno6 tnq 'suon I:: ::lnJlrJ peprlsar qJIV -eruro; eq1 ur :eq8rq ruor; saldruesur lu"urrxop -Jes parnsPau Jno Jo Jeqlre ur punoJ era^{ c r,:: --:qt lluouruloc sllun I"J pue luEpunq€ ,{13ursea:curauoceq (I eun€g eunzg yo 1ecrd,{tere 1eq1sarcads op rxoJlspurl rj\ iluaurluocsu€rJ eql Jo ;o slueuoduroc 1ecrd,{1aql pue '{eerC IooC aqt snpoloauo Io serlads e se ,{1a,rqe1ue1Suru8rsse Ea: .,lereue3ere s{toJ uerJ Jo aseqJql r^oqe u Z0Z rcsr snto.ttldotponbg ele e^{ leql suoqces-ssoJc 8ur(re,r ;o sluaur .I 5 i: :.joarv Jo elsls lusseJd Jo eJueJrncoo tse^{ol aql 'uo13u {tg U -ala uJroJruoc 'prqp ;o ,(1oq,tr slsrsuoJ uoqel r::.:: spuEISrpu€ roou-ees ,{q apelx'uort3as teql tuo{ uorlJalloo Jaqloue -ndod esreds srql ,{lrsra,rrpr'ro1 ,{:a,r;o eune; =---Y Ietuaunuocsu"Jl aql uI 'uorlcas 22 tensq?ry1eql uro{ paterlsnllr peqsrra,rodur ue paplerf, C eun?d qtua esoql T\ :nes ,$oll?qslnq peorq punow leql sluopouoJ aql Suorue peluasaJdaJ a^oqs sIJoJ aql suorl3es euloqsllo qtoq lv r?--1 :i{1 roJ ldacxe qs€,t\e ete aouDtp snpoD)DW pue 'dds outpo% 4n 'qelll urelsa^\ rr.|, -,ro-\ aql pue 'tualxe 'snto)rldotponb '9 'srya11otod'g se sercads ur $ 1rse ',(lelrlelrpnb pue d1e,r4e1puenb'euJes r -- :? se.r Ees )nES aql r{Jns 'ssalequele5,l lcedsns eJE O euneC aql qJnu sr €IxoqePlo ureqlnos ur 3 ?un€c 'Errdorl E:f ureqlnos eql Jo sluopouoJ Jo soJUeJJnJco pcrqde:Eqe-r1s ;o uo4rsodruoc eql 'surJoJ esagl Jo uolldecxe ::: '.- luor;er qtog rolenbe ,Lol aser{l 'luelsrsuoc lou eJet\ BxBl luopouoc eql q1l,4i| rallrw snpouuo)uara5; 1o setceds E:-r to lsaa\quou aql ol Jo suoueJrJlluJplsrq esnpJe€ 14 femq8rg e puE'seuof pue eJnJO srsuauoslopsnpopauo ts- ::': ?Iuu qJrq^\ le 'palrsod l? {eeJC [ooJ aql Jo lJed l?seq eql u]o{ uraql 'ngn11tr tt1u4lnq ourpouosoq) '\sruiq snsuap f-:-:ro-I learJ looJ pu? -(,{loteql l, ::-:'. Jpnulel S.0Z inoqB sarleu pue (a,roqe) laal ur sr ale.s :suortoes elnoqello eqt ur stuopouor uopnqFlslq-9 B1g E-- eqt 'uorlsl lPcrue^ Jo 9 -: --tuoqPl{o (9r6I) pr.r:\' ssod ,{q pezl l6 rr .-:rr \opro ,{yreg Surrnp E: :ru IEJlua, ;o 3uq1es u. =--qderSoeSoeeyed eq1 I T I I \olssncslo t'8 6 'suorlf,es stuoqBIIO ; rF:': iuo{ u,t\ou)l slssoJ _ol gtr--: -ruasa:d tV suorlJes I -; o r: '- saJedBlecJo suollces Fr: -:\a er'A 13q1 le^ralul I rrorrl eql uI suerurcads r:::r ilr = f, ::: p-\\ 'serPnlss.lllls uI 9'E 5-L9f "._]- 2a c; glE3te er l" F-:: rJJq lou a^eq selllpcol 9 | i -r! ., 9 ) :: --!rl Jql sElte,!\ se 72 f,en ' m.::,.jdn aq1 11 ,{e,t'q341 \'uo lnq 'q?lfl uJelse,{\ _r'r ; r-':-r7-: pue ouuns(qduttg , i:::du.roc eq leql uoIlJaS q: :. sar:ads1o se8elqruasse ;rc:'i-aol qtoq 1€ sellqolul 5 F€.:rlfer oq^\ (€86I) llFS cs ::rq ieq areq PeJeplsuoJ oc c: *rLrl IIH azuelrtr/rrtreqJ uorr.€s tt l{BArt6rH uort.6s loarc rerpuEqc sNtvtNnor,t S ttansuv sNtvrNnon vllHclM sllssoJJaqlo (J) [t 8 tes 8 qJ] aJu!^oJd luoulluotpll l aql Jo uBlr!^opro ra,||o.I aql uI sBunBJluopouoo u; a8ueq3 971 arrtinent Province [Ch. 8 Sec.8.4l Discussion L2l (c) Other fossils the cratonal platform to the south where the a:. uouNTAtNS -: Oklahoma sectionsare located (seeFig. 8.6). The Mckenzie Hitl Formation at the sections Open ocean conditions prevailed to the consideredhere has been studied carefully by north of the Transcontinental Arch during Stitt (1983) who recorded the presence of Early Ordovician time, and a broad, thick suc- trilobites at both localities. He recognized cessionof marine limestoneswas deposited. assemblagesof speciesin the Chandler Creek The sedirnentologicalsetting within which the Section that he compared with those of the House Limestoneaccumulated has been inter- Symphysurina and Paraplethopelrrb zones of preted by Cook and Taylor (7977) as a shelf western Utah, but only the former occurs at that was prograding away from the Arch, so Highway 77. The uppermost26.8 m at High- that the House represents progressively way 77 aswell asthe lower Cool Creek at these shallower-waterdeposition with height above localitieshave not beenproductive of trilobites its base.This shallowingtrend continued dur- E in Stitt's studies.We did not find identifiable ing deposition of the Fillmore as indicated by specimensin the more restrictedstratigraphical the increasingabundance of flat-pebble con- interval that we examined, although cross- glomeratesas comparedwith the upper House sectionsof carapaceswere observed in thin and by the presenceof 'algal reefs' (Hintze sections.At present, conodonts are the only 1973) in the lower 60 m of the Fillmore. fossils known from both the Utah and he Lower Ordovician rocks aqoss the Oklahomasections. broad central platform of the United States !--o : ]: E consist largely of dolostones whose low- ..: g divdrsity faunas and sedimentary structures :.i; (e.g. oolites, mud cracks,domal stromatolites) : !!; E.4 DISCUSSION ::. ; argpe for deposition in very shallow and .L The palaeogeographicaland sedimentological perhaps, at times, hypersaline waters. The I setting of central and eastern United States Lower Ordovician Arbuckle Group of south- during Early Ordovician times was summar- ern Oklahoma is an exception to this deposi- ized by Ross(1976). According to his interpre- tional pattern. It has yielded a diverse fauna rm :.=:,:bo\e) andmetres tation, the Oklahornasections were located at unlike that in coevalrocks in contiguousareas, about 20"S latitude when the McKenzie Hill and was depostedunder normal marinecon- and Cool Creek formations were being de- ditions.The Lower Palaeozoicsetting in this re- I p;n of the Cool Creek at posited,at whichtime the Ibex areawas situated gion of the southern midcontinent has been rse his identifications of to the northwest of them and closer to the interpretedas an aulacogen(Hoffman, Dewey, c hai consistent, these low equator. Both regionswere in the wind-belt of and Burke 1974\. an aborted continental rift rreuces of conodonts of the southernTropical Easterlies.At that time that opened southeasterly(modern orienta- qxr-r. -\evertheless, such the Sauk Sea was at or near its maximum tion). Subsidencein this basin was rapid, but ar:;.a. G. quadraplicatus, extent, and the North American craton was depositionkept pacewith it, for nearly 3000 m st .\lacerodus dianae are awash except for the Canadian Shield. The of relatively shallow-water limestones were ;lt. conodonts that Mound broad but shallowSauk Sea was interrupted by depositedin this trough during Early Ordovi- c Hish*ay 77 Section. In the TranscontinentalArch, an axis of shallow cian time. The frequency and abundanceof lon rhar section, made by sea-floor and islands that extended from the oolitic, sandy, and stromatolitic beds in the tle losest occurrence of presentstate of Arizona to Minnesota.Ordovi- successionand particularly in the Cool Creek r.[:L-rl m abovethe baseof cian rocks are generallyabsent from the region Formationsuggest that the depth of water gen- | 6e n pical components of of the TranscontinentalArch, and stratigraphi- erally was less than in the Ibex area of Utah a:rel.ingly abundant and cal units commonlythin towardsit. Clearly,the during deposition of the House-Fillrnore r from higher in the forma- Arch restrictedcirculation between the ocean- sequence.The depth of water in the Southern facing shelf then located to the north of it and Oklahoma Aulacogen probably fluctuated snpoloauo rf! s.rgrlspurl ;o uorleredas pcrqder8oe8 lreql altdseq qnnoqlp 'uoqelxro.{ IIIH elzue)tl l lea{ol I :.qr sslJads ITTJoJIUOJ 'xaql le seuoz elrqollll s.azlulH qlr/A eql uI pu" euolseurl esnoH 3ql uI u^toul puE €uroqsPlo qloq F,-I 'llrH erzueyrl l eq1 ;o lred raddn eq1 tnq 1e sr salrqolrJl Jo uorsseJJnspraueS erues aqa aql rstEPuduoJ lu€ururop sepnlcur qcqlti'uorseJJns s {1 al?[aJJoc ol alq? '8ur,no1pqs 1o porred e Surrnp palrsodap E:LT AAIqI Il? UI '{IISJAAIP s"A{aq pu€'auoz .erqadoqlaldoto4nddn ue pue s€^{ araq uorlPJeprsuocJepun sr l€ql uorlJes eql rc sJeds ur€luof, Eun€d c 'euo7 urySa1souay-o4uotapag eterpeurrelul 1o ged eqr '(SgOt) llns,(q palrodar sy pa11d gL $lJor aql :nJJo lEql te 'ougtns(qduttg Jo euoz rel\ol € apnlJur ol -dns se,r lueurpes snoerecl?J rlcrq^\ l? alpJ 'se^laqs uedo Jo Eruor.lsllo ul uolsssJJnse1rqo1u1 aq1 patrodoj eql uI suouerJs^ ol pue'fllJrpouad s1r o1 ro q q!-gnou salcads I lou PrP (ggor) llus'paldues a,t leql llun reltel eql ecueprsqnsJo flrsuelur ur saSueqc o1 esuodsar esnoH eql uI T^rlsaurl'I 1o ged aqt ur punoJ uaeq tou e^?q slrssoJesaql ur u€rcrlopJo flfta eql Suunp dlpaleedar rFis :ql pu" 'lellqeq lel?A\ lL1-r lou sr [IrH erzue)Jtr l '(9161 ssou relJe) atut uercr^opro SuFnpselels palrufl uJelse^lJo ,{qdefoeSoeeled--g8 8rJ P =u?punqe JeleerE qcnu lrrElllsei\ Jo sul€lunol tr sarloLrlotr)009 00t 002 0 rc*lap edols-lBlueuquoc sellw009 00t ooz ) ={5erd aql dq uorsnp 'J Trx !ra.t\ iryu4lnq tsd+ € ^eqlpereJul sJoqlns ll!:-rp slr peleFSal ]?ql EF\ rou prp feql 1nq 'uoxe1 !.{qdeJ;uBrls PuEl?f, rqdef It aJ:-"q uloJ Jrdolouels d::rur (yg6 y) qsledeg PUB rard a^eq 01 a aIIeq aA{ Qr slueq pnul e^11 *'s-rellollPqs eql Jo alpuoqrec rJsSO ut eJuspunqe relEeJB 13 lue uolSurqlg ,{q dPnls brr-'aloul aql UI aI€J aq '-iF:-ri slqlur xeql 1" peraAoc ds =-:qf ?uroqBplo uI fly', D D62-etsuap snlnuoLlo^DlJ r :cslJa;Jlp luect;tu8ts dPo s a; an (sn1o1n3uoJ 'ruos F(I:.qB rassel;o sarcadsaql 4. {ouJas aql Jo aaJql [? trEio er€ (vsuanoltuoru 'g G :ues eql 'seaJ€ qloq >:---l!rp luopouoJ 1o urelled (LI!-rJJI SV SUJaJUOCI€lueul 4.'-b SEUnEJIUOpOUO' eAIl n*--zduroo lruled ol lelltuls E--l uelsa^l PUP etuoqsPlo uotsote qrrr usrJrAoPJO reA\O'I uo^aOalel-ald scszreduloJ pctqder8rlerls lq pa^ouraU ftr::or€pd e^oqe eqt 'qclv :qr Io sepls altsoddo uo rr= ;uus€rluoJ pu" ru{ 00LI [t 8 res g 'qC] aru!^oJd lueupuoJpll l eql Jo uBlr!^opro ra^{o'I aql q sBunBJluopouot u! aaueqC ZZI tootitr€nt Province [Ch. 8 Sec.8.4l Discussion 123 1700 km and contrastinggeographical settings the entire population in our samples from on opposite sides of the Transcontinental Oklahoma, but they are associatedwith Scan- Arch, the above palaeogeographicaland bio- dodus sp.6 of Ethington and Clark (1982) in stratigraphicalcomparisons indicate that the Utah. In both regions a more diverse popu- Lower Ordovician rocks of southern lation that includes characteristicand long- Oklahoma and western Utah are sufficiently rangingspecies of FaunaD (G. quadraplicanu, similar to permit comparisonsof their respec- E. parallelw, S. rex, Ulrichodina spp., tive conodont[aunas without major environ- Macerodusdianae) is introduced higher in the mental concerns.As reported above,the basic section.The disappearanceof FaunaC and the pattern of conodontdisfiibution is the sa.mein introduction of the low-diversity fauna that both areas. The same species (4. linedtus, replaces it occur within a very short strati- R. manitouensis\are dominant in Fauna C at graphical interval, and no mixing of the two all three of the sectionsthat we sampled,and populationsis indicatedfor any of the studied the speciesof lesserabundance (e.g. L. bran- sections.The faunal replacement in each of soni, C. angulatus)are the same as well. The the sections occurs within a stratigraphical only significant difference is the presenceof sequencethat displaysevidence for gradually Clevohamulusdensus and Chosonodina herfur- shallowingdepthofdeposition. No abruptlitho- thi in Oklahoma. These specieswere not re- logical changesare present. The hardground coveredat Ibex in this study,and were found to observedin the Chandler Creek Section just be rare in the more comprehensiveearlier abovethe highestoccurrence of FaunaC is not study by Ethington and Clark (1982). Their duplicated at the other two sections,so we greaterabundance in Oklahomamay be indica- concludethat its positionrelative to the level of tive of the shallower-waterconditions that faunal replacementis not significant.We are we believe to have prevailed there. Ethington unable to identify a physical attribute that is and R€petski(1984) interpretedC. d.ensusas a common to the thre€ sections that rve can stenotopic form based on the known geo- correlatewith the faunalchange. graphicaland stratigraphical distributions of the Comparable general successionsof cono- taxon, but they did not identify the conditions donts are known from many placesin North that regulated its distribution. The same America and from the SiberianPlatform; these authors inferred a deep-water habitat for occurrenceswere surnrnarized by Ethingtonand C. herfurthi; they were influencedin this con- Clark (1982, p.608). Owing to lack of con- clusion by the presenceof this speciesin tinuous exposuresor to broadly spacedsarn- continental-slope deposits in the Ouachita pling, stratigraphical documentation of the Mountains of west-central Arkansas. The rangesof conodontsis lacking for all of these lE Ross1976). much greater abundanceof C. herfurthi in the sections.Al[ of these other occurrencesare --er McKenzie Hill is not consistentwith a deep- consistentwith the generalizedbiostratigraphi- water habitat,and the scarcityof its occurrence cal successionof conodontsoutlined by Ething- tteen found in the part of in the House Limestone suggeststhat the ton and Clark (1971),but the presenceof a re -I :ampled. Stitt (1983) speciesdid not flourish in the depthsof water low-diversityfauna betweenthe rangelimits of b :rrL-cession in Oklahoma of open shelves. FaunasC and D is not demonstrated.Recent mae ol Symphysuina, an The rocks that occur just above the rangeof work in central Missouri (Chapman 1984) b$ta-Xenostegium Zone, FaunaC containsparse conodont faunas of low shows Fauna C to be present in the upper Htopeltis Zone, and he was diversity in all three of our sampledsections. GasconadeFormation, the impoverishedfauna i srccesion, which includes The dominantcomponents of the populationin to be present but very rare in the overlying of the McKenzie Fr Hill, both Oklahoma and Utah are largely albid, Roubidoux Formation, and typical conodonts i zones at Ibex. coniforrn species that we identify with of Fauna D to occur in the still youngerJeffer- coeraphical separation of OneotodusLindstrom. These forms represent son City Formation. Recoveriesof conodonts Dg sllssoJraqlo suounq Jo -JuqxepuneJ srql leql os '?Heqrspu? €f,ualxv 1eq1sarcads 1uel8nurur dq paceldar eraa\ pup r :qr ur pepalJal eq feur qIoN uI sa4rl"Jol snoJarrrnurrJo4 u^\oul aJP lsurlxe euEreq seesA\olPqs lnq urrEA\ur aJrl Errr3JEldar eql roJ IEuneJ sluopouoJ.;o seruanbesJulrurs 'alqElrEAp lou oq peldepe selrqolrJl'suoupuoc eseql Jepufl Surleoqs ib-nrn Jo le^Jelur sr loluo3 pcrqder?rlerlsorqesolt q8noqflv 'eurporxJeql aql /(olaq roou Pas ,l\ollsqs Jo Ee B aql ur pu€ el"4sqns 'araq^\esle searepeorq lq8norq Eulloocpqo13 yo saposrde 6req; 'uerot.oprg f,peg rrJo+ s€erp petpnls eql olur pelzr8rur a,req lPql patseS8ns(ffOt) trns q8noqtp'u^{oul (t '?tl ' UAD ;rar-u6'ocar lsnur srrJJoJra8uno,{ aq1 1eq1os 'uorlelndod lou sr uonf,urlxJJo asn?Jaql 'spor:ad uoqs ururu pqo13 le,reyzes lsaplo eqt Jo srequour eqt Suorueperytuepr Surrnirro-og Ees eql Jo seerepeoJq relo palel r srql peldrues eq louueJ szuneJ luopouoJ reEuno,{ aq1 1o -lp€ aJel{\jUql saSueqcplueuuo-rr,rue -roferu Di"urrrolleqs :ql uorlsodep aql ]o sJolseru?snorlqo ereruorq1ecrd,ft e;o g a8elg o1 sasuodsarJo ilnseJ aql se pa1e.rd.re1uruaaq m{Jeqs eql pue 'peldutes Jo JrlsrJep€J?qJaq o1srurog a1qe1s ,{qecrSoyoqd a^"q 'peurJepe.re ,{iql qcq^\ ,{q sluaruaceldar bol aql qleeueq s{3or ul -Jorrl pu? SqEuer-8uo1qcns Jo uorpnpoJlur leunq aql pue 'seuepunoq arauorg rqqr roI aJuap g Perpnls eqt pereprsuocl11s 'aJuanbaspcrqder8rlerls 'ecue-reeddesrplduqe pecueuedxe'uJnl ur r.Ep eql qJsa €es Jo l€ luecr;ru8rse q8noJql peurplur?rxsl pu€ paqsrT 'll ueqt pup 'uorleJrJrsra^rpfu€uotlnlo^e luo^\ aurrl e Sugnp F::iord ;o -qe1sesr uoqepdod pelpsre, p e-rou.re 'II"lap ur -repun€unEJ ,treu eql pecelderdaqt t€qt uorlel rn--:r luaruecelderpune; pepnls areq s1e^ro1ureq1;o sdol aq1aloq? pue -ndod aqt Jo esoql uro{ sa8eeuqd:zuoqnyo,re :ouaprae pcrSoloqtrl s1e,ra1pcrqder8qerls leqSrq p1s lV uorssaJJns lueraglp lusseldar leql sarJeds slslsuoc 'sluoPouoc Jo s,ulfs Jo I a8ets ur punoJ salrqoFl aqt qllr{\ €uneJsrql :psJnpoJlursr eunel dlrsra,rrp-,tro1e s" ( Jla Ir lo srolelnEeJ 'iql alqeredruocsr a8elqruassepeqsrre.r.odrur srqa '-req8rqse-rleurpueJ ,{eJ uzr{l eJoruou ,{luouJ p-rrlurel'arnsserd) agord " 'Pun?JluoPouoc JePIo eql Io eJueJJncJolse -ruocpue 'speqreq8rq uI 'uorsseccnspcqdel3 ire r reql f,ueru aql Suoure -qflq eqi eloqe a4aur u"qt sseluortJas qoea -peqs lu?JrJlu8rse 1o uorlrsodapaql Euynp ru: :grcads d;4uept o1 " uI paJnpoJlursr ,{lrsre,rrp,ro1 ;o eune; asrzds uorlepdod e1rqo14eql pelzuruop leql sarcads rlrml 'alsrlsqnsaql 3^oq3 V 'ar:Luorqpasuq-elrqolrn e;o Jzrrdftsu pazu yo e8elquresseue ;o uorleurrrJelprel\dn ldnrqe e pardncco srust lfruaq -3ocar (y461) llls leqt se8elspluerudole,rep eq1 dq pazuepe:eqc eJ€ sereurorg selrqolrJl ol :-?seJuaraJul(ue e4eur eql Jo lse8uno,{ pue qunoJ eqt ol spuodsar u€rJr^opJo JeA\o-Iauros lo uonnqrJlsrp €ql pue al€IPeurrelul P IEqsPq -roc uoueruoueqdsrql 'slJor ?ursolcuaaq1 ;o tardraju ol tl pesnseq (ggef) lpS pue 's{ror EF Suorlertosseqll,!\ lsPJl ,{3o1oq111ur e8ueqc3ur,{uedu.roJre u" lnoqlt,,!\ uerrqu€C ;o r(qde-r8r1er1sorqeqt ur parTdde JeleA{-A\oeqs d uoqs-reeu uorloes eql uro4 dltduqe s-rzeddesrpsluop dlepr,nueaq s€q ldecuoc eJeulorqeql uo$saJ ddns secuauncco u,tou>1 -ouoc ;o uorlelndod paqsl1qe1sa11a,{e 'eseJ -rns alrqolrJl aql Jo slusural3lu€uruoP eql ur rlladag pue uo13qq1g ,1; q33a uI sernpunoq araurorqssorJ l€ql s{JoJ se8uzqc,{-reuoqnlole-uouJo srseqeql uo selels t nuopouoc usm^opJo ;o sacuanbasruo{ selrqolrJl,(q pe,{eldsrpsurel pelrufl uJelse,{reqt Jo u?rJqrrreJreddn aql ;nrp lectqderSoaSoezPd -1edaq1 o1 serlrrelrtxrs pelJeur ,{oqs €uroqBHO urqll,t\.saJeuorq, parrrJel oq leql slrunpcrqder8 pgr u1 (9161 1ctrrcg PUe qEln ur suorlJes peJnseeur Jno ur -q?rsorq puor?er pezruSocar(9961) rarup4 s*rq?l pu? seuJeg'IL6I stuopouocJo suounqrrlsrp pcrqder34€rls eql ;) iuopouof, ;o e1,{1s-e11 'secuzJsrp3uo1 raao uorle1 ssuswolfl reelJ sI SJIflO.IIU,I HIIA{ NOSIUYdIAIOC S'8 'ruaueel8esrq lou -eJJoJroJ elq€llertou eJEfeqt terlt pue 'sluele ro ra r\o'I ,fuolsq aql uI Jo crqdorlselec ,(geeu pegar 1ou op se8ueqc Gs srqt uoseereqJ -roy asaqt leqt 'sarreJorllll ur saSueqc,{q petued -persaS8ns(Zeef ) {r"tC -ruoJJsJrB SeUspUnOqararuorq te s:6ueq: 'ruJop?ld u€srJerrJv rllJoN aql Jo seuolsolop uorlEleroJ uounlosel + IEunpJaql l?ql en8re oqr'r (9961) do4se6 Jel31A-,{rolleqspeardsepr,t aql uqlll\ uElqo d :qr ra11ool suraessnql lI pue uasSr,rpnl,(q pereJJoueeq s?quorurdo 3ur osl" €uroq?plo ur dnorg apltnqrv ra/rrol eqt qsru:ng) salels 961 Palrun -luesslpV arnl€uur pcrqderSrlerlsouorqc[1pr1 JOI pu? qslfl ul uoD?rlroc eJouruc Pue euols eql q oslE p .ral€-{-A\olpqs -uasses" pel€e! ueeq aApq'snouoJr{J?Ip leq^{ -Jrurl esnoH Jr.lt JoJ aJeq p:pod:r suotlula.t qJnssersPJ eurreul eloql s3 -auos eq ol pereprsuorq8noqlp 'serEpunoq eqt lBqt uoqueluoc eql uoddns 'Je^e^roq 'sluop eq uEr lua^e eqJ lr"lIDuapl aJeruorqleql os 'uorl?JnpJarJq Jo ueeq a^€q -ouoJ esaql'sellqecol I"JaAas uo Pes?q uoD :t{1 uI , peluaser ?!I\ lePra\ ol peunss€aJe slueAe qcns 'sluerrJuoJr^uaPloJ -cas alrsoduroc € $ peu€lqo eJe^\ f,eql qtlqd' h ruere lueuraceldar-uot1 pue redeep o1 pale8aler ueaq peq ,{paru-ro; ruorl ecuanbas eql pu" ','Aolara,n f,pn1sleq1 ur g qJ] oJul^oJd luaulluoJp!tr{ arll Jo uBlJI^opJO Jarao'I aql u! sBunBJluopouoJ u1 a8ueq3 771 htinent Province [Ch. 8 Acknowledgements 725 rr reieeated to deeper and tion-replacement event appears to be rep- Stitt (1983) recognized a Symphysurinid r- Srch eventsare assumedto resentedwidely in the Midcontinent Province. Biomere from the distribution of trilobites in d ,iuration, so that biomere The event can be identified not only in normal the Utah and Oklahoma sections that we ;i consideredto be some- marine faciessuch as those discussedhere, but studied.He reported,following Ethington and i lare been treated as essen- also in the shallow-waterdolostones of central Clark (1982), that the conodontsof Fauna C lr4rhical innature. A dissent_ United States(Furnish 1938, Chapman1984). disappearfrom the Utah section at the same ra .-dtred by Ludvigsenand It thus seemsto offer the possibilitiesfor high- stratigraphical level as the trilobites of this rLr argue that the faunal resolution correlation that Ethington and biomere, and concludedthat the same event qe toundaries are accom- Clark (1982)suggested. was responsible for the extinction of both c:: lirbofacies,that these The reasonfor this significantdevelopment groups.The presentstudy dictatesrevision of reiect nearly catastrophic in the history of Lower Ordovician conodonts that conclusion.Conodonts and trilobites occur Ir .re not reliableJor corre- han -es. is not clear. Disagreementexists regarding the together throughout the entire sequencethat life-style of conodonts (Seddon and Sweet we sampled at the Ibex locality. There, the id ci:rributionsof conodonts 1971, Bames and Fihraus 1975, Klapper and extinction of the conodontsof Fauna C occurs I =.rions in Utah and Barrick 1978). In their review of the 4.27 m btgher than that of the trilobites of the ried similaritiesto the pat- palaeogeographical distribution of Lower Symphysurinid Biomere (top of Zone C of ,iriE-\biresfrom sequencesof Ordovician conodonts in North America, Hintze 1951), so that a simple catastrophic fuere boundaries.In each Ethington and Repetski(1984) concludedthat event, such as impact of an object from an ftbed population of cono- known occurrences support recognition of extraterrestrialsource, was not responsible.It &Tptlf. from the section shallow-waternear-shore populations that con- also may be significant that the other benth- ;rmrrn_echange in lithology trast with associationsthat lived in waters of ic group recovered at Ibex, the brachiopods, ds- This phenomenoncor- intermediateand basinaldepths. They did not was not influenced as were the trilobites and h-.1 and youngest of the make any inferencesas to whether the organ- the conodonts. I-ong-rangingforms continue lEs ihar Stitt (1971) recog- isms occupied a benthic habitat or lived through the sampled sequencewithout inter- r E*rbite-based biomere.A abovethe substrate.Further, they were unable ruption. We conclude that the independent r dnenit_v is introduced in to identily specific environmental factors responsesof the conodontsand trilobites are to h a metre abovethe high- amongthe many that vary along a bathymetric dynamic environmental changesthat accom- Ite older conodont fauna. profile (pressure,temperature, salinity, turbid- panied the shallowing of the sea over broad I r-semblage is comparable ity, etc.) asregulators of the distribution of the areasof North America. The two groups did , ft{rnd iu Stag€ 1 of Stitt,s conodonts. not respond to the same factors, or did not llsher stratigraphicallevels Lithological evidence indicates that the respond at the same level of developmentof ,d Theimervals here studied faunal replacementrecorded here took place those factors, so that the conodont extinction E-r-\irredpopulation is estab- during a time of progressiveshallowing of the occurredsomewhat later than that of the trilo- lrmed through a significant sea at each of the thee sections that we bites. EL-€. Stitt consideredthe studied.Evidence for this shallowingis present ri lone-ranging and mor- in rocks beneath the lowest horizons that we frrms ro be characteristicof sampled, and the shallowing continued after hmere. Obviousancestors ACKNOWLEDGEMENTS the deposition of the youngestrocks that we ndont faunas cannot be sampled.This shallowingrnay be leadingto the Engel'swork in Utah wassupported by a grant h members of the oldest global sea-level minimum that Yall et al. from the GeologicalSociety of America and by I the rouDger forms must (1977, tig.1) recognized at the end of the the E. J. PalmerAward from the University of ! th€ studied areas from Early Ordovician. Changingconditions at the Missouri-Colurnbia.The M. G. Mehl Award substrateand in the water column durins this from UMC providedfield expensesfor Elliott's lxuatigraphical control is intervalof shoalingmust have been respoisible work in Oklahoma.Landorvners' permission to k of conodonts for the faunal replac€mentdiscussed here and collect the Oklahoma sectionsis appreciated. r€rous-quences localities in North may be reflected in the stratigraphicaldistri- We are indebted to Louis M. Ross who pre- r- so that this faunal extinc- butionsof other fossilsfrom the samesections. paredthe SEM photographsof specimensillus- '088-698 '99-g dd'p yed XX salpruSISopaD 9t tSotoiuoalDa rlrol sluopouoc roJ fitstaatuI Sunoa tuoltgug qel{l',quno3 plPI Io t lapotuuuoppes rr8oloce uV 'Il,6I J n[ 'lee^\S pue 5 lrw 'eeJe xeql eql ul suorlces clqd€134€4s 'sld Z 'tS8-1,€8 69 uua ng Druauv lo uerJr^oproJlpprhl pue rs,lo'l fl6l I I'ezlurH &atcog erue,rldsuue;'Srnqsreqrueq3 'oo 66 loctSoloag t6E rceu uo4cesu?rJraoPro ie^\o'I 8E6I f la'opues ulla n(I f,aatng1oct3o1oout7^1 puo ToctSoloag 'S0I-€1,'JJIpleJ'seler\Jo lunesnn IPUoIIPN \Dtn '\etn uralsJl\ roJ suollJJsJrt{dpJStleils pue sse.rdseleld Jo,{lrsra^run tl6l ,aquatdas pallelap UErJI^op.lore^lo'l ls6l I '-I '.zlulH 'uoq8ulu|trg'wnlsodu.ttrS uoli'tJossv lDJlgolol 'dd Z9 ' LI rloog eprnC fe^rns -uoaDpd olo sgulpanotd :uais(S unaaop)Oi aqJ IEcl8oloeC etuoqeplo DutoqDplo'sulDtunon :( pe) uesseg c w :q selels pellun urelse^r alqcnqy aqllo f,8o1oaglouo8ag'6961'A A. 'IrreH aqlur uorlelueulpesoerJr^opro 9L6I I[ f u'ssou 'Zt 'It sF 'y7E-BIE Zt tSotouoapd Io tlu 'sld 9€ ' dd I9I 9 ulqtng {,to|slH ptntDN -.rao1.delpn rddrssrssrwreddn eqt Jo spequeqC lo 1tt&asnw tpoqDad &lsoalun aF,A 'seun.el np JrJrEJdJr.ll ruoJ] sluopouoJ 8f6I IAI/tl 'qsrurnl elrqoluJ slr pue qeln urelsPequoNur uo4eturoc 'Z0l -68 96I ndos loaadg fir3 uepreg aq1;odqder8rlerl5 1961 rI f U'ssol{ oJttauv {o &atcoglocrSo1oag 'tusllona!^otd pttl 's1d7 ' dd 171 gn salrDlolgtuopouo):( pa) )IIelS T O :uI s31€1S tnwan sDrnosav pDutw puo sau!n Jo nDErnB pelrun uretse^\ pu€ F4uer uI sluopouoJ uerc o)1xal oJrxew ,reN ureqlnos pue sexeJ -r,{opro flleg }o uoqnqrrlsp crqderSoaSotqoepg I ^aN tsoruuJelsel\ (uenr,lopr6 ra,no1) dnorg 'n86I g f 'qslede)I pue T U 'uofulqrA Jo *std osed tg urorJ sluopouoJ Z86l g f 'rlsrrdeu tt "dd SEI 'z9r-6nl '7 yed 97 stlpnts [8opag &lso^lun Sunol gt f,Soloiuoapdlo lDulnol "lrurrcrqde.r8rlerlsorq u1o1,l31rgqetn'ftunoJ plelll uratse,{ 'eerv Jo purl AleuV - arauorg 'Sq6l U V 'rrruled xeql eql tuo{ sluopouoc uelcl^opro .Ippll l 'ZZI-96 6S t8qotuoa1o4lo pue re^\o-I Z86I 'I (I'IrelJ pu€ 'I d'uolSuqlA pu)nol spueql )rtJrv ueeeueJ 'eulcu,{soaSonu 'sld Z''ZB-Eq l:ZI )lolaary ocuatuylo ttalcog uerull{uerc aql uro4 sluopouoc uercr^oPro loctSoloag 'tqdotSltousotgtuopoao) uo un$od ,tllPg puE ueuqueC ele'I '996I S C 'uel^\oN -rz,{9 :( spe) uro:1s8reg'14tr 'S pue lee^\S J I[ 'Ed,g'nen-E6E tt tSotol :uI ecugruv rtruoN ur sluopouoc uerchopJo -uoa1odotct141eruoqeqo'sur€lunol l aplcnqrv re^{o'I 'It6I 'I C 'l1rplJ pue -I lI 'uotSulqlg '(uel. opro) dnorC el{cnqrv re^\ol eqt }o '28-TS SZ uoltDr!lqnAplrads ,{qderSrlerlsorqpue sruopouo3 896I U'r{'punol l stslSolD.tauln puo stsl7olotuoalol Jtutouorq to '92-ZZ 'sld f,taoos 'stuauuot^ug alouoq$J tqoftl daal '9II-86'.DInoN, b^tsptDpzl' uttottold umDqts :( spe) soug d pu€ {ooc g H :uI epE^eN pu€ uerrqure3 reddn aqt Io elozoapd Dtlo'I aqJlo ttldot8uorsolg Jo uerJr^opro reao'I aqt uo DlDe d4aN:uI elsn8unl edeuuauelpod eql ur slusruuor^ue Jleqs pue eooF Ielueuquoc uosueduoJ g pu€ 'g 'H puE orerol l re^Id '(uepr^opro re^io-I) a8els Jo 116I I,\tr'rol,{el '{ooJ 'Id IsunqJ rql JosluoPouoJ '/96I v J'o{u3le4son I ''dd €/ 'unosslll'elq '6r g-I0€ -urnloJ'erqunloJ-rrnossUN Jo dllsje^run'flsaq, 's 7 o?ultaq4y ecuewv q oN uJequouur IP^reluI N paqsl.lqnaununossu^tr Ie4uer Jo uPrcr^opro rearol eql uorJ sluopouoJ t{'l'ueuro€qJ ,{repunoquErJr^opro-uerrqrueJ eq} Jo serceJorq .6VI_EEI elrqoluJ €g6I d S 'dorsei[ pue U 'ues8hpn'I '86I ' 1-7's1d'tr99-219 97 to8 g oroqtal sprroqdoguopouoc u€rcIAoprO ullolplJots suaSwualetr D443opa9 Jo trqeq cqlueqolleu aql pu€ 'sequnuuroJ tf eql Jo ,{qder8qerlsoqltl -uqpulqtol ! 'uepa^\s €le4s uPlcr^opjo lsotu 'serur^ordSL6I'5. 'I 'sn€rqBdpuP d J'seuleg I -+rrqollll €86I H f '1185 lE4uec-qlnosJo eql uo{ s'luopouoJ 's1dgy'dd 671 197oKnS oSoupnmql r rrllz{oaC 'std 9' dd 6L Wl -r3^\ol ss6[ I{ '|.!oJlspul-l 'Ez-sItl uoqpl'i qluaq snsre,rrr8eled 1d3o1oce 'ttSopaO'oiwusul oSotlsl4o^opaFsl'ouq?noN b-.ai ?t olu o 1,!Dpl O ew oqell..[) 'I pue 'reddeDl {pnq re^lt eue-I eql }o {roJ elppru eqt Jo fF{-l\'s JlrqoluJ uel.I^opro luopouoJ 8r6M 'Icul€g c 'ss-g' uo1lor sluopouocuprcr^opro [Ireg SL6I d 'C 'p^oEreqv FFI ale'I 116l 'H f '11!15 6l plcadg sts!?oPraulw puo s7st3o1o1uoa1o4 -M-8LI St t8o1otuoa1otr -1qn4 sgJNsusJflu rnaouotz [D!?os uottbuaulpas pulltulsoac E .rqolll ueuqureS ele'I lo puD utapow i|spe) Ie^eqs pr: iurreedag IZ6I H f 'nPS rualruv H d puE rt 'lloo H U :ul EpPUEJ'3le-t a^Btq rl9 l9I USSS lnoN lnuapolv 'uollEpunod ecuerJs [Euor]eN aql luo+ pesurue'I aqt (uerc leerc uorJ elduexe rlozorelora qit"i'."uirt ;o -pue" ttt"J8 red uI peqsllq?lse seal E OgeEaJI eql uI sluoPouo. -uf,sia8 or uoiretercrteue? rreql I€6/IZ8-Uva '{q 'tr6l ue{lng pue '.t "'u!aoc"1n!'d 'ueEJJoH flllceJ eql ereld u! pal?rl r$org 996I d S 'e,{ee8res ) ".!I ',{eae( 3Wn l3 I IEIS I 8 eEuBqC 8 qJ] atu!^oJd luoupuoJpllfi eql Jo u8rf,liopro Ja,$o'I oql u! sBlmeJ fuopouot u! 9ZI lortin€nt Province [Ch. 8 References 127 qr-.r, F.. and Burke, K. 1974. Sergeeva,S. P. 1966. Biostratignphic ranges of stone(Lower Ordovician),Wichita and Arbuckle k genetrcrelation to geosyn- conodonts in the Tremadocian Series (Ordovi- Mountains, Oklahoma. Oklahoma Geological example from Great clan) of the Leningrad District- Doklady SurveyBulletin lJ4 54 pp.. 6 pls. -rozoicIL ln: R. H. Dott, Jr and Akademii Nauk SSSR167 672 674. Vail, P. R., Mitchum, R. M., Jr., and Thompson,S. 1&- t: .Uodem and. Ancient Stitt, J. H. 1971. Repeatingevolutionary pattern in III, 1977.Seismic stratigraphy and global changes *:zion. Society of Economic Late Cambrian trjlobite biomere. Journal of of sea level, Part 4: Global cycles of relative J U,aPrulogists Special Publi- Paleontology45 178-18L changesof sea level. In: C. E. Payton (ed.): Stitt, J. H. "197'7.Late Camb an and Earliest Seismic Stratigraphy - Applications to Hyd.rocar- h-\-f. J. E. 1978. Conodonr Ordovician Trilobites, Wichita Mountains Area, bon Etploration. Ameican Association of Pet- : ia ll1.5-23. Oklahotna- Oklahoma Geological Survey BuLletin roleum GeologistsMemoir 26 83 97. E- C..oodonrs-nttuc.lerutrom the lower- lA 79 pp., 6 pls. Yiira, Y. 7974. Ordovician Conodonts of the t i- of south-centla,l'Sweden. Stitt, J. H. 1983. Trilobites, Biostratigraphy,and East Baltic. Institut Geologii Akademii Nauk i Srockholni Fdrhqndlin- Lithostratigraphy of the McKenzie Hill Lime- EstonskoiSSR, 142 pp., 1,3pls. F:--.-z* !-es:..rp. S. R. 1983. Trilobite f.e:.ian- Ordovician boundary r \o!-ih ,\merica . .4 lcheringa 7 Itf-- Conodortsof the Chunsk **--r:a-o). River Moiero and 'rs---ia. In: New Data on the f t*.c Lover Paleozoics of the I l.)-elsno 'Nauka', 98-1,16, -C.-qrr..lootsand biostratigraphy dr'.-;iie Group (Ordovician), rl Ol:lahoma. Micropaleon- L i 3-.- 15- Lrre Camb an and Earlv tE= trom the Franklinian La;ar -\'ctic Islands.Journal t ts-111. l- B:o:oere - A new kind of a Ji./mal of Paleontology35 lC- Conodonts ftom El paso 6i;a-rloan) of Westernmost \3q Vexico. New Meico i. Llmeral ResourcesMemoir Itz:sraphy of the GardenCity ba.iern Utah and its Trilobite &, Peabody Mweum of ad 6 161pp., 36 pls. Lri!.rician sedimentationin the l-s- In: M. G. Bassett (ed.): :: Proceedingsof a Palaeon- 't S)aposium, Birmingham, L-nnersiw of Wales Pressand d s ales.Cardiff, 73-105. Lcer Ordoviciansection near rqhania. GeologicalSociety r at !-i7-854, 2 pts. E \\-. C. 1971.An ecologic 3- Iournal of Paleotutology45 Lithological and conodontdistributional evidence for episodesof anomalousoceanic conditions during the Silurian L. Jeppsson ABSTRACT abundant and are potentially important in studiesof episodic.rapid changesin oceanic The early Wenlock extinction event at the end conditions. Taxonomic and biostratigraphical of the Pterospathodus amorphognathoides studieshave now reacheda level where some Zone was preceded by widespreaddeposition of this potential can be realized. of atypical sediments.All platform-equipped The early Wenlock extinction event, which conodonts, most of those with ramiform ele- sever€ly affected conodonts (Aldridge 1976, ments and severalof those with coniforrn ele- Aldridge and Jeppsson 1984), has not pre- ments (Panderodus hngkawicnsis, P. recur- viously been included in discussionsof extinc- vatus, Pseudooneotodtt"s tricornis and gen. et tions and contemporaneouslithological and sp. nov.) disappeared from studied sedimen- other abiotic changes (Alvarez etal. 1980, tary sequences. However, some apparently Mclaren 1983, House,1985). However, the escaped extinction. It seems that these per- extinction in the earliest Wenlock seems to sisted,confined to unknown refuges,to reap- have coincided with a change from more pear briefly worldwide during another episode oxygenatedoceanic conditions to'greenhouse' of atypicalsedimentation in the Ludlow, at the conditions,with a final fauna before the extinc- time of development of the Polygnathoides tion that.thrived during an episode of wide- siluicus Zone and,/or during the preceding spreadabnormal sedimentary conditions. Simi- Ancoradella ploeckensis Zone- Similar lar changesseem to have accompaniedother episodes on a more minor scalehave alsobeen Silurian events. recognised. Discussionsof extinctionsand other faunal changes must be based on terminations of lineagesand not on nomenclaturalboundaries. 9.1 INTRODUCTION Thus, in any analysis,the speciesconcept used shouldbe explained;for the Silurianconodonts Of the diverseorganisms that occurredin the consideredhere a speciesembraces the whole marine realm during the Silurian Period,cono- known part of a lineage.It is thus a muchwider donts were particularly widespread and concept than that used for many other taxa. 'ru 9t S ot dn seSuer ottdtod snptboalDd ploc eqJ puelloC uo arev eslF u.tD egl ul punoJ exel = (9|) 'uotcuqxe eqr ,{cl pepeJJeunuell^erl lE Exsl : ue{I^a{ rou ds la uoe z d 5 € le pasruSocerBxBl esoqt Jo scu€rgeddEsrpJo ecuEreeddPeql Jo le^el eauaja]er eql a^oqg etuelslp eql s€^r8uunloc pueq-lq8p eql druenbe{ uI sa8ueqcroJ lde.xa PepeJJ€unillueredde (t) 'atolpn'I'pltu rFe: a_: alo0 rJoo pnasd eqt ur ro/pueJ atrls Jo atup eql l€ Sulreeddeer'acEerncco petdnualul (e) ',ftBpunoqleddn eql ol esolcro llrtrr - - l€ {lqpqord (Z) ',{Jppunoqra^\ol aqt ol esol5{lqeqord (J) :saollo} seua^6 sI ecuorniJo u,llou)llsel eql -r 'euoz aql Jo pua aql lE soqaulxo eql elerrlFsere^ool lou r=: repro ur le^rte^rosuosuoeq seq q.eordde ,{W puBIloOuro{ elep udlo (tu puP'(986I) eBP^eS,{qledEd eql '(986I '[S6I e8puptv pue prullrqey{'286I peueqolAlpux e8pgplv '186I prpIIIqeLl pue a8plrplv '916I '9L6I'nL6l'ZL6I eEVrplV) sluepnNslq pu! eBPuplV,{qsuoqecrlqnd snolre^'(€86I) rtrur€A fq tueuituns leuor8ereql uo ,{lurprupeseq 'tsoz s2ploqtDuSoqdtou'd eq! urolJ u^\ou)l Exel luopouoC-I 6 elqEl F:: -:a snpolepuod I -. : .s u DllaPolaE 'speg ,{qsr^ reddfl reaol eql ot qSnorql 'I1LZ-Z|D sldures '1 tqsp]J.I-]-te suotuorq n:. seldu"s SurpeerJnsur Jncoosapuelnl snpoqlnd ,l S, :€IJ,S-SrC eldures '1 a,rres8ueg pue I :-:-::s -soratd elqqJJltld3prunqSnoqlp '(I 6 elqsl) 'f'IOI-ILC aldures'g repous lp slsualraold'V tl_::a:r6 la^el ecuereJereql e^oqe ur 96 € l3 punoJ :puelloc uo punoJ uaaq ,t\ou e^eq oa\l ls€l eql snpolrsdo0 ueeq elpq saplor1louSotldtouur'4'yo suerutcads 'proJeJ eql tol'snt0woq ;npullJur snpol ou : ::-; snuoarJora0 lj lsaq8rqeql 'snpo.DpuDdo1 8uo1aqsueurlceds -3ot\todg, pue 'slsunpaold 'V 'slqmn^ X 3 -_ v S:l33dS eql Jo 0/691uEql eiour alJq^\ 'sp:g ere peprocsr ex€l peddrnba-uro;te1d raddn eqt ur lnoq" ol sesearJep^qsr^ ,{luo eqt u,rou>l ar? eE? l?qt sluopouoo artrs : -:arnl Druaaloi tse,rol 00€-09 Jo ,{gznper8 ,{ouenbaryaql tnq '3I red sluaur ereq^\ seer? lsour ur lnq 'eaoz oDou]lqJ'S -ele luopouo3aJouJ Jo 00t uEluoJ uolltas aql aq1 ;o sa111o1dzr8qtr^\ reqlaEol snounfs d r =:a_,; ur pasodxa€leJls lsel{ol eql Ie^al ecueJeJeJ Jo sprocer ,{o} ? ar€ ereql (986I) uossddel - DUlPouo6l F--:. i-a l eqt e^oqe ru t'S tnoq" 'speg ,{qstl raddn [q pue (1961) oue,{n pu" uossulatsroql f,q pas E = .-=::r orpunladsv eql Jo slrPlxer{l Jo as€qeql le srmco BuneJpu" -sncsrp sV ' snJqn[s 2, yo ecueserd pzardseplt 'el,{ts Surreqleaa '.8op-reurut [e1c ur aEueqc Jo erl.rDaqt o1 dpo salelelz S d oq1ot uleJaq :-::q v [3^:l aJUaleJeJe se sa^Jes'yede w gp'7 ocuaJeJad ecueleeddesrp s1r q1r,tr dol eql -tnris-a-- r6 'sJa,(?l eluoluaq luaururord ortq Jo le,r{ol pu? uox€l snoru,{uoda eqt 1o acuerzedde aql ll :-:.::xa olJrpo>iJoz0 eql ereq^r'(Vg6I'p 12urpo'tr6l pleJn"'Iees qtrar seprJuroJ as€q eql-(€86l IJIrr?g osle :urpo.DpLrnlou |]l - --: uo[€rrrroJurf,r1pco1 rol) € ua{I^erl le uollJes eas) uo4rur;ep (t96I) s.ras el11 I eteq :_ _-rDr SnpOrUJDS ^{oIIoJ ID eql ur pale8tlser'uruaeq e^€q €ler1s tnq ''Z 'tuo'd eqt sorrepunoq eql suoll rr/c =: s:poqlodso.rold le l€lep Jo Jo Surpaeconsaqt olur euoz s {l q8norqt seaueqJ -rurJep lueraJlp ueeq a^?q aJ€ql'\'z s J) 3:r-: - .1-il SflSfdS IEunEl ur t I lszelle Jossau{rtqt e ol pesodxa euoz snrun[s saprorltou?Klo4 aq1 pue ('7'uto 5r.!!-:: ^JoJ ollalalaoY sr pu? 'speg ,{qsrn ra,to1 eql 'lrun lse,{\oleql 'd) auoz saptotqlouSotldtouto snpoqtDdsof,ald F--;: ur punoJsI 'Z'uo 'd eq1-'(9961 apa11) ut 999 eql eJe s6uoz luopouoJ uPrJn[s Pa+quepl le]-: snpoqlDdsorald lnoqe sr pesodxa slueunpasueIJnUS Jo sseu ,(1epr,r.rlsour pue cllsrJepBruqc lsorrJ aql Jo o,t\I b-, :r^ -rou60luadorlsV -Ictql IElol eql €l€rls ,i(rolpn'Ipu? lJoluaA lI =--:rf,s . DllaporJal YXYJ acuanbaselalduloc e pue (9961 uossddel) t s-:-s.jolor DllapolJf I ;o (SddINOS-NIuOJIY'Id HIIAI SYNNYd =_: a.j.JU6oqlDdsoa slrzru fue,ropuel1 Jo seleur euros epnlcul lo N NVTUnTS IUSAO(NV'I'I-JSOd Z'6 rl ?l€Jls pesodxa aql ereq^{ 'pu?[loc uo sr I J-r_v--: uoqdecxaeuo '?l€rls Surpeaccnsol uousu?Jl snqlou6ocolny eqt Jo ,{pnts pelrEtep ? SuBua^erd 'pesuap +.:L::-_oIs -uoc slz 'ulD 'd e\t seeredueru q I'6 alqel '?xel luaceu lsolu uI se = : -:3pcu snpoulolslo .Z'uID'J uo pelsq erB aseql pldg eql 'sercedsI"JeAas f,q pelueserder ara,tr feql eurl |1 :::-u snqlou6outlof qc?e os pra,res ePnltur D-Ee':-:q rDq oluD>1axnl ruo+ paproJeruaaq JApqsatrads 3o roqunu uI turod le lPql 'sa8eeuq a8rel y ,{1eorqder8oe3pue (9361 a8puply eraua8 lsoru ',{pepurg spruureru ra8rel Fr:-::1i pue pff11rqel '186I prellqe4 pue eSprrply) pue sprrq lueJag loJ pJsn leql saqrzordde I (uollelr?A s-= - 51r snqlD060Pr5dV dlpcrSoloca qloq 'uonnqlrlslp eprt ,{:e,r, ldecuoc eqt pu? lecrqderSoe8 pue ll -: =- rl U 0l1Y-1d e setT saptoqlouSoqdtoruo mpoqtodsotal4 produrel alq"reprsuoc e slrqrqxa satceds qceg puB lql-rsod l( o Jd) 6 rtJl uEIJn[S aql 8ulJnp oJuep!^e Fuopnqlrlslp luopouor IBJ!8opql!'I 0gI t tbe Silurian lch.e Sec.9.2l Post-LlandoverySilurian faunaswith platform-equippedtaxa 131 has a PLATFORM.EOUIPPED SPECLES hlrn.-F,-,,ohognothoides both ecologically Apsrdognolhusruginosus Mobillord & Aldridge 2 III=rd 1981, Mabillard and t!berculot(']s wolLisel 2 d reoeraphically. A large wolmsleyi A 1dridge 2 lo 2 37.r' ; h,:r e been recorded from Tuxekonio bo.borojeonnoe Sovoge 2 d ihese are listed on Johnognoth!s hudd lei 14oshkovo 2 to5 32m t zr--e-rthe P. am. Z. is con- Distomodus nodosus Helfr ich 2 slo(Jrognoihoides ( Wolliser) 2 to 5 96m g a Jetailed study of the Aulocognoihus bullotLrs (NicollS Rexrood ) 1 (G?) dEs strata. One exception kuehnr Most ler ,I ieie rhe exposed strata Lotus (Nicoll & Rexrood ) 1 lEs of Llandovery marls \ eospo'h ognol hoduE chop '1i Sovoge 2 d 3 complete sequence of lcriodello incon si ons Aldridge 2 ? br :trata. The total thick- lcriodello sonderst Mobtllord a Alorrdge 2 Ast ropentognol hus irreguloris Mostler 1 rLl:nts exposed is about Pterospothodus omofphogno'l hoides Wollisel 2 to3 96m ITte P. am. Z. is found in pennotus (Wolliser) 2 to5 76m r L..e er Visby Beds, and is Kockelello ronuliforrnis (Wollise r ) 2 from1 55m Ess .-i at least 11 m. Faunal E z--rneinto the succeeding SPECIESWITH BLADE AND RAI'IIFORIVlOR ONLYRAI'4IFORM ELEN,IENTS re:rigated in detail at the Pterospothoduscelloni (Wolliserl 1 cornulus r3 r:e-rr locality information Cdrniodus Wolliser 2 to3 96m 'Rotundqcodino'oft'Rl dubro( Rhodes) of l'4obillord& Aldridge 2ar3 Odin ;r c/. 1984), where the 0zofkodinoexcovoto ( BronsonI l'4ehL) L G n:rent bentonite layers, gu lletensis (AldridgeI 2l L1l ts a reference level. A hodfo {NicolL&Rexrood ) 2 -. E-I:l!rg)- . weathering style, poLinclinoto(Nicoll& RexroodI 2 lo2 37m rthe Sase of the marls of the sP o 2 to4 61m Aspelundiocopensts Sovoge L rtrout 5.4m above the 2 Ligonodinocf L kentuckyensisEronson & Bronson 2l L?l G E !-rgest strata exposed in f luegeli (Wollisefl sensuAldridge,1979 2 &-n' or more conodont ele- pelilo Nicolla Rexrood 2 to 4 61m I rhe frequency gradually sp b (= Sovoge,1985,frg 10 F-1,0-S) 2 Io2 37m 5(r--1if0 in the lowest Upper Tokeenio furcoto Sovoge 2 E rore than 7 5o/o of the SPECIESW]TH ONLYCONIFORT,l b P;nderodus. The highest ELEMENTS Decoriconustrogilis {Bronson & l'4ehl) G uD iogna thoi.deshave been Dopsilodusobliquicostotus ( Bronson8 lr'lehl) t, (G) troi e the reference level ProeciPUus Borrick ;i. unidentifiable Pteros- sporsu s Borrick uur in succeeding samples Wolliserodus cufvotus (Bronson 8 BronsonI !r f pp€r Visby Beds. sonc licloiri Cooper sp to5 56m Belodello n sp o of 14iller,1978 2 Ponderodus equicostotus Rhodes l.7n rf, t on the regional { ) ftom 2 $- -::+. 1972,1974, 1975, unrcostotus { Bfonson& f4ehlI 3orl G {lndse 1983,1985), spb 2? {G) tr=-: ,_onservative,in order sp g of .leppsson,1983 3 to2 16m longkowiens is llgo & Koike) 2 io 4 61m :t(€dar,!. (2) probablyat recurvotus lRhodes ) io5 32m * Shtef and/orin the 3 -.i Pseudooneotodusbeckmonni bicorn is Drygonl Ft-i=nd columngives the aL\- tara recognised at tr icornis Drygonl 2 io 0 85m t'-::e P am Z. elsewhere Gen et sp nov 2 to 0 85m -eurJ urqt o^u qtl^\ spue pue (tt 2g'7) auoT pue slueuele rrrJoJrrrrBrpu? 3p€lq qlr^{ asoql Jo luollzJ 'd ?fua^opu€l'Jeql ur suEls lseplo aql tsoru qtra ?uop '.readdesrpquaruala urropeld '(t96I reslll"la) ?lDrls pu? {solua^\ Jo qlu,r seroads [V '(I'6 alq€I) crleruerp ,(qznbe ^{olpn'I ur 17 eqt pue fre.topuel] lsorrJeddneql urqtra\ sr e1erls Surpaeccnspua Z'uto d oql uagl{laq Ei ii: \:l se1?rrPuIutunloc aEueqo aqt plro/r{ aql ur eJer{,t\eslg lsuo$I^lp euolseull paPpeqrelur qllAr slBqs Jo sllun leune; ! Et-_:lEl; a(uos urql rnoJ are oJeql(I'6 '3rg) errlsny ur uo11a3 'peAIoAur F 6 3rg pesodxee3e ueruo,raq f,peg pue uetrnyg aurrl apnlru8€u eql uolt"rlpur u" -rcotoqlr'I-I 1e ;o Jo ;o acuenbas auolseuq pasuopuoc eql uI ,{1uose,,r.r8 leqrunu srql pue 'ra^e,troq 'lu€lsuot :-: nuaJ)Ngz go lou eJe,!\ puelloc uo uorlElueurrPes Jo sel?ll stpuo.t 6 - sNoz s Io HJvN9oHdvoNv' d 'sreaf, o1 puodsarroc plnoar.,{Sopreurur 5 sHr do sflco.IOHJI'I €'6 699 96 s-l-1t.tqua:-1 ,{EIcuI e8ueqJ eqt pue suoDcurlxa I€Jol ls4l eql _as 4)Jna Jgz uaealeq plprls Jo ur g't aqt uaql '3I l s pelsel gta JDJJTJ qcee sqcoda pue {Joluel& er{t Jl pue -a.: H LZ ^{olpn'I € 'seceld,ta; e w (7'd 'y) 'pueltoC Jo u?rJnlrg aq1 -ro3aler aEera^e aqt s€ t:ptOtJ )ez euoz olnud ollalaycoy eql seop s€ 'spirp eur?s eql s€,n ue{rAaJI 1€ uorlalueurpes Jo eler -:-JAuutl )62 € dueruur uoq?luaurpes pcrd,Qe ;o sapostde Jql Jl 'uoltrurlxa IeJol aql Jo aull eql ur peerds o qll{r eplf,uloJ seuoz qlog'{coluaiA PtxS 3 Sr AJAqI SeleJrpur asuap ecsalla )oe lsalll€e I?eJ lnq leql = pue ,{re,ropue11alaleqtlor.'Z'uto '4 aq1o1 Kem -r^a aql 'JeAa^\oH 'sauo pcrSolorq putEuo saDtpDJ ) o J€lrurrs uI apostde leune; Jo uorle er{l puo,(eq leq,'aeuos se8uer pe,r.rasqo eql _ =-! tv ^\olpn'I " Putt I ) x " ged 'Z s pelJedxe 1 -uruqnc pue reley eql srrJJoJ d e\L p;pualxe e^Eq p^r uorlpqrnlolq lpql ------'-===?il' J))ua I tl '?l€rls lueJ?fp? Jo Z uto d aql q lnq 'Irol eq ol sr lr 'sespc IF u1 snouaBrpur oq Aeru 'aqnap ht -ued\ Jlel eql urojJ lou arB ?xer Jssql Jo 1(ueu pu? Jnoloc IPurJou Jo eJe eruos lnq 'pe{Jo,ttar JO Sa^nPlar u,!\ou)I lsesolJ aql lErll sI eJnleeJ a:e [aq1 1eq1 slsaEEns suaurcads preles Jo : .raPn l HZe :elnoqred V (t86I uossddal) qcode eq1 ruor; rnoloc {r€p eql 'spag fqs4 reddn aq1;o ged ur'rou1 serceds Jaqrunu eql Ja/,(\ol eql 01ur aSuer snpou.tolsrQ ' srufl m luopouoc Jo J[€tI PUE lnoqe yo Eurlsrsuoc'dnor8 ,,ro1pn1-pru, e uJoJ -ou8out1o1'snpolasulo&'snpoqtodsotaid exel .raqlo pue esaql 'sntowotltsod.swoutlcur '-req8rqeBuer 11a,r,(eur [aq1 pue uorlcas aq1 =- a owuls s mporllouSotqlodg, u,rroul ,{1rood eqt pu€ ;o ged srqt ur alol dre^ sr ,buanbar; reqt tnq @ 'srqDuD^ D aP)pox 'aepnuoPouJl ;o set:ads 'ru I9't ruoq eldues aSreye ur punol are z 'ds 'sapror1louS(1o4pue onapon)uv e:eue8 crd,{1 Durpo\ozo pue o\gad oulpouogtT 'slsuathol t- -ouour oql sepnlcur tsg eqa 'f,1pqo13 reeddee-r -Buo1 snponpuo4 lseq8rq eq1 ecuereedde s- ). u|aqoqE 4 - paddrnbe-ru:oJt€Id eql uI u?ql reqler acuerueddeeJ e sr ,r J)sl tJl N exel '^\o1pn'I lsJrJ Jo Ia^al € '(snqlouSonpV 'E a) exel frer'opue11 aruos srql pue '(,{lrrccol roJ '116l IIeUSaas) a.rocre51 € Jo esoqt selquesa-r ruropeld aql tnq 'u.{toulun eql ur B13J1sJeplo qJnu luo4 u,{\ou)l sr 1r lnq sr ?fulsaJuesll'lue(llele ur-ro;1e1dlectdf,l e seq 'ool 'eldures srql ur 3 ds snpotapuo4 aceldet snptS?r'2 aql ur luaserd f,11artq'o\uod o1 s-readde sntolsounba snpotapuo4 'uta1s = 'euoz o aplroy ,{po '1co1ua.,q eql Jo tseJ eqt uI -tu1ou snqlouSoplsdy pue 'q 'ds ou1pouo31'7 '^\olpn'I e13l aql ' oloutlcutlo d outposlrzzg ;o suerurcedslsBI eql ]o asoql'aldur€xe JoJ 'puE rellel eq] ueel(teq sureluoJ rrr 9t Z-99'g urorl eldures y szlroar.rl ueql s€uneJ lcolued\ raEuno,{ ,{lalerpeururr 'sd qtl^\ uo4€l3ossP ul e le punoJ uaeq lou PIJS Z'xuo d aql Jo s"uneJ aql uee/t\leq ser{ lnq 'er?r sr ,nza otynuot opala4tolq 'snadde secueJeJJrpreleer8 are aJeql erueH pouad -srp Aou ds la 'uaE pue suto q Muotulxaq eql Jo lsoru ro1 pelsrsrad l?ql seSeeull Psards 's4,,(q peceldar sr su.toJrtt mpoloauoopnasd sat!loldDJI -apL,Auel lnoqe f,q pezrJelJ?JeqJ arp s€un€J ereqrn 'Ie^al aJuaJaJeJ eql e^oq? v6tlDJlsotg luopouoJ uer:npg SurpeecJns'pepeJJ€un ru 0l I pue 06 0 uaa,nlaqrncco saBuEqJIeunBJ ere 'Z'tuD ';r eq1 ur sercadseql Jo 0/6SZlnoq€ lsrJ erll 'I 6 elqel uo pel€tlPur sI uopJes ,{pg'sluaruela IrIJoIuoc fpo q1r,,.rpra,ras € ue{r^arl aql ur 3x€1 Jo uo9nqulsp eql rI [€ 6 ces 6'rtcl uegn11saql 8ulrnp aJuep$a luuollnqlrplp fuopouoJ ptrB lsJFoloqlla ztl rlE Silurian ! lch. e Sec.9.3l Lithologies of t\e P. amorphognathoides Tnne I.].' o-r:orm elements. Only ;t';:: in the P. am. Z. are :&-: Silurian conodont Biosfrafig raph y rz:.i br about ten wide- 6rapfolifes [onodonts Ce llon Gotland a r€--:isled for most of the R greater differences Lst. --e .: the P, am. Z. and Marl F- \\'enlock faunas than G- :or example, those of SUNDRE : B:nlock, only Kockelella HAMRA B n: -:; the C. rigidus Zone, BURGSVIK E ilement. Its ancestry is EKE lel,-.:l resembles those of 3 rz =.t top Aulacognathus). ] H . :-:nbrm-equipped taxa E H.fr ifs chi linearis B bohenicus tenuis E ILe .::r includes the mono- J E4"j and Polygnathoitles, A.ploeckensis K.variabilrs i=t. Kockelella varirbilis, @ b.-:rrr'Spar,ognathod.us S chinoera NW r-. These and other taxa ' c--up. consisting of about [ !--aiodont species known ul t's-...n 1984). A particular KLIIITE_ 32H. I udensis BERG d-'r-.t known relatives of r-: ::.-rttiom the late Wen- L M dorthali MUL:y o1. Z. or adjacent strata. I u 6.nassa / H ALLA rte ater part and culmin- J 31( lundgreni S Siltst hu::: episode in a similar . ir :ie late Llandovery and C.radians L Bc.i zones coincide with 30 t. e lle soe o I in many 29 (. linnarssonr -.-dimentation = K. absidala T : \.:tkelella patula Zone 2aC.rigidus K pof ula b=s. 27l,,l ' riccarton en sis T OFTA 0 s rhenana 26C.nurthisoni HOGKLINT (.centrifuqus K.ran u lif ornis S OF THE 124 L.VISBY IEOIDES ZONE S. grandis Panorphognothoi P.(e![ont ll'nistone sequence of 25ll crenulata Drronran age exposed at g.1-Lrthological L- 9.1 ) rhere are four thin Fig. sectionsof the Cellon and Gotland sequences,related to chronostratigraphical divisions,some graptolite zonesand some conodont ,G'in I :nterbedded limestone rones and faunas.The symbol the-Gotland column indicateslevels where graptolites have been found. The marlson Gotland includeboth distal marls t Landovery and the 21 m and 'lagoonal'marts (e.9. the Eke marl). bg strata(Walliser 1964). the Llandovery P. celloni rDds \r'ith two thin lime- 'seuquelorq ,{q peteceur u pue fureg,{q peddBruse 'selSoloqtlluBImlIs 1tr-,.j1 696I p"orxe)I puP '(6t61) locnog punorBlreq'lpruroueqlJo uollnqulsrpeqJ saryntsluopouoc Iq peqsqqelseueaq s€q lI6reqn,{Juo pa1:eur "-: eql p€elsq uoll 7 d srsntprq v ecIreI[V q oN ura$eaq euoz raPtotpouSoqfuouto'4 agl Jo uollnqljlslP uir\ou{eql Z 6 3'C t=:: $ol sEA\31 rad sluaur rlE:-:r Jql leql lou 'uotl? !-:: :alEer8 e ,(q peseeJJut r sr::lJrqt eqt l€ql suopec Ef, 4=.\la reAaA\oH'scllssJc r i: :tBuoqJPt eql Jo uop r :F;! snoelesl€J Pue ale req tuor] e8u€qt :l= -:"uoqr€f,:qr qlIA\ 11a,t seer8e se:rrqr tEer8 oqJ, '(69 'd FE:\aU '696I lqSFX) r paxturq8norql -=.'loqrrl::luulqt 'etJnos uJelsee l,:=:-.:-. Jtszlc P :suolllpuoS r :-r- ?\eq saSueqJ eseql E:--ro eql Jo seelP leu , {::ilrur-I eql lnoqSnolql steqs to sarlau ouos E 1r.: salEuoqJ"J aql Jo EE :.=.\ JuSslJ lPql SlealeJ lrtql sertsu Iueyl alPqs lrll€l V f, f-: io ;uIJ€Il I?rale'I li-:: ilorlN PuP Psorxed) '' ** ":l:1""':,":"ilo q !:-.r-\) sseulJlql relnnls T,-O ueqlnos pu? ,{IJnl snle'H = +. :-1rsa \\ '(S16I PreIrIU r :ll. d '016I lotnog .tr0otop snoaEttr6rEto ,or r lF rr:1{ pue peofied :a[€qs auoz s.plot4}eu6oq.houe'd at4! )t ::,\ lErtue3 ur ,(lqeqord lI :r?iir€W Pue erut8lt4 kv.rr1 qrn5'fi ?lt'zttt u r-'Jr tu sz uBqleJou eq '1(Oleq dr ::: a \oqe pue 'uI ei :-r,oaJ€lllSre'luauuuoJ -,tasse f,1a8re1pedola^ep sI Z 'luo d aql'fuau :reqruatrAlslI?C sacud) suroqeHo uJel{lnos (9361 Surpnlcur AIS) ?aJ€ePI.t\ e ra,ro paztuSo ry- -:., illeur8reru ero1a1 -quoc aql uo,{1prluac a:oru '-ra,re,r.ro11 '(Z'O xoppPl^ -JeJUeeq s"q e^oqe pu€ ,!\oleq selsuoqlsJ lL^ _ ie ta saur€g :requatr{ lJurEg :rJqu:w Jo uPd Ieseq) lI ro ru ,{lecrdf1 '1tun 5O tn elnsurusd ecnrg eesseuuellerluec Pue'(696I peoffeu pu€ qtrr'r 'sseu{cql ur ssel 1 pooSsg pue requel snoecellr?:epJsuepuoJ {13uolts e sepadoJe,rep lru :u€ peoJxell :Jequetr l IIoJTN:reqlue4 -re,no1 l qt:o1r1 st.redu1 :.-:.-'O ere8€rNeqt '(986I 4eel3 ea1),{>1cn1ua;4luecelpe pue euerpul st'Z'utD d eql EJllaulv 1o lf 'uolleJ eql (sauoz r+':d. ) olt{o uJels"equou urals€eqtnos '(ZS'd 'gt1t .raddely pue l€ euoz eql Jo do1 le puPplerJrallBs elqolder8 g) snlerqroleur € sI eraql leqt tueP =:.']C PuE PsoJxaU :elml Irureg'916I '9461uosdruoql plelllqel rr-: zu?lpul ulelseldquou :reqruel4lapqg xrs-ftua,reg)ulseg srouIII eql -r,resr 11(9961 e8prrply pue l '916I a8prrply) arp olut saqcear dlaruq - Pue aqal'I Jo ollueJ Jql ol lrnossll u:etseaqlnos'(Z9d IJolued\ t -_.iI Peorxell 'z Ec.:-Erg) slouqll urels€e '9161raddepl pu€{crrreg'696I SIerJ:euols ulD'd er{l aruls (9461ra8ee1) sa111o1dzr3 ') 'Z d '; :uoz eqt ul{rol sJnoJo -errIrl rr"lJ 19 8ur,(1.teaoJo ru € 0 pseq aql (82 auoz) auo2 snp8lt sepnPu| X t 'Z'u.to'd SulPaarcns y -s:rLtxolop ,{preuJ 'set€ pue ,e1eqsuo11ng,) sesuel:y ureqllou '(086I 8ur,(y:e,roaql (ru €I'I) '/"J, dol eq1 Sunl-ro1(ru spaq euols t:!r lalaur (91 o1 dn) lzra uapsuv '9761 rcddz11 puB {Jlrreg eqt ;o 17'g) pue prlBoloq1r1 rI [€'6'cas 6'rtcl uBJrn[S aql 8u1.rnparuaplta leuopnqlrlslp luopouoJ NEI of ttLeP. amorphognathoides Zone 135 b Silurian [Ch. 9 Sec.9.3l Lithologies carbon- Thompson 1973, L975, Barrick and Klapper r 19?6, Amsden etal. eral (up to 10) metres of argillaceous horizon 1976,Amsden er a/. 1980,Barrick 1983), often le-r ('Button Shale'and ates, rnainly dolomites. A thin shale north- richer in conodont elementsthan the succeed- lerlring St Clair Lime- occurslow in the zone in, for example, Member; ing strata (e.g. Nicoll and Rexroad 1968, p 5, nick and Klapper 1976, eastern Illinois (Brandon Bridge include Liebe and Rexroad 1977, text-fig.3), and con- kouri to the centre of Liebe and Rexroad 1977). Other areas tains glauconjte and phosphate (Craig 1969, Member; northwestern Indiana (low Salamonie Dolo- Shale Sattedield and Rexroad and Droste 1982), easternand Rexroad and Nicoll 1972, Foo-nn-six 1973, 1975, Barrick mite; ('Packer Shell'; Kleffner Thompson 1975, Rexroad and Droste 1982, t p.62), southeastern northeastern Ohio Dolomite Banick 1983), minerals which are usually I Lentucky (Lee Creek 1985), the NiagaraGorge (Rockway and the interpretedas indicating a low rate of sedimen- Osgood Member; Nicoll Member; Rexroad and Rickard 1965), in Ontario (St Edmund tado;. Further, the change is not regional, , and central Tennessee Bruce Peninsula but global. An alternative model is that the ftr Vember; Barrick Member: Barneset al. 1978)' American regional changes in proportion between E !-ntrally on the conti- More marginallyon the North carbonatesand clasticsoriginated in a strongly iftseloped largely as sev- continent, argillaceous sediments domrnate rate of calciumcarbonate deposition' below,in, andabove the P. am.Z,wh\chmay lowered (Fig. 9.3), the P' am. Z has been be more than 25 m thick (Rexroad and Nicoll Elsewhere identified from the CanadianNorth-west Ter- ritodes (Chatterton and Perry 1983)' Yukon (M. J. Orchard, pers.comm.), southeastern British Columbia (Norford 1976, Uyeno and Barnes 1983, p. 10), southeastern Alaska (Ovenshineand Webster 1969, 1970, Savage Rickard 1975). West of these,in easternKen- (Miller 1976, 1978), central tucky and southern Ohio, the Estill Shale of i98+), California (Seddon1970), Anticosti Island (Uyeno similar thickness occurs between carbonates Texas Gasp6Peninsula (Now- (Rexroad and Nicoll 1972). and Barnes 1983), the northwesternGreenland (Arm- Lateral tracing of the stratigraphicalunits lan 1981), and press).In Europe it is known from reveals that clastic sedimentsformed, instead strong, in and the Welsh Borderland (Aldridge of the carbonates notmal for the area, Wales 7975, Aldridge and Mabillard 1981' throughoutthe LimestoneSuite and in margi- 1.972, and Aldridge 1982, 1983' 1985), nal aieas of the Dolomite Suite (Fig' 9'2)' Mabillard Norway (Aldridge 1974, Aldridge These changeshave been ascribedto regional southern Mohamed 1982), Gotland (Jeppsson conditions: a clastictongue spreadingfrom an and easternsource, thinning westwards and grading Spasov1966). From Asia very few conodontrecords of this increasedinput of \r-..-) tion of the carbonateby an clastics.However, elsewherethere are no indi- cationsthat the thicknessol the P' am' Z' was increasedby a greaterrate of clasticsediment- ation, nor that the number of conodont ele- mentsper kg was lowered becauseof the dilu- h.- \menca. A hiatusis tion. I;stead lhe P am. Z' is very thin (Nicoll {ttrr rormal- background and ats :.oken lines. and Rexroad 1969, Craig 1969' Satteriield pu" auolsau4l pazqrurolop ,{q umpa,ro s1 pue \_z-\__,2,-\__z-\__z-\_-,2-\_,- f,.Jelqe 'aFIIS proJsllld 'punoJ Ilaaq seq serjods Fuoz FF eql oreq^\ serlrlerol uo Altro peseq'e'doz saploqrDuSoqdtotuo aql uopnqr4srp u \ou)-€ 8rd r -Eilsrrc 'ue8rqcr1,,qu1 d Jo leqolS 6 '(0,6I rornog 6 $orsaurtl aql q '(116I I FosAAi gS ut epqs pue lr -iq patuasardalsI auoz ,lfrs'z'wo d eql lIJ El?p t -d rqr ueql uaou{-lla^l +rut- quoN urels€euI 51 uossddal)Eere uJelsee tn]lsarntl puE 'eele euols r qJetn 'lsa^\ srIl ol slJ?rrr 7.qqtrm) salzqs -ntaqoq-z't d 'puElloc uo "ql lI qnelugqJs)sdyg crure3 rF pra^as 1e padole,rep Z rt{I (t96I resul"l[ + qrlr i(lpexa saplcuros I :gorda elPqsqlrnoJ eqJ, !\UZ SNJIUNTIS 'd lJO S: CO'IOHII'I ''6 lt 6 3as 6'rtJl u"Irnls eql 3ulrnp aruop!^a lBuollnqglslp tuopouoJ prrB prfo1oqll1 9EI ith Silurian lch.9 Sec.9.4l Litholoeies of lhe P. siluricus Tnne L37 9.4 LITHOLOGIES OF TIIE SalinaFormation, typically about 0.3 m thick, P. SILARICUS ZONE showsconditions of slow sedimentation(Sand- ford 1.972)and from the meagrepalaeontolog- The fourth shale episodein Cellon (Fig.9.1) ical data may be referable to the P. s.2., coincides exactly with the P. s. Z. (2.83 m; (Berry and Boucot 1970, pp.779, 220; com- Walliser 1964). The P. s. Z. is similarly pare Rickard 1975). developed at several other localities in the In southeasternCornwallis Island in the CarnicAlps (Schiinlaub1970, 1980). Canadian Arctic a 40 m thick tongue of On Gotland, the P. s. Z. includesgraptolitic dolomitic siltstone and limestone, with shales (with M. bohemiats; Hede 7942) and M. bohemirus tenuis, is intercalated in dolo- marls to the west, marls in the marginal lime- mites and limestonesin the lower part of the r- stone area, and limestonesin the most north- P. s. 2., which is ar least207 m thick (Thor- easternarea (Jeppsson1983). steinsson1981, p. 5, Thorsteinssonand Uyeno In easternNorth America the P. s. Z. is less 1981,fig. 16). well-known than the P. am.z., brt available In oceanic areas in the Palaeotethys,too, data fit the P. am. Z. model For example,the atypical sediments interrupt the graptolite zone is representedby argillaceouslimestone shale sequence (Jaeger 1959, 1976, 7977, and shale in SE Missouri (Rexroad and Craig 1978). Thin intercalations of normat shale V 1971), in the LimestoneSuite area (Berry and produce the only graptolites found in these Boucot 1970). units. In the deepest areas, e.g. Sachsen |i :. where In Michigan, Ontario, and New York, the (Jaeger 1977) and the Carnic Atps (Jaeger -\aliries the Pittsford Shale,a black shaleat the baseof the 7976, 1977), about 5 m of grey-greenshale t 19!+). The singlesarnple rs-1 qas from immediately rtercalated in a limestone frua Lower or Middle rruan (Igo and Koike I cellon &< P. am. Z. makesup at I -<-'n. of an unchanged }{-er 1978),to date the rgJ.-\rted. e r';ns of Greenland,car- r -=L.€d with the P. cel- re r'r press,Hurst 1984). Etrlrn is that before the : r.:e of depositionbarely Ockerkalk = ..r subsidence,and that I rb:idence during the Thuriogian F acios ,i^1", girr the seabed below the rhtjrinsen --^\-\-.""6: Bre Some s. s.g.-dimentation. in the Welsh Bor- ri A,Idridge1985), seem Fig. g.4._Schematicdiagram of sedimentationbefore, during and after the p. amorphognathoidesa^d an:d model. P. srTrricls chrons.Some of the areasdiscussed in the text are marked.Thicknesses aie not to a uniform scale,but relative thicknessesat eachpoint indicatechanges in the rate of sedimentation.Distance below sealevel is not to scale, Eq[P $) [lPtIIs 1r ruo{ 'suonrpuoc pe1eue3,{xoaroru ;o leds yer-rq pue sa111o1der8yo sarcadsJo reqrrnu aql uI E aqt 'JeAe,t\oH rl\olPn-I e slusse.rderdlqeqord (euoz ruo ard eq! suorlenpnu 'seAuEqJ[a^el €esJo ecuapr^erllln\ qf paJeplsuoc ecueserdaqa !_.l:laq uol go 1p ro ged pue)'7'tuo d aql'snql sauoz req1e3o1urelr:g ur epqs IrBIq Jo F resunlr pu€ pJlql eql aerql Surpeeccnseql pue .Pu?lloc {co1re1\ USHIO 5'6 Surperardeql ur ueql (auoy stlouds^Jelopuel-I'S aqt 8uI SSCNVHJ CIIOI{Y !crde-6- aslNeqlo : seuols -lerodrocur aSesnrreql ur) auoT otolnuan'W 'a[x4 : Fr(Ie leql le PUEI aql ur sepqs :1ce1q1o fcuanba.r;Jeelol I{Jn(u P r a[eJeq uopol.td snldD,t3 srereql teql (I'aIJ'I86I)' p p lJa??a1 'euoz ^\oqs b1t parrageun peurcrxeJ 'euoz leql ulqlll{r Ila,t\ sI l?ql Jo Pue eql J3u3 IIoa penulluoJ ll ellq,{\ E aqt aJor{sJJo eJorrr (9961 pue11o11)peugep ,(11uarrncse lcope,,1\ 'eraq,$,fue^a'Z s d er+!o esEqaql ot esolJsl ! Jfil-aroqs eql 01 rasolc eql Jo esEqeql oslP,{qsreqt pue ''z'tuD 'd erl! aposrdaeleuoqrec peerdsapr,tr srql yo u?ls eql p>Jo1a-rap r are speq eql ;o do1 eqt tzql pue (696I re11e;nsues) auo2 'lrun euolsorrrqeql olur tx S se8uErpu€ elEqs 'e^oge 'Z'tuv 'd @ u1 nlnnds 'g aql urqlrrr\ puno} eJ? seuoz saprcqpu Jo eJleu reddn aql urqtr^\ str"ls 'Z s d eql I Ffr-fur aql IJ ' Z 'd 'X -Sotqd"touto'd puB ruollal 'd aql qtoq teql etEc '(9961 ''ururoc 'srad 'ra8ael 'g) saluedsotq I uE3 mqt qJrq^\ 'puBnoo -Ipur pu€lloc uo eros I reN eql q sluopouoJ eql Jo esEqeql ,lroleq ur € lnoqe eJueJJncco bauor pue I alIS (€86I pue ('ruruoc 'sred 'punl?o{S 'g) salqolder8 lsaq8rqs1r ser4 slnau! ?qrstf4 'W '(Og6t qnet ni ipqdqs ur vtvprsqD X ;o se8uer paqslqndun's;qr qll/r\ tueueer8e -uoqrg aas) snuat snJnuaqoq'J,Vsuleluot llun ;l5 uI sJnrJo uas! o$'x al 'avoz s11ot1ds'geq1 1o gud raddn eql srql .4\oleqet?qs Jo tu t PUI] eqJ '(086t qnel alPP[u erll ur uasu FF -Io Jo e^qerlpu ?un€ge1y1o1dzr3 e qlp\ pel€bossu -uoqcg) seluedsolqJo trun ur tI e JoJldecxe 'y '(y pq'2, 3g '9961 rlcu 'Z'wo -d Jql Jo sluopouor payodar a,req aFqs qll/r\ paleleJlelul aJe Speq euolseulq rT eqr Jo Jluq Je.t\ol eql ul (9961) saureg pue oue,{11pue (9161) proyog 'fuen6 B{^ollntrAlaqt q (086I qn€Foqcs fl t-Sl uretsea eql uI prre :(016I qnppoqcs pue ra8ae1)euoz luop 't96I JoSIIIEII)alqEle^? are sel"p luopouoc f I I I3:-TIIEA\ (Z'd 'X) -ouoJ tuolP)'d Jql ur punoJu33q e^Er.l (13^al '€rueqog ruo.rg (4461 '6961 reEael) sluaur ^C l| * plqu3sep s€,r\ p^ s11ot1ds'g e ,ro1eq rnq) esoz Dplnuan'W -rpes 1ecrd,$ueql uqll/$ suort?leoralulepqs rlE __'0) trun epqs e qth\ eqt ur ya,re1qSrq e Surlecrputsalqolderg suou urnlE lsa/r\ol 3ql ul PunoJ st suD)utl tq)slt4 ln,pe1d an-l1e szq qJrq^\ -euoz luopouoJ pue a111o1dzr8eql uaa^{teq 'yt1'(976y n?aefi euoz onDultlr '71aq1;o do1 p :ooereadde JerJq aql suort?lerroJpaqsllqnd ,t\al e are areql 'paceld eql pue es€qeql uee,ru3qfr?A slueurPesIesI 'paJrnJco aq pFoqs,{repunoq lcolueiA/dra^opu?l'I -dfie eq1.uo1eq seune; alrlolder8 lsa8uno,{ aqa I c-Je i€u stue^e 'eltqns aql euozorqe111o1der8 qJrq,tt ur urcueJun snql '(9961 p1a;ne1 pue uossddaf) salr1o1der3ou -rIIxD [ uee^\leq I?^Jelur q tI (€I'd'91,61 'pp sesseg)1co1ue16 eq1 tnq sEun?Jrlrlr IIll,$ pue 'auolspuespu€ euols ' E]t{lq lo J3{"e,t\ reqlla ;o eseqaq1 ro; ,{1qeco1ed,$ eql rxo{ peqrrrsep -aurl 'seuolstlrspue seuo$pnu snoeJeJI?JsB Fnu"{rmJop lleAl ssel 'oA{1 ueeqte,( se e^?q salllold?J8ou tnq 'selllold€j8 qrns'sar8oloqlll snorJel Jo dnorg esgsrue; 'a^oq? qrpPE uI pessnt [q petep eft sel€qs ic?Iq eql se 'uleuetsE -pa^o eqt ol d€^\ saarEepqs elrToldeJ8'ueP J .xrr muaprla IzJrEoloqll ol llncrJJrp sI lue^e Irolue^\,{1ree[re,rop -e,{s lsouruJerllnosuI'auE{s uI '(9161 rcEaefi are areql E luareJJrp -u"l'I elEl eql Jo Eurrurl esrcard aq1 tsse eql ur ersv 01 lse,{ aq1ur eSrtqaSrelarqcS '(u?rlsJoc eqJsruraqll aql puE otcoJoli{ txo+ 'punoJ pu? fer8 seare(pogs) A\-- If,O'INS,ry\ gHJ NI fpee eql ur paceydst epostdaeql eral{,t\'086I sr {pluereJoquo {cBIq {Feg-uerrnpS I flflSsod USHJO 9'6 11eEBe1areduroc tnq) 'Z s d eql Jo le^el rel{o eqsuI lp^ralul u"Iuolaq eql re '(I '39 'fgOt 'p p rra33a1)uelproJpn'I lsarlJ€eeql ur sluaulpes pcrdf,lz qll,| aposrde 'd ,tlrea eql ur aposlda{ea^r'Jeuq e JoJ ldecxa ,{po eql srt1 acuenbes ;o sediftqloq ul'(gLz euoz $suauouDJJllw atp e^Oq? uElrrg '9761 n?eel) ls€e aqt uI Bple^olsoqrezCPu? ur epqs {relq ou sr erarll pe1zue3,{xoarour quou eqt uI uesgJespue uaEurrnql ol 'qlnos FfouoJ ur pu€ seltsoloqll ol',esnoqueer?,'luzu8elsruog paEueqcsuourp aql q uIuIpJBSpue (zuolecrzg reau) uredS atal uorpeuuoc 5ql Jo -uor u?ero '{Jolued\ eql eseq eql ol esolc ruo{ ue ur acuenbesrellurts e sldnrrelur Jo "eft ;e u dlaq 1ouop snql pue teqt etecrpur'(796I uoss?uofPu€ a\ollaJpooo) II€IJe{JO Jo rx 0€-0I 'setreg uetSutrnql prd ssal a^eq suoBrpuoc uo{n^ ul spue4 JelnJessr€g pue '(186I lr ra eqr uI (t 6 3;g) e1p,q pue eleqs tunp cB rcnrs1p peqsqqnd reqlg pe88rl) elep punzJ reqlo pue'spodolqrerq -qo1der3pasuapuoc Jo etuenbeseq1 sldnrralut [r'6'ces 6 q3l ue;.rn1;Seqt 8ulrnp aJuap!^e1euollnqlJlslp luopouor pue papoloq;1 8€I t Silurian [Ch. 9 Sec.9.7l Conclusions h launal data (I-eggett Other publisheddiscussions of Silurianoceanic present it cannot be related with any confi- 3 secular trends in Yukon conditions have less precise time resolution dence to any lithological or faunal episode con 1984),indicate that and thus do not help in establishingthe nature elsewhere. t \l enlock, ocean con- of the connection between variations in Ergnart,'greenhouse', to litholosies and in conodont faunas. ke is no black shale in lI. riccartonensis Zone 9.7 CONCLUSIONS episode in the early -at ia ol. 1987, fig. 1), at the Four similar episodesof unusualoceanic con- (but !. compare Leggett 9.6 OTHER POSSIBLESIMILAR EVENTS ditions are registeredduring the interval from placed IN THE WENLOCK AND LUDLOW rft is in the early the late Llandovery to the end of the Ludlow. They differ strongly in magnitude,the oldest of the late Llan- There are different kinds of faunal and and the youngest being the best developed, :l event is difficult to lithological evidence for the two events dis- with the most distinct faunas,and with the best - E :hales are dated by cussedabove. In addition there are probably documentedand probably also the most pro- Taolires haYeas yet been two, lesswell documented,and thus probably nounced lithological deviations.The fact that locality for F the base of either weaker or briefer events in the time conodont lineagesthat survived to reappear actol.1975, p. 13). It is interval between them. Further, still more during the youngest episode studied did not b graptolite biozonethe subtle, events rnay also be expected to have appearduring the precedingtwo episodessug- ( boundary should be occurred. geststhat these were of a smaller magnitude. pubtished Br correlations The brief appearanceof Kockelellapan^rla, The much lower number of platform-equipped h a-ud conodont zona- which has a true platform element, coincides and other taxa during the youngest episode bdng a high level in the with a shaleunit (0.77 m) in Cellon.This inter- comparedwith the oldestdoes not in itselfindi- (lm below a S. spiralis val was described as the K. patula Zone catethatthe youngestepisode was weaker, since d m the P. celloni Coto- (K. p z.) by Walliser (1964). not all taxa are likely to have managed to d Schonlaub1970); and In the easternUSA, K. patula is found high escapeextinction and to hold on in refuges Llyeno and Bames (1983) in the lower half of tlte K. amsdeniZone (Bat until favourable oceanic conditions recom- fros of the P. am. Z. rick 1983, fig. 4). K. absid.atareplaces K. wal- menced. However, most faunas representing pine fauna indicativeof Iiseri in the middle of that zone. On Gotland the youngestepisode are dominated by long- lb S. spiralis Zone. In K. walliseri occurs in Slite Beds unit f and ranging taxa, with taxa of the 'mid-Ludlow- g uapublished ranges of K- absidatain slightly younger beds (Jeppsson group' numerically subordinate; thus the fnd pers. comm.) and 1983). Slite f and contemporaneousstrata on youngest episode also deviated in some way i I core on Gotland indi- Gotland,which thus can be correlatedwith the from the oldest one documented here. In dloni and P. amorphog- K. p . 2., fit the model developed for the common to the oldest and youngestepisodes qrithin bod the S. rpild/rs P. am, Z. above. In marginal limestone areas are the facts that they both include more than l!X9) and that the top of the beds are developedas a thin, marly unit; one conodont zone, and that most taxa have rrebr also the baseof the closer to the shore-linelimestone occurs,and locally varying, overlapping ranges. In com- t defined (Holland 1980) more offshore the marls seem to have mon to all four is the reduction in carbonate E. remained unaffected(Jeppsson 7983). Mono- sedimentationin areas where limestone was L fig. 1) showthat there is graptuspriodon becarnewidespread on Got- otherwise formed (Fig.9.4). During the two E of black shalesin the land at that time, and occurs even in lime- strongest episodes the resulting starved i rheir usageincorporat- stones;otherwise graptotoids are very rare on sedimentation usually began contemporan- than in the preceding Gotland. eouslywith the final conodontzone; however, -)rceding three Wenlock The third and thinnest shaleeoisode in Cel- locally it started during the preceding zone. st. Z. (and part or all of lon consideredhere is closeto the baseof the There is a very stronglocal aspectin the over- N probably r€presents a Ludlow. However, the conodontfauna studied lap of the taxa of the zones;this may indicate : or\ senated conditions. from it is small (Walliser 1964), and at the that the appearanceof the zone fossil of the ,(IUIeuraJ€ seJuaJeJ Fq--Tqndun Pue paqsllqnd aJrnbal suoDculsuoJer-snlsndd?'fleleunl plp asnoH :a,rge1ardre1u1 | -ose-i:t Iuol pu" 'pr?q3ro -roJun 'punoJS ?urureS ,{1,no1ssr ool 1?ql pu€ -Jrpeql sluaurrpassnoplxoue Jo acuas:rdeqt aql t -C iaupog ',{qdrn11 y ',{ruouoxpl ur sluerrole urJoJleld-uou SuIAJoAa ur puE ?xel Jo Jequrnu e ;o acuzreaddzsrp l!! -uouetl€qJ g '(I uelra ,(1,uo1sarour eql Jo esn sllured sasnl€rudde ul (936I) asno11,{q petueu.sluaAe,u?ruoaeq ',(UaSOluO alquaseJ saposrde aseq; fun1rur 'aldoed;o taqrunu JO UOrprulSUmeU UO UOn"IIITOJUT asoql Jo eruos E! Suorssnrsrpq .punl JaqunJ a^rF 'elq"I?^e uaq^{'seues qulorc '1apoupaugar e 8ut lrq O 11aeqcr141'ra8ael 'uoque e Eursearcur paurc3 ,(1,tro1sa,req qrru,{ -aarqreur lusuodur aq o1d1a4l aJeuoqnloser ra- -5 p:emog fq ualrE s?,rr 'sernpnrls IeuJalur Jo edeqs eql uI pe^Jas aurrlq8rq alqeredruoJqlra{ selpnls IsJruraqcoa8 lF r-UUOrsSruJed 'JeAeIAeJ -erd sr slueurelg tuopouoc aql Jo ,{ueaoluo qluri paulquoJ sJlpnls luopouoJ arntnC aseqd 'sernlcnJls pel{J"al ! rr"o-lxall 'g IftC 'uoss aql Jo qcnu ,(leleungog aall Eqleralacce-gas Jo ploqseJql e ,e8pupl\/ " EtE:*-:lrg W 8nS -e,{resuoJ uo pue fueSoluo uo psseq ualJo aJe sa?ueqr pnpzr? eql lEql seleclpq seposlda i gueuruoJ ,(q pa,rordur sdnorE raqto ur sdnlsuoqeleJ r{Jns Jo sepnls rolsur oau er{l Jo pue eql le a?u?qJ IeunzJ rsE 3qr 1o ,Qlenb eqa 'pessncsrPex4 erlt Jo lsolx JoJ psqqlqel$ eq prder ,{re,,r.eql 'reaa^\oq :lapoul elels-o,{l ol uletuar sdqsuorlsleJ luu puess3p-l?Jlsacu? aydurrse ,{q paureldxa 1ou sr saposrdernol sl\frtsc(Is'I,!toN)rJY 'ssel Jo sJ"et{ uoqlrur 0I lnoq? {1uo sr ,{lqeqord aql Auunp esuodsa.rpunel luopouoJ luaraJ oJuesqEJo eulrl :qt lPql lJeJ eql Jo alrds ul -Jrpeql 'suourpuocrepurs fpred lszel 13rllra\ 'luesqe aJe^\ {eql ueqr'r seprcuroceposrda lse8unod aql puE'suonlPuot pelerdrelur fI s?qs) ayqrssodra,re,noq seuoz {uetlI Jo poued 8uI^\oIIoJ 'seuoz o,,{rl crueacopeleua8dxo Jo a u€JIpuIse " eq1 no-rftrrl suoq"npng leJol Jo euo Jo eurl aql Eurrnp ,{1uo pealdsepla ele-rr seEueqcqlr,n sepbuloc epostda1sep1o 'sluelu lE!-Ee {laJ€s aq o1 sreef ereq pessmsrp ?xe1eqa dnorS aqt 1o uolnloae ; r+ iq paleredas eq lsnur eql ul lueuodrul eq ol Pelredxe eq pFo^{ -1paslueu8els purou Jo slledsJauq auos roJ nI 'SUOrlPlalJoJ peseq aEnJar palelosr auo u?ql aJorrr uI paqJueque tdaJxe'sBes Jpolercrde eq1u1 peseec peq uoqe fum - :lqeaerqceuoqnloseJ eueceq serJads€ Jr llnseJ plno,t\ uoll"rceds leql -lueu{pes pe ?ls-alEuoqJ€caql JelJe sauoz a.EqL -suonrpuoJcru"eJo pooqle+l 5ql pu€ ezJSuo4epdod uI saEusqc ;o aldnoc e Suunp panupuoo uou?luerulPes FF! Supnlcur 'seSuzqc e8l€l dra^ eql 'raEuol qcnu aq lou plnoJ eleuoq*J puz suoqlpuocurogoq paleua?dx6 'ruaql I'I P(]e €xEl luopouoc lue proJer eql ur sd33 aql leql e^aleq ol uoseer Jo eJorr Jo euo ulqll,!\ pel"4rul sPt\ aleJrpul leruauruorr,luaeql 8ur ou sr eJeql 'Je,\e^{oq lsreed uolllru I?JeAes8uI uo[€lueurpas al€uoqJ€c IBJoI l"ql El .qpnrnur eq 01 sasrurord -mp lseel 13 roJ proJer [IssoJ? 8ur,real 1noq1r,n deru lapour sql ur papnltur tou la^ sB -ar "lep E rrropouoJ 'se{?ulou€ lsua plnoJ sea?eull luopouoc 'dlsnol^qo 'Je^e,{oq:ueeto l?ql uI seposrdeSurpacerd eql 'JlqluaqolJeu pedola,rep ete EJ :roIII ]O UoneJrJrluepl el3,![ sJeqlo eql Surrnp tueue rclurrs Jo leluelq 'eAoqP elrq^\ J€pled aJe,r\ reuroJ aqt tpql pepnlJuoJ -uoq.recoN saoeldlsadeep,{ra,r eq1 ur ldacxe pessntslp el€uoqrcc ,z ?DDtutqs lf eql qlra\ oq^\'(9L6I) sne?rqBd pue souI"g [q 'lr ssoJce ueEaq uorl4uarulpes pu" IllroJluot qll,\l pue peleueEdxo urzSepue i4)urllls 4 ;o acuoserd s€,r ex4 Jeqlo slueuele eupJaq 'spJs,tualJe ;-, r go pea:ds pe1tu11 e 8ur euos ueeaueq ecueJa]JlplsJl8ololqoeBled ueunp5 Surpocerdaqt SurrnplueuSels 'ueaco "x€l 'seSueqc pelcPal aql Eurrnq t r Jo :lduexa elqrssod aug r7 reqlo ol '[lqeqoJd auo lseel le 'eposlde lseEunod 'sluaas Ir43!O3 UOlSrpuer suo4Pl aJor['Jo '?x€l J3qlo u?ql a^rlrsuas ssal reqlle Jo asJnoJl€lllrns e qlr,$ luel gard- palelost puesuorlel erea ex31 paddrnbe-uJoJruoc lsol^I seBueqf, -srsuoceJe seposrda re1e1 eql uro{ pu? suolEeJ LrrN se snoeuo[a aq [ €cI crueero ol e^Illsues djea are^{ sluoPouoc Jeqlo ruo4 ?l?p peJe Ess aJolx aql elns etrurolo(I pur8reur aq1 otul os1e pue ellns Frrr--tIo spuD{qtog alq"rlal sssNsnossNoc I rEr.rspsnoq raurtoJ eql auolseul'I eql Jo seJe aloql$ eql ssoJJ€Jleqs l?ql 'IYJrCO.IOr{OSV.IYd Sr,{OS 8'6 45d Duopr^e eql .elqellet elEUoqJpJaqt Jo a8pe aql uroq lleJ se^\peJJe peJnpeJaql FoJ Jl? ,{11zraue3 qcrq,n aql-uo4eluau{Pes el?uoqJ?J Jo ol ere eposrda !a\a uo lou 'se8Eeurl Jo luelxe eql deur luarcqlns lseplo t :oseJeadde eql uo peseq 'esn"f, eql roJ eJueluv quoN urels"a ulo+ Ple(I 'IJoIJ Jelleq I Uopouos u€rJnlrs lsotr l elerpeururr cruxne ro Jpelsna peq e.leq deru '3X"] slue^e ueruo^ecl aql l€ql &$qrssod eql psssnc E eq d"ul esuodserpcr8oloqlll eql ueql 'os pafqns s?,t\auoz puIJ lErrlsa '{Jor\ JequnJ qcmu -srp eq pu€ 'secueJ?add€al due f;quepr 1ou JI OO1eCUenEIIr IeCOI Ol 6 qJl uqunlls aqt Eugnp ef,uep!^epuopnqlqslp fuopouor Pu" IBr6oloql!'I O?I 'ft Silurian [Ch. 9 References t4l lpf€arances, and he dis- much further work, especiallyin regardto rare ment Fig. 9.2. Ema Hansson typed the manu- i tbat the Devonianevents taxa. script and Christine Andr6asson drafted the li or euxinic immediate Most Silurianconodont zonal boundaries are illustrations. My research is financed by the basedon the appearanceand disapppearance Swedish National Science Research Council of lineages, not on events within lineages, directly and through Project Ecostratigraphy. which generally are considered to be more To all my sincere thanks. reliable.The evidencepresented here indicates IDTOLOGICAL that the former boundariesmay also be highly reliable.Both kinds of correlationmay theoret- rrl .ensitive to oceanic ically be erroneousas surviving archaicpopu- REFERENCES taxa were lationsand isolated'premature' advanced popu- kn-cquipped Aldridge, R. J. L972. Llandovery conodontsfrom lbl other taxa, or, more lationscan distort correlationsbased on events. the Welsh Borderland. Bulletin of the British D other changes. A One possibleexample of a minor event allow- Museum (Natwal History) Geology 22 125-231, lEme betweensome taxa ing a limited spread of a lineage is the local 9 pls. ttrs ald other taxa was presence of P. silurirus in strata correlated Aldridge, R. J. 1914. An amorphognathoidesZole conodontfauna from the Silurian (1975), with the M. chimaera Zone, as discussed of the Rineerike d Fahraeus who area. soulh Norway. NorJk Geobgbk Tidlskrift Lmer were pelagicwhile above. 54 295-303. tbenthic. Identificationof more recurrentsedimentary Aldridge, R. J. 1975. The stratigraphicdistribution hr lineages could exist anomalies, conodont faunas, and lineages of conodonts in the Bitish Silvi.ar,. Journal ofthe ril record for at least dur- promisesto be mutually beneficialfor identify- GeologicalSo.iety l3l 607-618. 3 pls. Aldridge, R. J. 1976. ing the environmentalrequirements of differ- Comparison of macrofossil Es: however,there is no communities and conodont distribution in the I rh€ gapsin the record ent conodont taxa and for studiesof oceanic British Silurian.In: C. R. Barnes(ed .).. Conodont r longer. The very large changes, including identifying recunent Paleoecology. Geological Association of Canada r size and the likelihood oceanic conditions. There is a limit to the SpecialPaper 15 91-104. I result if a speciesbecame resolution achievableusing presentconodont- Aldridge, R. J. and Jeppsson,L. 1984. Ecological specialists among Silurian r based correlations, as recuffent conditions conodoDts. Sp"cial tba-none isolated refuge Papersin Palaeontology32 1,4I-149. b be important in the must be separatedby at least a few million Aldridge, R. J. and Mabillard, J. E. 1981. Local 1r The taxadiscussed here years to be safely establishedas not merely variations in the distribution of Silurian cono- tduring the time of one or local fluctuations. Improved correlations are donts: an example ttom lhe amorphognathoides a period of many zones however possible(Shaw 1964, Sweet 1984). inteNal of the Welsh Basin. In: J. W. Neale and 3 M. D. Brasier(eds.).. Microfossih rccentqnd lrl. from fossil shelf seas,Ellis Horwood, Chichester,Sus- t thar the time of absence sex 10-17. 10 million years or less, ACKNOWLEDGEMENTS Aldridge, R. J. and Mohamed,I. 1982. Conodont -t rElationships remain to biostratigraphyof the early Silurian of the Oslo region. E of the taxa discussed. The quality of the manuscript was greatly IUGS Subcommissionon Silurian Strati- graphy.Field meeting, OsloRegion 1982,Paleon- fuships in other groups improved by comments from Richard J. tological Cont butions from the University of Oslo roeeny and on conserva- Aldridge, Stig M. Bergstrom, Ann-Sofi Jepp- 278 109-120. 2 ols. ftmately much of the sson, Carl B. Rexroad, and an anonymous Alvarez, L., Alvariz, W,, Asaro, F,, and Michel, dont elements is pre- reviewer. Permissionto cite unpublisheddata H. V. 1980. Extraterrestdal cause for the Cretaceous-Tertiary E of internal structur€s, wasgiven by Howard A. Armstrong, Hermann extinction. Science 208 1095-1108. increasing attention. Jaeger,Michael J. Orchard, and Roland Skog- Arnsden,T. W., Toomey, D. F., and Barrick, J. E. lund. In discussions -rdr available, give further during ECOS IV, a 1980. Paleoenvironmentof Fitzhueh Member of EEn]-. Reconstruction of numberof people,including James D. Barrick, ClaritaFormation (Siluriao. Wenlickian) Sourh- I use of the more slowly Brian D. E. Chatterton,Kirk Denkler, Michael ern Oklahoma. O,tlahoma Geological Survey Cir- cular 83 iti-iv + pls. L elements in taxonomy, A. Murphy, Rodney D. Norby, Michael J. 1-54, 7 Armstrong, H. A. (in press),Crnodonts ftom the gaimng Orchard, and Tom Uyeno, informed ! ground. Unfor- me about early Siluriancarbonate platform of North Green- He€onstructions require published and unpublisheddata that comple- Ialid. Grqnlands geologiske UndenOgelseBulletin. 'dd E--rpMuE tpreJlrqel 'nnl-Izl9l btotlg puo sl?fsoc Puelloc Ez I A t l uoun ltsuJ ols18opnury1s!8qoag spun'I ! t:?-1f I SuDztx-lottBul1 aqt urorJ seuneJluopouoc uelnls €86I 'I 'UOSSddeI 1,ql (,ft euuns qsrT8ug ' 06-98 016 I u){Dqrsuassth up4 uapu,ldpparyur oslv 6ZZ-50Z'Z16I-2861 F.,-r1of 'puepog ;uezrx ur uelsds hp anuapotlv uaqJslq)ta also taP assolx uaqJll qqry)t4o..l DlstSoloa1 spun'I Jo D ':d-LL) h:-.=uo11 nro pue nd-o€g ur.I -{o4)Judsst,intDu-tllsuowa4tirtt ldp )aSDzuV uerrnls Jql JouorlElr.r.ror Jql uO Zt6t 3 f'epeH 'Z7Z-8LI pun '_iI-stl ototlspuo 'rn1S ueJelleul JrSolouolqJoquO-ueqlrloldeJC 1;t uaBurypuoqqva8oloiuottlod sI (ta1:nllsemprop) I.( aqr ,{qder8rlerlsotq /er8olouorqcpjEd-ueluopouoCslutl9qJe^ tunz aEoloag n.{ qcaqtqrl sazap ;o sec *ar,J 'n,(-o€g ur.I 8€rlragurA 0f6I d H'qneluoqcs pu€ H'ra8eef uerudqlrguo,l runlrozoeled-llv 896I lA'sPeH $r- €86I '069-889 KSoPaD 's]dz ' L9B-nnB ' LOI-68 '@L6ID 69 {otps asag Zl )ap ua8unlaj 'uoln^ 'urseg u,{^!es 'elu€q puEelu,{d uI spuell ,B.E: I eete 'sroul[I le[of uaqcnSoloa? uaqxstq)otntso ueqcllpJoN relnJes peuuepuo0Elnue^ uorrEuSels !-{--=I pue uEupuexelv uro{ -rll.{ (uedFlsg) euozue{ce,{rn€r9 srrg,(q PUe =-!I(au pue rep mIS tuep sne ueqlrloldErc 9161 g 're3ae1 ueero t86I )I I'uosspuofpue ( A'^{o[e]pooC lJ I u'.qerl 'lo uopuol 'Id I '6S-St 979a a{aqsBun4tstog I 99-6n 9 llutoqs t9 _ EI '&aDoS lDt '(e1opo4 yo suoz .t =-zat Suueu crozoeeled n&nqtatg (uesqceg) eS.rtqa8ueqcsr.lr2:e8requel In4sa28o1otuoa1o4 ueunp5) pue 'g 'u 'spre{cQl pue -uprd rur IrJoJd-^o{qco'IAnlISsec 'LL6I H ' re?eef -uoH Alsuqslntr{ .{1spo.ro8re1r;1'6ls^elseu ai-: _ (uory rLne)JI^J') 'std€ eql sluopouoJ 696I ht C 'tue8iftO ;--i f 'Ue88e-I '829I-rZ9I '99r-6'r Lel '662-egz tZZ rN St a8ptr anau outplodoa1 &r:it avr punoreaueJ D1JV o^oN Sunlnepag aqcsrSoloaSleuor8er Og ulatqg o21ou1v lo lotSqoag lo 's?suplrv 'Qa1cog >.--,ed rraql pue seFqs eures pun ueIuIpES uI d,{I ueqcsduunql 'lcr.rlsrc e[r^sel?g 'elpils u?unlls Jo pun seo 'leneet uorssaJJnsluopouoJ pue Jql!-l '8re:-; tI iliJg 086I ) I 'De38e-I uJo^ uo,{eprelun Inls 9/6I H 696I L1 ,1 'stdZ'9Zl-lII DryrnrDJ(uedtv 's1d' 'dd DaDlPDuoo Sr.a:z 1 o1n3o1oag sa8uaag Z 99 771 I uo,req/rnl5 nlqdotSotuoapd sulPlunol elzue{cel eql 5 :c = Go1oa8 pue ,{3o1o1 eqcsrurE)) slqoJd-uolleJ sep l l '9261 uorJ selrqolul puneldoluopo u€unlls pelJlJllls rl =E:.:eJaU t[6I S 'pleJnP-I rrrr Sunrqnlueqqlolderg arq 11 'nBee1 'sld dd '€86I 'd!ed pu? g (I S 'uouell€qJ d --_:-_t 6 ohlol'unasnw tI ' LZZ C O pun a8oloag'awraq) n{ assolx '6-9 Ddod o?lDulv &a!?os | -: ;rroruatll (tuetutuns Z alaololg OSI lurads lo uvtag nz uatlotlcsuassl^np attuapo4vuaq)slnap eeumo pue 'puel€ez rrc?r qt uEdef 'nlo{rqs ur 1oc8o1oag ^{aN ^\eN l-:*_-i :ql ulo{ se8ulquesse np ua7unlpuo4qy srnpS re8urrnql uels8unf 'pqe4snv Jo slcou u€unls 3ql Jo uorlPlerJoC rou€a{n) pun '6s6I :(spe) pue'tureg g "lue p i=:ara 9L6I I saparqder8qerlg ueqllloldErC H'reaeef locnog f V N ,{A 'std (3o1otuoaop{ :ut sruopouoJ 14': g ;tst8o 1 oa g unap./ta d t 'IZ-I Lt otsv $Daqfios lo -lel v f sL6l o J c'JJoqJsls Dlj irl oJqo urelsEaquou puo {.&opaD -IX 'plsv tseeqlnos Jo fSolol pue suorlnq dd ndo4 ocuaug p fsalc$ I E{---:rrrsra^\o.I Jo /€oloea -uoeepd fSoyoe8eql ol tuoJ IIu€d ' 68Z Z0l Tozcadg eql tuorJ sluopouoc uEun[S upcueuv q oN eqt rEi 696r 'A ,,!\ 1q3!u) 'pdepw'spu€Fl r^\€I8ue'I IDJBqoaC's{Jor Jo pue uem,ropJo 'I te{rot puP loBI uoq€laroJ pue N g id'dueg * : - '::-OfZ orq6 'snqum ueun15 896I H 0L6I'f V'locnog 'dd gVt uua ng aqaSqstaPun 'std voday F9 l]1qo eql ',9861-uo! tt Z'dd 6I etl pueFeerC quoN uretsPe snuaos atulisul seuas t .-t :t +a) reJpqs g lA aln8o1oag tpuoluotg loJtSopaD {o {colua,/l\ pue eleuoqEc adfl eql 'I puE' g I Eslol\-el v ',{er5 'I :IlI 'uoqnlo^e seltpJ',{qdEraqPls Jlaqs 5/6I d'uellelA d'spre{cr-U r i-: rue (aJeJrnsqnsorqo uernls pu€ u?rcr^opro reddn t86I w t'lsrnH 'H 3 'pue1o11'W U 'I 'qroC 'C I tr'Desseg 'zz-LI 'sld ortSolovoaopd p ocEoloag Fd- ltre .uolurlo, de4s eql ele a,rrrN suo4sqrnued t '66-6S nI .;86I .tr ,reuJJelt ^&lueulpesplouoluule 'o aprpox dueSol,{qdpuE 'euoq€Plo 'sul31 +E!11 V t IpqolS qlh{ slua^a ,fuEuoqnlo/\e Jo 'S861'U splcnqrv 'uoqpuuod €1I.Ie[Ceql sluop o' i a I uulllol'tlooqap|n9 Jrozo.eled-prurJo uouele.uoJ I 'JsnoH -unol l Jo 'gEZ (uer1co1ue16-uerJe^opuene1e1) uetrnlS f 7a _o Jlozoaolod )aao1 El -ouoc .-.. oag gZ0 ?1tpg at1 ouqtal' kTdetaqeqsouorqJSuluJetuoc suo$tcep ]ueureleqlnhl916I C'raddeu puP A I'{rurPg Io '6f.2-8OZ t3o7uuoa1o4 -rr?i- ':rrJ rllqndaa $401?os :se8elspue seu€s uellnlls 086I H J 'puBIIoH LS lo 1ouno1 'sld gg t3o1o7uoa1o7 'easseuuel eu,{el!\'serceJoqlll aI ,rtoh ys'sn rolssrs Z '695-15S to 1ouno1 F.rluec'uo4euiroc 'pue1,{:e1r1pue elpuoqr€c pue'setceJorq'^qd€fqP4sorq *"" a5 louottoutatu1:(spe) 'elu€Jr^ lsel\ 'EIurArIA 'suo4 eql pue luopouoc(uer.rn1t5) ueqcoFeld g f'{tlueg !11 I 'ofte; 'q :u1 dorc -eturod u^{oluqlFl I eql Jo uEd l?seq IIIH €86I .I g a-:S 'A 'uuerxeED{ eso)I eqt Jo ued reddn eqt ruo4 sluopouoJ{col L_E9 f86I .ZI_I )adDd Klapog utsog uo8!1ct,'ttr LEEJ -ua,11f,peg-,fta,ropuel-I ele I 086I I J 'qrl4leH E looads loctSoloag '68-rn 'eerv uqnolruEl aql yo dSoloeg oueluo 'elns 2 t>s3oloag sa8uatg uap fiuapa s'pun'I l puo [8oloj -urueiJecnrg pue puelsl ul1nolruetr'dqder8llers 5 = dno:g esBsupu-ps^O to 'tSopag f.touownA l .S '9tuaau11 uo4 suouD) -orq uer.mlls puE UPIcI^opro '816I f i TlelnE.1pue -I'uossddef -uoaop| 't(3o1otany1Io satwnsq aLl luopouot nlnuv -{g/rd ur oslv 'LuloqI)ols p uapa s looqaplnD 'V C'luel]elpuE'C d 'p.IoJIeI'U J'seuleg E Xt 'adDd lDDadS '6NI_EEI t -c-:sl'mrojd puB serceJorq 096I uaprcN uolfisasIXX ssat?uol 1oct3o1oa9 g otoqla'1 spttor4dotuopouocu€IJr^opro tlqeq Fl) -'l -c :q louollouDtul puelloc Jo uellnls eq.L Etu-Pcs Jo aserf,eJorq pesodold eql pue r._!--jJ-s38e€u{ luopouo. ]o.rozoeled Jeao'I aql :epeH g I pue II9ude)I cqlu€qolleu 'sellrunruruoc 'C 'se.uhord pue * :+ppn5 7961 1'uossddal :uI puepo5 Jo ueunlls eql 096I A'I'epeH 9L6l'if 1'snerqgJ lI C'seureg 6 qJl uplrnlls eql SqJnp atuap!^a lBuognqlJlslp luopouor puB leJleopql!'I 7nr ' ! Silurian [Ch.9 References 143 s Siurian of Gotland. In: G. Jeppsson,L. 1984, Suddenappearances of Siludan ous foraminifera from the Llandovery/Wenlock Ld:: The Lower Paleozoicof conodont lineages-provincialism or special boundarybeds of the Wenlock Edge area,Shrop- r .-i Gotland. International biofacies? In: D. L. Clark (ed.): Conodont shire.Iournal of micropalaeontologyI L29-L36, provincialrsm. : -l--\7 Sessioz Nord,en 1960 biofaciesand GeoLogicalSociety of 2 pls. I *t>ckholm. Also in Plrbli- America SpecialPaper 196 103-112. Mabillard, J. E. and Aldridge, R. J. 1983. Cono- fuet of Vineralogy, Palaeon- Jeppsson,L. and Laufeld, S. 1986.The Late Silurian donts from the CoralliferousGroup (Silu an) of Geology, Uhiversity of Oved-RarnsisaGroup in Skene, Southern Swe- Marloes Bay, South-West Dyfed, Wales. t-!---' n. den. Sveiges Geologiska lJndersdkning, Serie Geologicaet Palaeontological7 29-43, 4 pls, Lae Llandovery-EarlyWen- C^ 51 l-42. Mabillard, J. E. and Aldridge, R. J. 1985. Micro- r ee upper part of the Rose Klaamann, E. 1984. Stop s:4-Jadivere out- fossil distribution aqoss the base of the It ..i rhe Mifflintown Forma- crop. In: D. Kaljo, E. Mustjogi, and J. Zecker. WenlockSeries in the type area.Pqlaeontology 28 ar \ d'sinia, and Maryland. (eds.)t Intemational GeologicalCongress XXV 89-100. lg, Sl -i-i7-569, 2 Pls. Session USSR Moscow 1984 Estonian Soviet Mashkova, '1. V. 1977. Novye konodonty zony I Sisrian series and stages: SocialistRep ublic Excursions: 027 Hydrogeology amorphognathoidestz nrzh-rrcgoSilura Podolii. ! d:.lnostratigraphy. Lethaia of the Bahic 028 Geology and.mineral depositsof Paleontologicheskijzhumal 4 127-I31. Lower Palaeozoic of the Eastern Baltic area. Mcl-aren, D. J. 1983. Bolides and biostratigraphy. h:elation of mid-Palaeozoic Guid.ebook-Tallinn 1 984. 62. Geological Society of America Bulletin erents with global Kleffner, M. A. 1985. Conodont biostratigraphyof 9431,3-324. the stray 'Clinton' and 'Packer Shell' (Silurian, -n-1ba.os- \drule 313 17-22. Miller, R. H. 1976. Revisionof Upper Ordovician, lF: Ordovician and Silurian Ohio subsurface)and its bearing on correlation. Siludan, and Lower Devonian stratigraphy, =raah).'. faciesand evolution, In: J. Gray, A. Maslowski,W. Mccullough, and southwesternGreat Basin. GeologicalSociety of *=J- Grtinlan ds Ge olo giske W. E. Shafer Gds.): The New Clinton Collec- American Bulletin 87 961-968. I r{t -3 pp. tion-1985, The Ohio Geological Society, Col- Miller, R, H. 1978. Early Silurian to Early Dev- A16!- Ordovicianand Silurian umbus, Ohio 219-230,2 pls. onian conodont biostratigmphyand depositional B l;_lrhawi Islands, Malaya, Knight, W. v. 1969. Historical and ecoromic environmentsof the Hidden Vallev Dolomite. lhe geologyand palaeon- geology of Lower Silu an Clinton Sandstoneof southeaslernCalifornia. Journal oJ i'aleontology --(i!t \.ia. XL. Geology and northeasternOhio. The American Associationof 52 323-344. 4 ols. i,.:t {iill M-21, 3 pls. PetroleumGeologists Bulletin 53 142l-1452. Nicoll, R. S. and Rexroad, C. B. 1969 [dared rEdFn und Stratigraphiedes Kuwano, Y, 1976. Finding of Silurian conodoDt 19681 Stratigraphy and conodort paleontology r Sl]urs- Abhandlungen d.er assemblagesftom the Kurosegawatectonic zone of the Salamonie Dolomite and Lee Creek c&t ll'issenschaflenzu Berlin in Shikoku, Japan(In Japanese,with an English Member of the BrassfieldLimestone (Silurian) in 1 Geologie und Biologie 2 summary). Memoirs of the National Science SoutheasternIndiana and adjacent Kentucky. Museum, Tokyo 9 17-23, pl.2. Indiana Geological Survey Bulletin 40 73 pp., E Graptolithenfiihrung im Laufeld, S. 1974. Referencelocalities for palaeon- 7 pls. Cellon-Profils (Karnische tology and geology in the Silurian of Godand, Nicoll, R. S. and Rexroad, C. B. 1974. Llandovery lIIl-116.2 pls. SverigesGeologiska Undersbkning C7O5 172 pp. (Siludan) colodonts from southern New South I Sur und Unterdevon vom L€ggett,J. K. 1980.British Lower Palaeozoicblack Wales. GeologicalSociety of America, Abstracts r ra Sardinien und seine shales and their palaeo-oceanographicsignifi- with Progrums6 534-535. I B.deutung. Nova Acta cance.Iournal of the Geological Society, London Norford, B. S. 1976. Faunasand corelation of the Ids tS Nr 224 263-299, r37 t39-156. uppermost Lower Silurian Tegart Formation, Leggett,J. K., McKerrow, W. S., Cocks,L. R. M., southeastern British Columbia. Geolosical $: Lochkov-Profilim Fran- and Rickards, R. B. 1981. Periodicity in the Associationol Canada,Mineralogical Associition Fi\rge (Sachsen).F/eiberger Palaeozoicmarine realm,Iournal of the Geologi- of Canad.aPrcgam with Abstacts | 39. 5 -5-59, 1 pl. cal Society,Lond.on l3E 167-L76. Nowlan, G. S. 1981. Late Ordovician-Early lB{1rhen aus dem Silur der Liebe, R. M. and Rexroad,C. B. 1977- Conodonts Silurian conodort biostratigraphyof the Gasp6 deozone (Ostalpen). Mir from Alexandrianand Early Niagaranrocks in the Peninsula-a preliminaryreport, In: P. J. I-espdr- a.eichischen geologischen Joliet, Illinois, area.Journal of Paleontology5l ance (ed.): Subcommissionon Silurian Strcti- trr- !9-107. 844-857.2 Dls. graphy, Ordovician-SilurianBoundary working bt. H. P. 1970.Ein Beitrag Lin Bao-yu, 1083. New developmentsin conodont group. Field heeting, Atuticosti-Gaspl, Qutbec hrf, onten-Parachronologie/ biostratigraphyof the Silurian of China. Fo.rJils 1981, Vol. II: Statigraphy and Paleontology, ituonologie im iilteren Silur. and Strata 15 745-747. 257-291.7 ols. *aons ch -nat ut wiss e nsc hafi- Lin Bao-yu and Qiu Hong-rong 1983. The Silurian Odin, G. S., Spieldnaes,N., Jeppsson,L., and Tor- *'eic hk chen Akade mie der System in Xizang (Tibet) (In Chinese with an shdj Nielsen,A. 1984. Possibilitiesof time scale t E_<-90. Englishsummary). Contributions to thegeology of calibration of the Silurian in diverse geological conodont faunasfrom the Qinghai-Xizang (Tibet) Plateau I 15-28. (P.T.) environmentsin Scandinavia.Bulletin of l5 Mabillard, J. E. and A-tdridge,R. J. 1982. Arenace- liaison of I.G.C.P. Project196 3 6-23. -ianf *rau 121-1,44. '88-I Z6Zuga nS Dpouo) -uoda11 pc1?o1oag nosv'W :f,ildot8q'tls 'bung 'rJlros to ta^ns lurtSopaC ,{qdu8qerls ol suoqnqul ut salpryS '{a^tns lot8opaC ltnosstn e8p9q -uoJ I ued oSeledrqcrycrlcry u€IpeuEc'ulldn -srl l rrelseeqlnosur (uerrn15)uoq€turoC eqloog aql suon^ue eql uI slcor ueluo^ecl -urEg eql requelu e'ePqS xs-flua^es Jo Jo ^\au Ja^ro.I pue uer:np5 reddn Jo sluopouoJ pu€ '9161 '1 'L 'uosduroqJ pue U I 'ple+ralles .{qder8qulg (O8et p"t"p) 186I d'uossurelsroql 'gjg '9767 tol sruot8ud 411/Asltoqsqv .I09-E6E 6t Dtluolod ocuatuy lo &al?os lot8opaC unossly\lulels€e oltSopa{) ot?V selqold€18 uo peseq puElod -qlnos uI uortpuroC e8puquleg eql Jo eseq eql e ,tqdEfqe4s pue sluop ;o dqder8qerlsolq upunFs eqJ 696I 'I 'rellel 1e (ueunpg lrun apqs Jo '9t-tz 56Zolso lo [,ts -ouoC €16I 'I J'uosdtuotll puP U I'plegrelles '95-Zn'Zl -Ja^tun aqt ulott nounqnuoT 1ott9o1oluoao1o4 das 0l- zL [P12oS 'ua${S uDrJt^opro aq1lo slcadsy :('pa) uo1ug 1 fiag 6'7267-uotst4)xg Ppttr Pnuuv lDt :dqd 'O :uI V'S n'ecul^old luau$uoJprl^Iuecueuv -8o1oa11utsvg uogtt|rr.j,V'ol.leluo 'uolItueH quoN 's{cot uEItl,{opJO raddn puP elpplw -er8qerlsuere8erN :( spe) uunq V 11pue IleBeS 'I :uI struolcel pue ,{qdPr8I}E4s(ueuntls) reddn Jo uoll?leI]or cIqdPrC !86I J '14e,!\S d 'slo E ls-EE^\ ll |tu uerrpuexaly-uere8elN Zl,6I J f 'ProJpues 'srr'lnlrt'nssrttoPg -so13qqqoag (er.usog,1\N Pu€gS uI crozoeeled 'sld Z '9ZZ-6\Z ! ou otr re8unof pue Jeplo Jql Jo euneJruopouoJ eql) llgpqtJd olzosaw1uozoapd olllotSrtbnsI eusog zs r rl pryozoel?d8epe1u r Sefuelseune; '(eltll slqt q1r,m,{reuumsqsl8ug ue qll,t lnq 'uels Blsluopouo) 'f tr^odrlU pue '11 ',rosed5 -sn11ur) etuenqtrl;o slrsodep(ueunlls Je,lro'I) 9961 'A 'Id I '16-68 9l tSoPaD lo atnj ueue^opuel-lJql Jo sluoPouoJ g16T 'snrzpelEs 'dd,gl Dqlunu sauas -ltsuJ .^otiluttg nutr;wlg , a4 lo 4lapng (uetsstty ' ,Z puo alols u-r)crozoeeled aqt ;o ,(qder8rlerlseql JoJ PuneJ uorlJ pub don a MaS aJuaPS unaslw eIIt ecu€cryu8ls 'H '^oseds 'elPls IroI ,$aN uI s{cod ueruoaeo luopouoc Jo 996I TrcI ^aN 'qd6'dd6g 1ued pue ueriqrg aq1;o uoq?lerroC 916I A'I'prerlsl; 'yZZI-LIZI (SoloruoaPd a8rc5 le f,Zopa) ur suounqutuo) luloqtlJots slsuatu1 6E Io toutnol pJEBerN -lol,ppots sqoltsa^lrp zley (uepa,t5) puellog eql Jo eluJlsueunlls eql ruo{ sluoPouoJ selEolueq u?IrIAopJOpue ueunls,t6I S 1l9us puoz'5961 A 'I 'PrPrl.rUpu? g J 'p€olxel{ Jo 'slo 'dd AN'{ro^ lIH z 59€ ^{eN 'twouo -net3oyl 'Ktqdot8tlotlsm aulr '?96I g V'1{€qS 't L- L9 IES 'nlSololuoaoPd p o)\8opag pue (orqg 'sld 6I "dd 0€I gg suoltDSltsa^ul -xEl lu.rlJlelllnur uo SuuBeqrraqt eFqS eql EorJ lo yodag 'urysnv to nxal lo {lED^Mn aqJ 't3o pue flcnluey 'ueunFs) Illsa pue g -lodg )lulouoJaIo nDarng 'sexe]. 'uoI?JU ouEI'I sluopouoC'Z16I S d'lloclN J'peoJxeu 'dd Kaa eqt Jo sluopouoJ yeddeq3-er6 016I C 'uoPpes 67 97 uodag Plcadg 'e.rrrrprtl uI 's!d9Z' dd EIZ 9E otsuosaP -ng pnSoloag ouptpul solsei{qllou -ung uaqtnSoloag np ua8unlpuoqqy slzo4sqv (ueun[S) e]tuJolo( eIuourPIESaql pue euolseurl pue looqapm{) '01 SOJ un$odutl(S tuopouo) {eerc uo[es eql ]o ,(SotoluoePdluopouoc puB g uoadotng puotag 086I ( pe) d H 'qnEluoqis dqder8rlerl5 Z86I g f 'elsor( J'pBorxeu 's1d'7 ' dd 911 gg 71o1s '28-6L sld 'EOL-nBg9V tSolot -uosapunguaqcstSoloag np ua8un1puoqqv'Wl.r -uoa1o4lo|ouno1'g nossm 'unr-qlr-Il€ (ueunls) -ialso ur urn{tozoEledsec 6t6T d H'qneluoqJs uoq€turoC e8puqurcg aql uro{ sluopouoJ sldz'9r8 90€,'0L61 Jo dpnlseg Ir6L I nd.'8IB.IJ Pu? g C 'peorya)I tlotsuosapung ua4cstSoloag )ap uaBunIPuoqPA 'qd S 'ZII-S8's0toloutlH ut uolt '(qcrerelso) uedlv ueqcsru.rc) ueFrluez rep -o?gsatut aa4otadooX 4euatg'ourg :( spe) npnyl serze$euoqlexueqcsunls lep euIqEIIJnEneNAIp '-I f pue uer-3upn1;r1 :u1 ( tcerlsqe qs{8ug uD jeqn 3unpe1lrtrrJe3ryne1.ron 016I d'H'qneFoqcs ql \'eseuqC uI) (leqlJ) SuDaX ur sEun€Jluop 'LZL-llL sa)ualrs qtrog -ouoc crsseul pue clozoeled t86I Suol-8uoH InO ZZ .nLt-OLtJ'J-e1L uotpouDJ'ulselvuJelseJqlnos'Juols tadDd E@rq) pue rellonf aql .lo lDulnot I eleJeH eql esEq eql uo.IJ sluopouot Keatng Toc8qoag satog palun -saure{pue 'I'I'ouefn -eurl Jo louolssalotd a-l (1co1ue16-,{ra,ropue11)uerrnl5 'P{SPIVurels€eqlnos 'punos rello Eas luequou u6tmg opuuo) tb\ns 5961'W N'aBE^ES a Io '€LI I slrbrtsqv ur euolseul'I elecaH eql Io fqdel8qe4s pue or suo[nquluo9 leil tEls I ssat8uoS ttilz'lsqY) eBIy''OL6I C C'relsqei[ puE V I'aulqsue^O r EpEueJ 'glldn eqloog loctSoloag fruottouotul e)ISeIV tuorJ sluopouoc (seun[s.) 'lS't uod '6961 u sFor lleruo^ec re,|lo'I lsEeqlnos I u€Dlcoluel\-uPue^opuel-I'?86I'I^l N'eiB^es to! swotSot4 tl|ljtr spo4sqv 'D?uatuv lo &alJog $Eopouoc qe4s t PuE,{qd?la '6rt-60r L9 7o43o1oags{selv uJelseeqtnosEo4 sluopouoc iqdEduErlsolg (086I pue 'V 'I + suot\oggsatu| ttoday 'sa nosaY PtruoN lo Juaul ueunls '696I O 'C 'relsqa^\ 'aurqsue^O tur,i:l puE U 'uossul9lsro[l puB 6 q3l uBlJn[.s aql SuFnp ecuap!^a lBtrollnq!4Ep fuopouot lBrlBopql!'I vnl References r45 r Slurian [Ch. 9 Thorsteinsson,R. and Uyeno, T. T. 1981 (dated Siludan). Anticosti Island, Qudbec. Geolo4ical 6. Rep or t Inv es tigations pls. 1.980). BiostratigraPhy. In: R. Thorsteinsson: Survey of Canada Bulletin 355 49 pp.' 9 and conodonts of UpPer Silurian and vtua', i . 1977. Conodonts and their distribution in llrDdoverian-Wenlockian Stratigraphy Lower Devonian rocks in the environs of the the Silurian of the East Baltic (Ohesaare,Kunk- lm southeast Alaska UDlift, Canadian Arctic Archipelago. oiai, Ukmerge etc. borings) (in Russian).Pcdii i d Geological Congress Boothia Part I contributions to stratigraphy. Geological Fauna Silura Pribaltiki,'fallin'.J, 119-792 Survevof CanadaBulletin 292,2l-31'. Walliser, O. H. 1.964. Conodonten des Silurs. r lLlandovery-Wenlock) r Uyeno, T. T. and Barnes,C. R. 1983 Conodontsof Abhandlunsen des Hessischen Londesamtes l i of the Heceta Lime- the Jupiter and Chicotte Formations (I-ower Bodenforschung41 1-106, pls. 1-32. ta- Canad,ianJournal of f2-. Uaufi ge Mitteilung iiber ikischen Karbonatf azies b -\lpen (Osterreich). *sischen Bundesanstak ts Pdaozoikum in Oster- t G eologis c hen Bunde s an- 19E0. Secozd European (ECOS ), Guidebook, t der GeologischenBun- 5 pls. l+?el Conodonts of tbe hreau of economicgeol- F 19 pls. Svatigraphy. McGraw- pp. -55 | @ovician bentonitesof t L'nivercitatis Stockhol- oibuions in Geology 3l blce of the conodont FI of the Palaeozoic(in t' fudi imir D in itrov' I nsti- ll7- | pl. t, J. 1966. Konodontska 3 Paleozoikaji i sz Bosne J rh€ older and younger ft- Bosnia).Geoloiki glas' It|c correlation of upper l}do\ician rocks, North t Pro\ince, U.S.A. In: D. .o! rheO ovicianSystem. ifuions from the Univer- lrian biostratigraphy of Folires. -4clo Geologica Sted 1980). Stmtigraphy irr Silurian and Lower : eorirons of the Boothia : Archipelago,Part 1 Con- $. GeologicalSurvey of -34. 10 Cyclesin conodontevolution from Devonian to mid-Carboniferous W. Zieglerand H. R. Lane ABSTRACT rO.1 INTRODUCTION Pectiniform conodont elemenrs occurrrns Clark(1981, fig. 58; 1983,figs. 1-4) discussed worldwidein penn- rocksof Devonianto early variations in the rates of conodont seneric sylvanian age record at least sevenevolution- evolution in mid-Cambrianto Triassictime. ary cycles. These cyclesare: (1) late Silurian His graphs suggestthat the origin and extinc- to Lochkovian; (2) Pragian to mid-Givetian; tion of early Palaeozoic (Cambrian and (3) mid-Givetian to Frasnian; (4) Famen- Ordovician) conodont genera reached an nian; (5) Tournaisian; (6) Vis6an to early impressiverate and that a tremendousdiversity Namurian; and (7) posrmid-Carboniferous was attainedduring that time interval. Accord- boundary. Each cycle,except for the last one, ing to Clark's work, rnanymore conodontgen- consistsof successivelow- and high-diversity era survived during the Cambro-Ordovician episodes followed by an extinction event than became extinct. Toward the end of the before returning to a low-diversity episodein Ordovician, however, his graphs show a very the sqcceeding cycle. Generally, low-diversity sharpdecline below'Index 1', that is the level episodes are characterizedby the dorninanceof at which survivinggenera numbered the same large-cavity pectiniform elements, whereas as those becoming extinct. A period of rela- high-diversity episodes record evolutionary tively low-diversity was maintained throush flowerings of pirbearing forms. After loss of the Siturianand into the Devonian.A maior the pit-bearing pectiniform elements in the recovery in conodont generic suwival is early Vis6an, high-diversityepisodes, such as dramatically illustrated for the Devonian and those in earlier Devono-Mississippiantime, Mississippian,but thereafter generic extinc- never recurred,except perhaps in the Triassic. hons mostly exceededappearances and, as a Instead, following major extinction events in consequence,the group tumbled from crisisto post-early Vis6anPalaeozoic and Triassictime, crisisuntil its final demisein the late Triassic. total taxonomic countsretumed to pre-Vis6an By its nature, Clark's method of using gen- Devono-Mississippian low-diversitylevels and era for his documentationof originationsand remained there until the next maior extinction extinctions.and using the lerigth of each event. geologicalseries as the basic time increment f'e'roz .elc{c -oto.lnsaDtd elpplyq eql Jo uEd re,$ot eqt uqtl,h sr uEqc eql Jo uro oq aql te uoucurlxe eqt :eroN] reqloue Suquasardarsdetlted a'roz -Drolnuan re^ro-I aql ]o dot eqt t€ ,{lrsra^Ip sI dorp p sr ereql leql s^\oqs F flEqc eqJ (tr6I) aue.IJo 0I-8 slrufl IeunpderE 0I nJ ol 8 nJ slue^euopcu$xo Jo suolIsod alouepseql o {cEIq rarAEeH lue^e uopcupxe fuepunog snore}ruoqrec-PrE eql eroJeq Pue lue^e uollssllxe ue?sr^ eql ieue le^rolu! ueFnruPN-ue?sl^aql uI srn''o eposrdo/hrsro^p-q3rq punsrp os leql aloN sletolJIreueS ^Fee Fttoz reqlEr slunoc pue uo peseq$ uro{ sreJlp osF (sqdB€ I ueql rglcedsqns crJrreds I leql uI sueqc s(IrElJ lI s,l4lC) seoaseql ro (I 0I 8rJ) seposrderhrsre^rp-Arol pue -q8q ueqt raqla'auoz eql sl tueurorcu errrlt (i 1186I) crseqeql leql uI (, 8rC'S86I) ]eeas Jo leqt or reFurrssr pu? (€86I {relJ Jo leqt pue I 0I 3ICtuo.IJ dltuerrlu8K sreJlp lreq. srrll etuB (V ueFnureN-uersrcurnol:) ueru€^ldsuuadlseIIEe ol uerddrsslsslltl roJ slunoc sercodsqnspus sebeds ptol ur uoneue^ euoz Aq 6uoz eql 8mt€4snlF rrerD-z 0I 8lJ 'ueluo^eo aql uI sprEl\on seseejaur e selsJlsnllrz'0I Erc l"ql eaallaqeia sle^ralur seposrda ,qrsr6^rp-q8rq urqlkr sorceds Jo leqrllnu IunE arrrrlJeuoqs snql seuozptlqdeJSll€Jlsolq -rxEr! aql lEql uoqrppeurs,{roqs e^lns eql sauese roJ Pu" pes€q (z'0I gunoc snoreJruoqr"J .l lBtolauoue8 e^OpFunc E splueuSes euo sel€llsnll qcnlrn uo arz ?rg) '(Z Bg'€86I '8S 3r.1'186I)1rEIC ]o ]eqr ol rse4uocsr 'ls"rluoJ uI 'fpnls sql In aurJap a^{ l€ql pue o 'ueruo^ac eql ur seposrdedlrsre^rp-^rol -q8q Fre^os slueAe {lrsrearp-a\ol pue -q8q ur-qll^\ slunos soleqpq e^rnc eql Eunoc cueue8 uEql reqlsr cgrrads F -qnspue rrJDedsuo paseqsr oso qr0 leqt ldecxo 'sqdef I31o1uo p3s3q q (I 0I'8ld qd?r8 ueluo^eq ,{trsrerp srqSuBeolc roJ (€86I '186I) {rED,{q pesnenbru eql 'I?adde Lr\ qd€JA eqt uI suopenPng z p^relur -q3a1aues eql s,{olduo1I euozro lecqdel8llells eql rDPru?Jparou aql 'slunoJ JrJrJedsEuq rod Suurncco selaedsJo requnu pexe eql ueql reqter @ eposrdeejrlua ue urqlr,{ seraedsurJoJrurlaed Jo reqlrnu -lold roJ ueds eluq aql rateer8 eql'Sulqder8 unurxetll eleuuordde ue 8lluellsnlll eNnJ-I 01 8rc uI pesnluaurarJur a(Ilrl aql Jo ssJUJuq aql uodn z "\.1"/ luepuadep $ slunoc sercadsJo 3ur11oydaqt rIIo4 uorlerrrJoJurJo f]rpnb l"ql elellsuoruep (Z Ot pue I 0I sSlO sqdzr8 o,n1 rng '(z0I 'I 0I sEId) uerddrssrssr6pue uEluoae61eql roJ rrereq peluesardesoql ol suoq?npng ,{lls -ro,\Iprepurrs d:a,r s,noqs lr :lrun Jrurouoxa eql se sercedspue pNalul aunl c$"q oql s? sauoz Sursnux.erp sear,$rsrarrrp luopouot uErrnIS pue u?rrrlopro roJqde$ (4 3g '9961) s.tee,v'5 (t, '(sauoz-mt1tn11s pu? -saploLltvuSotldtoulo tt + -tuolpJ'e'r) seposrda,{lrsra,rtp .reqSrq o,tr.1 ,{qpolznpund se,n1r 'uerrnlg aq1Euunp peupl z -ureurse,n ,{1rsre,rrp-,tt'o1 ,{1a,u1e1ar;o aposrda ue qEnoqlF 'oslv 'pez$1n spouad aruq 3uo1eql Jo esneJeqp4?nlueJc" sr r se 'slsa88nsqdeJ8 s.{Jelc sE crlErmJp s3 aq ol u"rJr^opro aql Jo pue eqt lp eseercep,{1rsrs, p eql a^eJleqlou z op arrl'suosBJl eseql JoC p:aco.td ,(eu 1:,ra1 rrraua8 aq1ltoleq selpnlspe[elep eJolrrqorqal uodn srseqe Surpprordse 1e,nse 'setres1ecr3o1 (---- -oa8 ,{q procer pcrEolorqeql Jo Surpu"lsJapun tr -- -,-- iffi_- praue? e ur pre leql seler ,{reuorln1o,ra ut suolleue^ asaqlJo aer^ peoJqdJeA € ele4snll z o1 qceordde oneuralsdserour e Sqdoye,rap 're^ea\oq 'op sqd€J8 s.{JelJ eulll q8noJql er? aa{ 'Jrdot sql uo reqtJnJ Supuedxa dlrsra,rlp luopouoJ ur suoDenl3nu lnoq? dpnls ? q suollenlcnu /$rsre^rplueuela u]roJ suolsnlouoc pa4prauaE ,fua,r acnpord dpo -rurlced;o lunocce asrcatdpu€ eleJmJe orour upo eperu aJe^\ slunoc orraue8 qcrq,n qq1r,n lI oI res 0r'qcl snoJeJgoqJBJ-plu ol uBluoaa(I uorJ uopnlo^a luopouor q salJ^J 81vl ftous [Ch. 10 Sec.10.11 Introduction t49 lcb€ account of pectini- 7Auctr-rations. In a study r this topic, we are lErematic approach to sz & D z;1:,:.:;./.1,;; 7 .1, z ral o EXTINCT ION z a[ approxlmatemaxrmum ak sirhin an entire episode - o tr of speciesoccurring per E- h employsthe sametech- l9!: I for oeating his divefiity z : x :rr.ed on specific and sub- il E cDun6 Tbe curve indicates p iv episodesin the Devonian, t I 196r. fi9. 58, 1983, tig.2), o '.r?r::..;, I rs a cumulativegene ctotal F rr In addition,that the maxi- itin high-dive$ity episodes I Fb,{pecies coults fot EXTINCTION kr differs significantly , Fg- ;) in that the basic s10t520253035 l) n $e se.ies(Clark's Li6c countsrather than Elatr interval after the ZOr)d )lo. sPEclEs r eletrr. Heavier black fl-ale (1974)-Tbe chart F rePresentinganother t \hddle pruesulcata- 'pue ldruqe uE ol lI Sul8uuqelc,{J P ulqlli{ eJeqa[uPlncco plnoc lue^e uogcuUxeue flpcqoqrod,{q'el.,{t elelduoc reqler e $ slqt q8noqrlv seseqdcpsllenpef puE e^IlelpEr Jo peslrdruocsr eposrdei(r$re^Ip-q8rq aql lsql pue epoqda ,{llsre^lp-,1Aoleql Jo Ued reddn eql ul srn$o lll\ol eseqde^qe^ouur ue leql eloN e^Ile4snll eq ol pesoddns$ IuerAEIpgql esneceqPepnlJul als seuleuselJtds IIEloN uoqelllll eredsJo esnecaqpue Poolsrapunlle/rr ssal erE feqt osnetaquleleq Pelellsnllltou are lnq 'snpoLDIpne snqpuStlod.eleue8 oqt uqu { f eroedselslxa sreqlo pelerlsnlll are s8urdnof ctoueSol,{qd roleu a^rd Ip^relul etup uElusel{ ol ueqe^IC-pIU eql uI alsdc ,{reuor1n1o,raue;o elduexg-€ 01 3I{ trcorw o -rsod (9) pu€ :u€rspurnoJ etoz-Din1dn? -3f1o4 ''3'e) uoqecrJrsre^rp ;o seposrde Irl -aJd pue u?uuerrJeC etoz-DlDJlnsaoJd eseql ur peJnpord era,t sdnor8 pcrSoloqd rl Jedol-lsod (g) lueruueu.regeaoz- subp8uDltt -rorrr a\eu snoreurnN 'eurrl uE9sr^ lserlJee reddn-erd (y) lauoT-snctort.u€ne^rC-pru(€) ol panurluoJlsql slue^e uorlJurlxapu€ uo4ec 1ue13er4 aaoT-suatnqaVai. (7) :u?rn -rpsre,rrproleru IeJeAesJo auo se,n eraqdsorq z -o\\ao'l euoz-oqap-efi (1) :are uerddrssrs luopouoo eql olur tm deep srql luaAe uon -sr6-ouoleq eql ur seposrde,$rsre,lrp-r'ro1 -surlxaJol?rrr E lue^uapun sEuneJePnlue[-,!\ol '€ 0I 'AId uo paterlsnflIsr u"ru snoeuelodrualuoc'eroureqrnc '(€6t 'd 'gg6I -sErl ol uerla^rc Jlel eql ul alJ{J elalduroJe laang) p.rocerluopouoJ eql ur ute?epereadde Jo eldrx?xauV 'elc/{. IprxJou€ dn eleu saseqd Jeaeu f1qeqo-rd pue peleullxlle dleleld ,{reuor1n1o,raeerql'snql aseqdcrlsrpnper8 e -ulot aJaA{o eqlq u,l\ou)[ s?unq luopouoJ ,{q pa,ro11o;'ped dJfte {aql ul espqde^tt€rper epnlqelq8rq aq1'serceds luopouoo Jo Jequrnu e-saseqd dreuo4nlorrao,n1 deldsrp saposrda eql ur eullJep srqt Suunq 'se8ueqccrueaco (lluenbasqns G ,{lrsrer'rp-q341'Surra,tog freuoqnlora 8ur :oleru palca;ye leql suorlercelS t3/\ ol -pearcnsroJ {rols looJ eql er? pu? auruado Jo serJese o1 SurpeelaEueqc crleuqc roleur n" 1es€qpuuou ueqt ra?re1z a,req,{1ensn aseqd p qtr^{ al"lerroc ot sruees(gOl-OOl'd 'SSOf r€86I slql uJsluopouos e,u1e,rouu1 'Eursolc raql reeu tee.{{s 'I86I {ftlC eas)eurl uzrcrrroprg eseqdf:zuorlnlole elrlelouur ue a,req,tqecrl 1se1e1ur f,lrsrer'rpluopouot ur eurTcapeql i -suelJ"rerlJsaposrde dlrsro,,lrp-r\o-I'luela uoq 'sropel crlauaSpuE rruopal 'lelueuruoJl^ue -Junxau€ ur salpulullnJleqt saposrda,{trsrar'rp parJqueprunle( se Jo requnu a ol raqlel -q8rq pue -,no1 1o laydnoc elrssaccnse s" lnq '(986I) '1rr uosuqol fq redpd eql ut E paurJapereq sl a1c,b,{:euorlnlor'e luopouoc Y padolar'apse saiueqo1e,re1 zas cpslsne ol dluo v peleler eq ot tou ueas (1'91 '3rg) ,{ltsrer'ry o SS1JAC IIISUSAI(I ur suon?nlrng aql 'saposrda,{1rsre,r1p-,no1 pue z NVIddISSISSI I-ONOAS( Z'OI -qErq 1o Surlsrsuor'selc,{c d;€uoqnlo^a luop -ouocJo ecueJJnccoaql equcsapad\.eJnlsrelrl z eql uI pue suolpelloJ rno ur elep ;o ,(poq 'sluopouof,eql Jo uollsullxa IBUIJeI{l ol e8rel eqt ;o srsfpue a,ulsneqxeue upql raqler rorrd lsnf crsserraeql ur pue ueru€^ldsuuedaql arurl uerddrssrssq,,q-ouola(qSnorqt,{1rsra,r1p ;o sgzd ur 'sdeqJad'ldacxa 'euru uersEurnol luopouoJ ur so8u"qJgo uos4eduroc€ slueseJ -lsod ur luasard lou s?a ,{IdrulsulaJeq pessnc -dar raded srql ur qceorddea,rlzlqenb rng svgt9 -srpslsrnq freuo4nlola crozoe€l€delpprtu pue '(ual I lddn ,{lrea 1z.ra,rasur uaas yeds dreuo4nlo,raaql 3urmo11o1eql u! lue^a pue 'eseqd 'eposrde 'JeAe]YroHsle^ol ueunls-aJd paqteer 'seuln 'elcfJ Jo suoqlurJapeas) aposrdepuu a1c,{cz svgt9 lE ls€el le 'flrsrelrp luopouoJ pue '(snoJapuoq ;o saseqdsnorrel aql urqlr,r sueld ur.royurped lsow d! ddn -reJ ele-I pue ,(Feg aql ur e"p4uopoglsu Jo arueJrnrJo lualsrsuoJ fue ruo-r; osp lnq -3orp1 pue eeprluopoqleuSorpeg :ueruo 'aposrdeuB dn Suopru seuozurqlra{ serceds 1o -^ao elpprl J pue dFeg eql rrr aep4uopoqteu qunoc p1o1uror; dluo 1ou'a1cdr qoee Sursdpue [z oI tes 0r'qcl moJaJluoqreJ-plur ol uBluo^eq uro{ uopnpaa l|Iopouot u! self,,{J 0sI [Ch. 10 Sec.10.21 Devono-Mississippiandiversity cycles lll -rocs Erlv and Middle Dev- futidae and Idiog- 4ltTlr\CTloN 4V3).lT r Earty and Late Car- UPPERMOST tnt diversity, at least at E 6tGAS hia-n levels. However, t seen in several early UPPER :.rolutionary burstsdis- atoa5 Es not presentrn post- TJJ {r- p€rhaps,in parts of o li the Triassicjust prior ataaS d tle conodonts. z t TSSPPIAN lrJ (ttz o & I rr cvcle is here defined r o hr of low- and high- I tllminates in an extinc- l_vepisodes I characteris- M IODIE dre evolutionary phase z rad1 e conodontsin this --))i rEEr than normal basal z mt stock for succeed- IOWER ht Gring. High-diversity ,d Folutionary phases-a z r earlv part, followed by -? hs tbree evolutionary ir F d clcle- An exampleof ! late Givetian to Fras- .E uJ ? 10.3. ,\- bs in the Devono-Mis- n e4eha-Zone Lochko- o- errs-Zone Pragian; z n l-Znlae; (4) pre-Upper B ct6rAfus \ mial; (5) post-Lower F I t Enoian and pre- Irl rliar: and (6) post- o P}IASE o VAR'U5 \"/ INI\OVATIYE to/ sap^J [z'ol'ces 0r'qcl snoJaJJuoqrBJ-p$[ ol ueluoao( rrro{ uognlo^a luopouoJ u! nsr troos [Ch. 10 Sec.10.21 Devono-Mississippiandiversity cycles 155 r er-olution.The bound- Including P. ovatinod.osus,which heralded a paralvea,K. ovalis (Ziegler et al. 1.976,Ziegler r end Middle Devonian new type of ornamentation,about nine species and Klapper 1982). b first occurrence of existedin this low-diversityepisode. ni'trs. a subspeciesthat Palmatolepis: Evolved at about the Lower gradualisticphase. (ii) Late Givetian m Frosnian high-diversity asrmmetricus-Zone fron P. asymmetricus via lat the high-diversity epbode Palmatolepis transitans and then into many E in the more open speciesin the Frasnian (Ziegler 1962, Helms ydrcus-Zone, d the Polygnathus- In th€ Upper Polygnathus latifos- 1963,Helms and Ziegler1981). rallel developmentin satal evolved from the varc J stock by develop- atrd endemic genus ing a rather large basal cavity. Ozarkodina Ancyrodella: Evolved in the Lowermost ; (e.g. Weddige 1977, sannemanni,which has its first occurrenceat asymmetricus-Zote from Ozarkodinq san- 979). The rnajority of this level, also has a broad basal cavity. nemanni into Ancyrodellabinodosa and rt Middle Devonian Although iterative in regard to Early Dev- in the Lower asymmetricu"s intoA. rotundilob r a small stock in the onian evolution, the large basalcavities in both -Zone a. The latter is the marker for the beginningof 4- I. corniger Group) these forms were once again an innovative the Upper Devonian and it continued the line ue geographicallyand morphological feature from which important on into many younger species(see Ziegler td Viddle Devonian subsequent rapidly-developing radiative 1962,Klapper 1985). Ziegler 1976). How- evolutionary bursts arose. Following the em- rirdid speciessurvived placement of these two forms in the Upper I ertinction event. varcus-Zone, a phase of radiation into new Ancyrognathus: Evolved frorn Polygnathus morphological types was initiated in the into A. ancyrognathoideusat about the change 'lieh-diversity episode, i interval is character- hermanni-cristatus-Znne.The time span up to from Lowermost asymmetricus- to Lower ; than 60 specieswithin the base of the Middle asymmetrirU.s-Zonewas asymmetricus-Zoneand then into typical lobed aa as Polygnqthus, a period of rapid radiation of many new mor- species (Ziegler 7962, Klapper and Lane I Sannemannia,OTar- phological types, some resemblingplans first 198s). i Tortodus. developedin the late Lochkovian. The Mid- dle-Upper Devonian series boundary has Polygnathw: Continuingfrom its flourishingin pia-n cycle recently been definedat the baseof the Lower the Middle Devonian and near extinction in asymmetricus-Zonewithin this radiative phase. the mid-Givetian,Polygnathus recovered with k of the varcus-Zone Examples of these new morphological types many new speciesin the Frasnian (see Klap- b? of the Uppernost follow. per and Lane 1985). A number of new Polyg- nathus speciesgroups appearedand died out Schmidtognathus:Five speciesevolved in the within the Frasnian including, among others, 3-ersity episode hermannitristatus -7nne (Ziegler 1966) from the d.ecorosus-, webbi- , brevis-,unicornis- , and narrow polygnathidswith large basalcavities. angusti.discus-grorps.One of the severalFras- Easitv occurred in the nian conodont biofacies seernedto be popu- d Widdle varcus sub- Wide-platform Polygnathus (including Meso- lated dominantly by speciesof Polygnathus mrs polygnathid plans ta;ri): Evolved from narrow polygnathids (seeKlapper and Lane 1985). Although seem- rerious high-diversity in the hermanni{ristatus-Zole. The first ingly occurring in more restdcted environ- r remained (i.e. Polyg- species are P.limitaris and, P.cristutus in mentsof depositionthan the contemporaneous lP- r-arcass./., the latter ascendingorder, then M . asymmetricus and M - Palmanlepis biofacies, the relation of the and P. xylus xylus) in n. sp. (former P. ovalis) (Ziegler 1966, Ziegler Polygnathw biofaciesto that characterizedby r S'eddige 1977, text- and Klapper 1982). Icriodw (Sandbergand Dreesen 1984), is not I - diffcilis, Ozar ko dina fully understood. Nevertheless,Polygnathus rE other associatesof Klappeina: Evolved at the beginning of the seems to be one of the most evolutionarily *alliseri and. Icriodus disparilis-Zone ftom P. cristatus and includes resilientpectiniform genera,haying survived at r rather endemically. the species K. disparata, K. di"sparilis, K. dis- leastfour low-dive$ity episodesand extinction .tsEI:Z, .voz-tuolnBu,lq 'pedole,rapseseqd cpsll€npBf e^qerpErrelel qcq'{\ IUo{ strrroJ k i]!q\ eql or IIerPur Pue eterrpul s.+ slua^J uogJullxs aleJlpu!s!+ uolleuoz luopouoc r r o,rI1e^ouulJo uoqcnpo.rlul;o suoprsod Pue irsre^p-A\ol pus pue seposlda e8e ueruuetlle{-9 6I{ :I|[ tuEpunog ueruo,.ec prepuels eq] ol uopeler rleql solJ,{r 'slur^a 0I -$rda tB-:r-@\: urql--9 0I 8rd 1 NOrrfNllxS I Fl Al rSdl Alc-Mo] 1p W suo/nouout od W I(, NOI1VAONNI \p n n t!g 'I I lr fi^ W oP.rda tt W n n 'I 1 ?4 - oaP.togtuoql A!SllAlc n 'l I n ueJlutarow n m lsoul,n lsouJ,n z ;9 1 I lrl o Je! z - '{tllotl n ll z 'I I H9IH oJelsod z z n n 'I 'I P W osuodxe W rlr: JrrrEJaqsprqleuSdlod rsDqlapor{cue'sprdalolelu n n z 11etsotrr1e uoqrsod 'l '1 )q' rt_6t uare1c1A1)sauoz NOll)Nr1Xl --d I olo2psao)d 1 5'ue -sz8r3 lsorureddn Ul rF !e 'sr leqt '(9 3rg) NVTSTVNUnOI A.LlSUSAlC -/v\O] 0I I n n Il{rEruseJJ eql laolaq NOIIVAONNr I uoqcu4xa dreqs rg [:ra ]NOZ B4 umuuau)ol Kltog (!) il)Af lcosldl IN]A] '(796y)Sreq xl,rz- ot oJlnsaDrd elpplW ? an snquuSottcuv pue olppothuy esoql lua^\JapunsluopouoJ 'spJE,!\dn Ia^el lEql rIIo{ pue auoy-ntop8uoul Jo t 's?aJ€ euueru uedo aroru palrqequr Eurr'eq pue paceldureeJa \ sed,&F r.Soloqdlou arau qr re ue8aq e1c,fusqa 1 se papJs8i'J i(lluernt an s1da1onut1o] lsour 'ueluolaq reddn eqt ;o ,fi?punoq Je^\ol eFiJ uBruueurBd (p) Jo serredslsoru s?areqrl '(1461 '7961 nfiar2) eql eAoq" leqr\euros'euoz-snJutauuttso secuanbescrleueSoldqd eseqt ruo{ serteds e1ppl1,atreqr Jo aseq eql lnoqe ,{g uo PasBqsr uollBuoz luopouoJ pnPuels erJJ 'EUeur 'eIlIE 'usruseJd eql tnoqEnorql selJeds snoJeuJnu I er-rcr leqt\euros ur punoJ slue^e pecnpord leql luaudole,,r.epcgqpnper? e u?gsr4 olur uBIuoAeOalPpltrAl Suunp res sep'(3 [zoI 0r'qJl snoraJluoqIBJ-pFII ol uBluo^oo llto{ uopnloae luopouor u.r 99I Hirous 10 [Ch. Sec.10.21 Devono-Mississippiandiversity cycles l) / dnpment that produced found in somewhat more restricted environ- Palmatolepis (Pa. trinngularis) is present in the touehout the Frasnian. ments. Lower triangularis-Zone,probably having its lm zonation is based on ancestrywithin the Frasniansubgeneric man- E phl logenetic sequences ticolepid group. A shortlived low-diversity Bhereas I most speciesof (d) Famennian cycle event in the Lower triangularis -Zone followed drrently regarded as This cycle began at the base of the Lower this major extinction event. Near the lower rre open manne areas, trrangularis-Zofueand continued to within the boundary of the Middle rriangularis-Zone,the I and Ancyrognathus arc l{.iddle praesulcata-Zone ol Ziegler and Sand- appearanceof one new speciesof Palmatolepis, berg(1984). Pa. delicatula, and of one new species of Icriodus. I. iowaensis.heralded an innovative (i) Early Famennianlow-diversity episode ISODE CYCLE phase. A sharp extinction event occurred somewhat below the Frasnian-Famennian boundary (ii) Mid- Famennian high-diversity episode (Fig. 10.6), that is, at the boundary of the Uppermost gigas- and Lower triangularis- After the appearanceof the innovativespecies, zones(Mcl-aren 1970, Ziegler 1984). At this Palmatolepis delicatula, in the Middle position almost all previous speciesof pal- triangularis-Zone, a rather abrupt radiative matolepids,ancyrodellids, ancyrognathids, and phase began at the lower boundary of the polygnathidsbecame extinct. A new speciesof Upper triangularis -Zone. Ma'ly palrnatolepid z 1 z z aY: uJ E IT:i: TY F:,':aS,TY Fig. 10.6-Chart illustratingthe extinctionevent in Pectiniformconodonts neal the Frasnian-Famennian zonal boundary. The b $e standard DevoDian Boundary. The extinction event is neat the Uppermost giSas ard triangularis phasecolresPonds approxi I of innovative low-diversityepisode ls within the Lnwer ftiangulans-Zone.The innovative mately to the Middle triangulnris-Zoneand the radiativephase begins at the lower boundaryof the Upper -EGluction triangulark-7hle (Ziegler 1984). Two morphotypesol Pa. subrectaare illustrated t:rrl,:-l!oH 'trd e Suqrqrqxe asu e,re3 (mtoclapaord 't4) sppoqteuSotord 'uegsr4 ,(yreeaql u etotollxuo erll "soz-mnf !lE- ueuonnlo^e JoJ [€I1 aql ereq^\ euoz-Dtolnuannddn-oqarosl eq1 Jo pua aql ol panulluoJpue auoT-ototlnsaotd D peuJEJ (olotlnsaotd 5 Jo eseqaql lnoqe 1e ueEaqlI 's?uneJrrrroJrurl a1ppl4 eqt ulqlra{ ue8eq elc,b slqJ, '4 nt; '3 a) lllaw d -cedeq1 ;o uoqrsodruocaql ur a?ueqJJrleuJ?rp (lrsra,rtp-,no1 t m-ucra -ro € sr aJaql 'u?rsreuJnol eql Jo asrnocaql uI ep,{J uBlslBuJnol (a) 2 ;upnpur'sprqleu8(1od '(snuud s4 ' suDur8totu' sd Jo ''3 e) sarlr,r.zcFseq ralpurs passassod,(gecrddl 'spre,ndnpenurluoc pu€ eurl srql la eceld1oo1 -\rlrezo'sprpoqteu8+Gr ?e[neluar sluopouoc (ueqooqrepury) uersreuJnol(1-rea aq1 ur 8ur snoJeJruoqJ€Ceql q lueuodurr auroJaq ol D ii:\ y setcads lsoru ;o -do1e,repserceds snqlou8 tlodopnas4 arp' (snlou ara^\ leql (snpoqlou8o1o"t4pue oppouotldrg) Jz-iuorslDloqzuo ew lo -uadnatq's4 'slsua8tnqtow'sd ''?e) seqr,rec eraueS onl ;o luorudola,rep ,{lrpa eql F .cqr€r pua u€ ot tq8nolq leseq a8rel raqler dq,(qeleua8 pazJalJEJEqc 're^ea{oH usruueuredelel eql uI p3qsllqBlse r r.r-.luqrad 0t u"ql erour eJea{u"ruoAeo }seleleql ur lcuqxe eIuBJeqleql euoceq lsnf peq 1eq1sarceds snpolltodslg Jo $ rrrvarrp-q8rq stql snqtou8K1odopnas4 yo sercadsseereq,q zurporl dlrroleureql ur parrmJo osp uorlJuqxg (t86I 'JrsssIJI al?l eql -tozo pue'srupou8,fiodopnas4'snqpuStlod Sreqpueg pue :eper7) aaoT-otoclnsaotd Fl nuopouof, ;o lsrnq ,{re aJ€ suorlJurlxa uerddrssrssrlAl-ouoAeqeql elppl4 aql ur ,{repunog snoJaJruoqrpJ ts bzl aqr eq ot surees pue relJz peJeAocerteqt ereueSruro;ruqced .req1g -ueruo^e6 eql lsnl 'snllvu ^rolaq It sr rrErn€uJnol al€l eql ul 'aull (uprlooqrepulx) g€ul ot qpl u?rsrpu -3t1odopnas4 pue snqlouBKlo4 ;o sarceds L Eruq setteJ fueur sso-rce -rno1 Suunp uoqnlole rllslsnp?J8 ;o porred qlln Euop 'l3urlxa aureJeq,{eq1 'peureruar ft:--ls -rog yn;d1aq ,{.ra,r E ,(q pe^rolloJ s€^\ teql (y'7t1 '9761 'pn sarcadsprdelolerupd eerqt,{po lqun sarceds EP i'i.qB{rEuer 3 sr uersleu Sreqpueg)aseqd e,ulerper palzrelecceue,(q ;o ssol,tols e ,{q papleraqse,r aposrdadlrsra, p g eql ecuelsrxaolur erueJsedfi pcrSoloqdrour,neu -qBrq ueruuerueg-p{tr eqr Jo pue eql latr'-',:uoPoqleuSorls? le1uouelpel eql'sn10u lsour 'esoz-o1o?rydnpeq1 noqe tV uorl€uoz 'sarceJorqeJoqs *le= :lysnrlztrf Qo4 e o1 luopouoJ prEPuslseql JoJ srseqeqt sr qJrqa -l"au eleJrpur suoqnqrJlsrpesoq,\\ pue u?ru { -,-*61 '12 ra euel) dnorS '(8L61 'pe Sraqpueg) u?rsrpurnol eqt ur -uerueCeql urqlln reedde leql BJeueB erc " Ait:-7'apn"t, eqt Jo uoqn sanads ollapouoqdrg 3o 8ur-re,nogdnuoqnlo^e mqlou8ttqdocgpue snqlouSouo4' snqtouSop(13 rc.xL1f, uEtsIELInoI,(Ift e u? ot osrr e,re? re11e1eqa 'percadde oloc 'sercadsasaql uo pasequoqeuoz reteA{-/i\olJ€qs mqtou3t1odoPnas4 ollapouotldtgpue tuqan4 snpoqtouSotot4 eroqs-r?euE qsrlq€lseppoc (y961) uesaerq 'Fa:' 1ns I F qorlJnpal ezIS aql 'eposrdar(lrsre,rrp-,to1,{repunoq uerddrssrssltr4l pue Sreqpueg t?ql os l?q^reuos pet?relecc? '){JOZ-Worsrptol.puo -u"ruo^e(I eql Jo pua eql spJ€^{ol (snryouSoluV'snpo1tc() lueurdola,rap pr1 D a-l eql ol panulluoJ -uoporJJreql ua^g '(snulw,flaatq'd 'snpisoJ ft ,.1q1 '(nqtouSor1ocg aponda K uaarp-tq8trquorstounol (r) -tutas'd 'sunwran'd 'DqoF'd 'a r) sdnorE qc-L ot n otg' mqruu? o11oq salcads snqnu8t1od raqlo Jo lpql s€ IIar\ &,:Jq'g) aep[uopoqleu '((rsor'eeer)s? '(€96I suryell)aruq srql 1eaczld 1oo1dnorS t :cl ro ruauraceldule eql Jasrll?1!\eas 'sJaqlo pu? 'selrqol4 ul?uat -snto$o)opou eq1 ;o uorsnlord ,{reuo4n1o,ra nri --.-prpoqreuS;o Sululos 'sprououru?eruos 'sprouaurf13) lno parpsdnor8 eql 'aldtrrpxe rcg 'mqlout(1o4 ro; f,ltsre,rrp 1.-r.etd{t ra,^ao1 eq1 yo do1 IrssoJJeqlo ,{ueurqcrq,t Surrnp 1uea.euoucurlxa qEq 1o aposrdaup oslp s€^{u?ruuerxeC eqJ ;I?F=EaJuo flrsJellq1 sseurB qlr,r\lueprJuroJ sr sluopouoJ Jo flsra,rrp '(1961 n1flar7 pue srulag) xeldruocsercads '*\-Z - srnD 9lo t oLlx u D erll - f ,r\ol aql Jo eJuerJnccopexe eqJ frepunoq prdayocqueruu€rus€rg aql ur asoql e{lJun E| !or1 etB[ e Pue evoz-0101 lectqder8qerlsluelrodrur srql ssoJJepelrAJns ere 1eq1sed,{1 lecrSoloqd:oru pacnpo-rd qu pue rd!-r rqr rE 3ulpue auo ,(Fee 4to1s snqtou8tlodopnas4pue'snqtouSotivd ereq ur8r-roJreql ppq sdnor8 sanads sldalolout I E?r:rPuInoJ eql SoPIAIP 'snpoqwdstg e pue o1oqnsaotdopapouoqdry -122,;o se8eeul dreuoqnlo,recrrauaEqns uetu ; r-.qs ieur,{pnls Jequng 'sarceds snpoqwutototd eenl!'swnuauoJ -ueuredrnoJ eqJ (tr961 ?raqpuegpue n1Bar7 1n-: ipuec4tuSts osp s€a\ snqtouStlo4 ,{po (1e1o1ur gg1 ueql aroru) 'lL6I '296I ra13ar7) paseq sr u€ruueured ) -xl.-z leqr Jo dol eqt t? spll sercedsueruuarueg ;o uorsn;ord eql II? JO eql Jo uoqeuoz luoPouor PJ?Puels plJo^\ E}.run\a [BurJpu? suIIJep eqt qcq^\ uo aseqd cqslpnper8 u,{oul-lle^{ F]EL-D sr sIqL snPoqlouD apostda t|tsta,up-uo1 eql ol esu a,re8pue lelel srql l€ ecueJJncco ftt rusuodur aql ot tnpunoq uorddtsnsnl4l-uotuotaq (r) lsrrJ rraqt s,req sercads (dno.rEcuauaSqns) [zoI res 0r 'qJl snoJaJluoqJBJ-plrx ol u"luoaa(I urorJ uollnlo^a tuopouoJ u! seIJ,{C 89I aroff [Ch. 10 Sec.10.21 Devono-Mississippian diversity cycles t59 pian boundary to the important Mississippian genus vative stock becameextinct at about or slightly Gnathodus.This is coincident with a marked above the qnchoralis-latus ftexqnus zonal decline and final extinction of the siphonodel- boundary. Thus, by the time of the lower of Famennian species lids at the top of that zone. Conodont diversity texanus-Zone, the ability to produce high- ral t only Polygnathus was also significantly reduced at this time. diversityblossomings associated with basalcav- woq athod.u.s species, Further study may show that this event sub- ity to pit-forming evolutionary lineages was & and a Bispathodw, divides the Tournaisian into two cycles, an lost. We postulatethat this lossof evolutionary andopoly gnathus stock early one endingat the top of the Lower crenu- potential resultedin a suddennaffowing of the tnnant stratigraphical latq-Zone and a late one ending at the top of genetic spectrum and was responsiblefor the Esurrence of the low the anchoralis-lans- Zone. low-diversityconodont faunas generally known J cDincident with a mass Diversity once again increasedtoward the from Vis6an to early Triassictime. rhich many other fossil top of the Lnwer typicus-Zonewith the blos- nids. some ammonoids, somingof gnathodids,pseudopolygnathids and (f) Vis6an to early Namurian cycle I otiers, see Walliser th€ emplacementof the new family Bactrog- nathodontidae (Bactrognathus, Dollymae, This cycle began at the base of the texanus- D oliognathus, Staurognathus,Eotaphrus, and Zone and.continued to the mid-Carbonif€rous ai ersir!^episode Scaliognathus). This high-diversity episode Bounday of Lane etal. (1986). By earlier continued to the top of the succeeding Palaeozoicstandards, the entire cycleis a low- of the Devonian- an c ho r alis -l atus-Zore. diversity episode. 7 lo,r-diversity episode, The size reduction of the basal cavity in i and Siphonodella sul- Pseudopolygnathusspecies observed in the (i) VisEanto early Namu an low-diversity rner gave rrse to an early Tournaisian continued within the evol- episode ot Si p ho no della species ution of the Pseudopolygnathusmultistriatus Sandberg et al. 1978), group (Lajneet ql. 7980, p. 723, fig. 3) and led After the major extinction event at the end of tte standard conodont to a Polygnathus-likeshape and pit in Ps.pin- the Tournaisian,a long period of low diversity e d up I i cata -Zone, rnosl natus. The radlation of the numerous speciesof was evident through the whole of the Vis6an rs came into existence the Bactrognathodontidaein the late Tour- into the early Namurian.In termsof the North itise phase (Sandberg naisianis a remarkabledevelopment, which is American succession,this includes the entire rt *'as followed by a very helpful for stratigraphical correlation time spanfrom the 6aseof the texanus-Zoneto r'olution during Tour- acrossmany facieslimits. This radiative event the top of the late Chesterian Upper (Linderhookian) time. in the late Tournaisianis distributedworldwide muricatus-Zone.This encompassesa number ra rhat recovered after and seemsto be the last significantevolution- of low-diversity,long rangingfaunas represent- lian extinctions are ary burst of conodontsbefore their demisein ing at least eight zones(: Faunal Units 7-14 *-znathus, and Ozar- the late Triassic. of Lane and Orrniston 1982) including such s of Pseud.op o ly gnathus This high-diversity episode, during which speciesas Gnqthodus texqru$, G. bilineatus, b latest Devonian were more than 40 pectiniform speciesexisted, was G. girtyi, G. pseudosemiglaber,Mestognqthus 'b\ rather large basal brought to an end rather abruptly near the top beckmanni,M . bipluti, Taphrognathusvaians , Jr?ensis, Ps. brevipen- of the snchoralis-latus-Zoneby the extinction T. trqnsatlqnticus,Cly dagnathus cavusformis, rorhls speciesdevelop- of most species.A very conservativestock of C. gilwernensis, Cloghergnathus globenskii, rtial (Kinderhookian) conodontsremained, including small numbers Cavusgnathu.sunicornis, C. altus, C. naviculus, |er basal cavities (e.g., of gnathodids, ozarkodinids, eotaphrids and Hindeodus scitulus, H. penescitulus,H. cris- ral l. polygnathids,including P.mehli. In previous h.tlus, H. spiculus, Ozarkodina campbelli, Tournaisian, there is a low-diversityor extinctionintervals, forms like Adetognathus unicornis, Paragnathoduscom- romposition of the pec- P . mehli (e.g. P . pireneae, P. latifossatus, mutatus, P. homopunctntus, P. nodosus, r at about the base of S. praesulcata)carried on the genetic poten- Rhachistognathusprolixus, and R. muricatus. zlata-Zone where the tial for evolutionary radiations of new forms Thus, an averageof only three to four pec- t-delicatus) gave rise exhibiting a pit. Hovr'ever, the P. mehli conser- tiniform speciesoccurs in eachVis6an to early P Ln Pa{ull sI lueAe uoll qJrq,$ur'as?qd lsqznper8ulrel-raEuol y_ (g) 1o aldnoc e Jo auo slueseldeJr 'JeAa,$oH ( r pq: Sg6I) s.raa^\S'oslv 'eseqo '(Z Of 3lS) tunoc sarcedsptol ul awarrap Iy\-rrEruo^ad eql Jo esEJ crlsgenperSe ,{q pe,ro11o1eseqd a,ulerper lq8qs e ,(1pnpe sr eJeql'pessnJstp dpeerle qfe-\l or IelIuIS sI qJIq^\ e 'slwd o,rl Jo pesodtuoose,t 1eq1aposrde seue^oJar JaIlJEe qly( uosueduror ut pue rF r" luer\a aPqs {JElq € ftrsra,rrp-q8rq e 1o EuruurEeqaql se,n 'lueae uoncuqxafrepunog snoJeluoqJ?J-Plur uuruuetllEc-usruseJc uoqerpeJsrql 1rde ol sarcadsearlelouur eql Sut,r\olloJftrsrearp ur d:e.T.ocerlsaporu l)-tq raFrz 'ald[IExe roC eql Jo Suredo pseq a8rel eql Jo Susolc ,{re,l e sluaserder uolssecJns d:zuo4n1o,re Il uoddns uorlnlo^a luop eql qlryr lueplclnot eJe,raslsnlll eallslp?J JqI EJueuV quoN ur ueru?^lfssuedaql Jo tF nFsar eql leql aAallaq asaql'l?InJqr?d ul'uerddlssrssrlr'{-ouo^eq aseqeql s€ aII]"s eql sr pue (9961) 'IDta eue-l l :-rEqrrn uE p€q slue^e eqt uJ'sureJJatlecrSoloce alqzge,re pelrqeq ;o ,{:epunog snoJeluoqreJ-plur erll Jo Ie^al 'irBpunog D FEe uerddrssls -ur puz ol paldepe 1eq1sruro;,nau fueur aq1s1 alcfc s {l Jo as?q e\L'swnulu snpoap , r=c uoqJrntxa ImFolorq lnoqe SurSuuq'padole,rap aseqd a,tlegper -u!H pue 'm)ltautu{s mpoqlouSoaSl' snpnuls Gi6I ) rasIIIelN 'u,t\oul1 lJoqs P 'sr.uroJJ^rl?Aouul aseql uoic (z) saprctpouSotp1' mtnuua' A' lta$qa&' g'snutlt d nq ilapm eJ? stue^e 'eposrda,$rsra,up-/rrol € Jo pue eql Jeau snqlouSolsl1cotqy'mtat4npou snpolllDu Dr hllrjlnlxe u? pe,{r\OIIOJ pJocal eql ur *edde seJnlseJean?Aouur -]ompa(I 'sanltods'V ' sntnq snqiDuSotapvere I FP-e ]€ql ffelc sI 1I qcns Suueeq sorcads 'dlpreueC 'sernl seldruexaeuos elJfJ snoJeJruoqftJ-Plur-lsod -zagpcrEoloqdroru areu alrlelouur ol esu aq1 go sSuruurEaqeq1 Suqrzur 'per?adde ad:lru rau snoleurnu e,re8qcrq,tr yo auos 'sEuruadopseq a8rel uzrue,rlf,suua4Jql Jo JllslJalJ€reqc ureue8 fil se uouoarp fte ,{Je,uleredurocSurr'eq sarceds ,(q peleu pue sarcads,.aeu 'proceJ aql uro4 Suueadde t:3-ar! s€-t!sasef, eseql -rurop eJB seposrde dlrsralrp-rriol esaql -srp eJe,lr eraue? luopouoc eseql 13ql eur4 I.ai :)qurnu au?s eql ur seunBJ ',([IeJeueC 'lueAe uolpuqx€ aIII"s eql lnoq? lv 'lue^a uorlJullxe uEur eql rrr,.l rruuxa Surczlder ue fq tnoqe lqSnorq aposrda ,$rsra,r1p ar'oqe,(pq8qs 1no pelp (1su133:'t7-9pae xalduns rg.r xuu uPrslP[unol -/l\ol'urel-Uoqs e qlr,n surSaq alcr(c e !Ku!8 g ' E a) snpolltoug snueSuerddtsstssrl,,q I Etfi:xroxa rolzur 'uerddrss$sqI-ouo,raq uegsrn-erdoql q (I) luepodurr aq1;o sarceds1se1 aq1 pue 'pereadde FI -:Un UetSr?UJnOI -stp snpoqlouSoto4 puz snlllDu&sn^oJ Ereua? Ep ;!eq ol urees :sa{olloJs€ stuaAapu? seseqd'seposrde luopouoJ eql ;o sebads 'la^el s$l lV 'pe{ool rr-\rl 01 uBuqruec olul pelpl^lpqnseq ueJ qrue puz gg1'?rg -JaAOUeeq seq uoll?leJJoJpu" uopeJlJlssslt llr*j: .ursre^rp-q8lH (9) uo pele4snlu sr selc/(ceseql Jo euo ;o eldurexe lerrqder?nerts aplappo^r ro; arueuodrut slt 'pua ldnrqe uy uralled c11c,{ce .uo11o;ol srrlees lelre}ur ,{puef,orlrlun '(Zg6l'pp urogoqsruegaas) F . ugrrl a-raqx,{ue srql uI uoDnlo^e luopouoc 's1ueure131sI?Je sreJrurueJoJpup'spodorqc?Jq'splouoluuJ" r -r=qd :r1s11enperE -uaEauros el?ru ol sn s,no1pseroeds ruro;rur1 osl? tnq 'stuopouoc ,{1uo lou pe^lo^ul pu€ h=:s i errleJoeql -cod snoreJruoqPJ-ouo^aq urql^r dlrsre^rP frepunoq uetue,r1,(suue4-uerddrsslssU{aql :trlull\.-. a asaqa 's8ur luopouoJul suonenlf,ngaqt;o srsfleuern6 le paunJJo selJ?punoq uegsl^-u?rsleuJnol q :zan su:oy fq paleu pue'uerddrssNsll^I-ouo^eq'u€Iuuetueg-usru Ili-iol B ol rrJnleJ NOIIO'IOAS -s8Jd 'uerJnlls-uerJr^opJo eql 1€ esoql a-*J: -irrsrea.rp-qErq JNOCONOT SnOUs.drNOf,UYC-ONOAfl( Suqz,ru 1ue,rauollculue lu?uodurr ,fie^ v t :gt Jo uollsuqxa .{o NuaIJviI r'0I pi. i-reuorrnlola uy (7) s'If,.f,C rUY(NnO{ 0'0I rrrjorrurDad,{q paleu SnOUST NOSUYJ-OIn'ISOd 1lqenpe-6 asaql 'uerd 'crss"ul ol uB9sIA rrq ue:sr6-ard eq1 ur aql ur padola,rap aq ol uraas 1ou ssop d1d 4\r{ro Jo sauepunoq -ufls saurl uegsrn-ardut pelou slunoJ selJeds '(9961 -raEue6ipue auel) ,{qdur?qer1g u":q aazq saseqd rFr letol u! uoll?nlrnUapr^\ auoz-ol-auoz eql 'IJEJ snoJeJruoqnJ uo uorssrutuocqns eql ,{q llr ,teu Jo uI lceJJoruraq plnoaraposrde dlrs-ra,rp-qErq e perpteJ,{llueJeJdrepunog snoro}luoqJeC-pltr 11 -s=qd-nads a.\rlerp€r eql uorperrp ,fieuor1n1o,laur aSueqc,teu slql IIIre! aql 01 pu" Bf,H3lu\/ quoN ul uzrddrssrssq,,qeql r Fcr:ls lo suorlJurlxe oJ Jrss"rrJ pu" JlozoeelBdal"T eql uI uo4n Jo dol eql ot spuodsarrocaposrda f,lrsrarrp-,to1 u.reued reln8ar e 'euoz ueunlxeN ;o -Ioaa luopouoJ Jo uorlJerlp ut se8ueqcroferu srrll Jo dol eql luoPouoc 0r'q3l moJeJluoqJeJ-plut ol ue!uo^a( uorJ uollnlo e luopouof, u! saltf,C 09I kous [Ch. 10 References 161 Eion of conodont evol- a regular pattem of appearancesand brought on a geologicallysudden cooling of :uoic and Triassic.To extinctionsof speciestook place,followed oceanic waters. However, we recognize that i evolutionarydirection the radiative phase.The first occurrences possibilitiesfor biotic control in the form of b rould be incorrect.In of new species within the gradualistic abrupt radiations of predators, for example, rile fluctuation in total phaseshave been widely usedfor defining need to be investigatedand could have contri- r pre-Vis6antimes sim- boundariesof conodontzones. In general, buted to the evolutionary patterns of cono- t be developed in the in the pre-Vis6anDevonian and Mississip- donts. pian, thesegradualistic phases were domi- nated by pectiniformsthat bear pits. (4) An evolutionary cycle closed with the extinction of the last survivors of the ACKNOWLEDGEMENT EROUS CONODONT high-diversity episode (Fig. 10.3), and a We thank Allen R. Ormiston, Amoco retum to a low-diversity interval domi- Research Center, for critically reading the nated by forms that bear large basalopen- manuscript and suggesting several modifi- h0ations in conodont ings. These extinction events, although cations. n-Carboniferous pec- theoretically occurring at the end of a I us to make some gen- gradualistic phase, may actually occur iotrt evolution in this anywherewithin a cycle,bringing it to an r a ryclic pattern. An abruDrend. REFERENCES : c-sclesis illustratedon (5) High-diversity episodes,common in Late and its l be subdividied into Clark, D. L. 1912. Early Permian crisis Cambrian to Tournaisian time, do not bearing on Permo-Tdassicconodont taxonomy. tnts as follows: seem to have developed in post- Geologicaet PalaeontologicdSBI 147-158. Tournaisian time. Instead, recovery after Clark, D, L. 1981. Biological considerationsand r Devono-Mississippian, major extinction events in post- extinctions.In: R. A. Robison (ed.): Treatiseon it a short-term, low- Invertebrate Paleontology, Part W, Supplement2, Tournaisiantime wasvery modestindeed, of America and hought about by an Conodonta. Geological Socicty replacingextinct forms with approximately University of Kansas Press, Lawrence, Kansas, Generally, faunas in the samenumber of new taxa.Recovery in w83-W87. t episodes are domi- thesecases was more a changein evolution- Clark, D, L. 1983. Extinction of co$odotts.Journal r ha\.ing comparatively ary dtection as opposed to a burst of of Paleontology S7 652-661- some of which gave Helms, J. S. 1963. Zur 'Phylogenese'uud Tax- ;s. numerousnew morphologicaltypes. ionomie von Palmatolepis (Conodontida, t*' morphologicalfea- Oberdevon).Geologie 12 449-485, 4 pls. species bearing such It is clear that each low-diversity episode Helms, J. S. ard Ziegle4 W. 1981. Fig. 62. In: R. 5 appear in the record followed an extinction event. Causesof these A. Robison(ed.): Treatiseon InvertebratePaleon- ba-diversity episode. events are widely discussed,but still not tology, Part W, Supplement2, Conodonta. rative forrns, a short (1984) GeologicalSociety of America and University of known. Walliser has discussed the Kansas,W98-W99. bringing about KansasPress, Larwenc€, biological extinction near the Devonian-Mis- Jeppsson,L. 1985.Greenhouse conditions and cono- -loped, lhat adaptedto and in- sissippianBoundary, and concludedthat global dont distribution.In: R. J. Ald dge,R. L. Austin nlogical terrains.In the events had an impact on bioevolution. We and M. P. Smith (eds): Fourth Eurcpean Cono' lial- in particular,these believe that the resultsof our studiesof cono- dont Symposium(ECOS N), Nottingham 1985, ETE coincidentwith the Abstractr. University of Southampton,14. dont evolution support Walliser's conclusions. genus ge basal the Klapper,.G. 1985. Sequencein conodont opening of For example, Ziegler (7984) discussedthe Ancyrodella in Lower asyrnneticus Zone I to a pit. This radiation Frasnian-Famennianboundary and illustrated (earliestFrasnian, Upper Devonian) of the Mon- I of a high-diversity a black shale event at the time of extinction, tagne Noire, Frar:.ce-Palaeontogruphicq, Abt. A mposed of two parts, a which is similar to Walliser'sconclusion in the IEE19-34, 10 pls. bwed by a gradualistic Klapper, G. and Johnson,D. B. 1975. Sequencein caseof the Devonian-MississippianBoundary. conodont genus Poly8nathus in I-nver Devonian Also, Sweet's(1985) end of Ordovicianextinc- at Lone Mountain, Ne'tada. Geologica et Palaeon- *ralistic phase,in which tion event is linked to climatic chansesthat tologica9 65-83,3pls. 'sloE'z6t-e9t puE 'uoq€rceds'due8ol,(qd gl,6I hl S 'Urg gg fuaqua4cuag lntltsuls8unpstol raunc) pue ')I 'zluelnrl ' l\\'reIEerZ "V J 'Sreqpues '9a11Du.ruDluOAaCI PAtDPt Puv SutUUnOnqstuaqv 'stot 8tl-€tl -ouuaptv aW lo [Sqoiuoabpd puD ttldDrSltotis g6t ndoI lutads oz1,tauv lo &anog pc18o1oag ulluoaa(I uO :(spe) reu.re,16U pup relSerz 'uisqDtxul^ud pu' salJololq luopouoC t(pe) ',11 :u1 drepunog u€ruo^eq reddn-elppq,{ {ry13 -I (J :uI uoneuozluopouoc Jelp^\-1t\olPqs eql ro] Ie^el pesodord eql 'euoz luopouoc et€ujelle pue slepou sercpJorqpquopouJr u€ruo sqotodslpeqJ- Z86l C 'reddeD pue lA 'relSerz -^eC alEI ?861 U 'uesearo pue V C 'SreqpuEs '€1,'I ou 9I sutotSot4t4tn xcottsqy '9SI-I g uouorltqnd tqdDrSlto4ssnualtuoqtu) 'nuauv {o &apos loc8qag srsrtaupruuetu€J uo uotsstxuulolqns,{-repunogsno:a;ruoq:e3-pt14 /ueruserCeql pu€ sluopouoJ 't86I L\'reIAetZ e roJ elep crqder8rlerlsorg Z36I 'g 'sue^\o 'tgz- Lzz Lzl tlouan pue 'g ltL 'srapunps 'J H d\ 'urolloqsuIeu oruauv lo &anos lnlSqodD {qdotSltotis 'sld 8'€8-t t'l sazuaoslDJtSopaD u1 -olg tuopolloJ uo unlsoduttg :( spe) uro4s8rag suoltoctlqn4otuto{tto) lo t1!so^!un 'epeAeNl?tl 'w s pu? ree,t\s c i[ :uI ueruoaec uEeoo.ng -uaC'(seuoz lalpul\-snqadsaq)sluopouoc u€ruo eqt lo dqdefqErts tuopouoC lL6I l 'rel?arz -^eg re^r.o-1Z86I J I'pler1l pue y trq 'fqdrnlq 'srd g 'gLg- Lng 'sI8- I08 6 uapltsa,y pun puopraqy uo^ a!8qoaD jg {3qo1uoa1o4lo lm nol seuepunoq pue Dp u! attutltsltotr uo^epreqo/1e$rw ezuerC eJll 'euIll :sserppEl€rtuaprsard 0/6I I C 'uerBIcI{ rep ue SunreperJ8ueluopouo3rep SunraureJ 'qd 0I 't9I-rII j1 octSolosoaolo4 -ra^ eura (996I ruudrurJo elep) 996I rlA 'ra6elz n on8qoag'uorlpuoz ollapouotqdtg-1sod 'slqtI prppupls,fi€urqirlerd pup sluopouoc snoreJuoq ' 991-1' gg Sunqcstoluapog )II satu.osapuo1uaq) -:eC re^\ol euos ;o dueSoldqdpup druouoxeJ -slssaH sapua?ultlpuoqqv Bunlnep"g eq.srqdeJ8 '086I l['ret8erzpuE' V C'?reqpues'O -F€JIS erqr pun ueluopouoc reqcsruo^epreqo t#"."t eruaSoldq4 pup eruouorxsl Z96I 'l!l' 'r.l1erz 'Z8I-SII 'osed sereJ yo pui .V91_19191 Ig le ftrs.realun ol8qa dtetcos pcrSoloeC ospd lg srorJ uuuoqnofll -uoaopd ta DJSopaO snpouJl pue snqi'uStloJ aW to uoinqlnsl1 puo Suutag lDtualuuon^ua e:eue8 luopouoc ueruo^eq elpprtrAlur su.re1led -oalod aqt uo unlsodruts :( spe) euoure-I d:euor1n1o,rg6L61 1,7'rc13er7 pue ) 'e8rppe.iA 'A C PUE'eue.I U H 'uollo8 ) tul zS6I'9-7. '66I- L8I q.4tr{ 'reunues plerCFuoDeujelul u€ JoJaprnC SI ndod looadg opouo) It uotl :ocrxel^l ,|aN 'suPlunow oluauerJps serJEJ -DtJossVlottSopaD' f,8qaaa\o{ luoprzto3 :(pe) uertroslne^|Z86l U'V 'uols(ulo puE lI H'eue.I seuI€g d J :uI ,{uEurreg 'uelleJrg eddl .ql 'sF Zl '96-68 g1 oct8o1o1uoao|o4 trro+ sauolseuqrn sortal sn4lvuSf,lodisnpanllo n Dl8oloa) ?)lselv [P4uaJ-lseg'eerv re,{rd acuecr;ru8rseqa 916I lA'relSerz pue')'e8lppelA tno:I uoules eql Jo ,{qdefrleJlsorq ueruo^eg 'srd9 '6rt-rLz -ornls 616I d V 'uotsrrrro pue lI H'aue-I BS oaDqtaT ouu8nqualcuag uelerqe8serz€C '9II_ZII ueueqqJeu.q ur pun lerqeSsnd,{a rur g saposldg drypurog snoteJluoqre3-pq,^IE sJnls-la]!3rap ueluopouoJrrc '//6I )'rSrpPrl^ roJsrsEq aqJ 9961 1 .,11're8ue61 pue 'U H 'euE-I 'gnz-ft1 Lg StaqublruasJnt!$uls8unqcst og '6Et-EZt DynoC'(topunog snotaltuoqto2-uotuorcg aq1 I pypDn'tSopaD puD ttldorSqt ts snoralluoq ;( spe) teers rg pue qlorde4 A :uI -toJ uo ssat8uo) lDuollouoiul tU1I 'npua) -C/( prnleu e :o;6urpee14 t86I H ^repunogO 'reqFi[ afiulo) fJeptunog snoreJruoqreJ-prl4ll?uorteu 'oz- LI n olruSoc olDJ slue^e -relul uP roJ lEsodord 986I I.,i{'lue,{eiApue'V lpqolSpuB sasseJord crSolooC €86I H O'rasrllpi[ 'g're8ulllreU ' J H ld 'urolloqsur€U' g'suer\O 'r6t-s8t 'J 'Ef,e)ls^orureN' iA '{nqcrsspN ' l[ 'reguEt{ 69 t8qaualo4 Io lownot slrzleq,{ elgll "A 'tqz-8ulf 'J V 'su€8rH 'A 'puel eH ''I Surleurcse;esoql :sluopouoJ O id 'lee^\S 'O'rourg ' 'appuarg ' 'UeE{rnog ' 586I lttDis d f lI H'euE.I 'std t 'It-6 Bzl ual^ 'zgz-69z gs slsldopag unalotad E: r '0f I-60I Ot wlzqot -uosapunguaqcn8qag Dp qrnqtqot (et:4.e^op lo uopu)ossv uo tauv 'uga ng ldolJe1ue8pa,r IEE:qt- quoN puB eoolna -oqcezc) eeJV uerpuEJjEg aql Jo ueruo,lec -uo-e8paa\V-sexeJ lsel\ pue oclxew AraN I-1.+tppry^l'stuopouoC) re.llro'I eql ur dJPPunoq uPrSerd-uer^o{qco'I urelseaqtnosJo uErdd$srsslr'\tr'tl6l l{ H 'eue-I G\:!\l nspue uoqrurJepeu .qI i p n'?-Bdnlqc'I :uI 's86I 'd H 'qn€luoqcs '196-tO6 {3o .OZI-ZOI 65 '13-9 raddel; ' l^'rel8"tz L f,t1dotSuottguo loJuoapd lo lourno[ epeueJ J i['N 'sarceJolq I rz,nd3 ure1sd5u€ruoleq sagaFa aN '(snoreluoqryC re^\o'I pu€ ueluo^eo mqtou8Klo1 eqt Jo sluopouoo (ueruse:g) '5 -:rJdery pue iA 'relSerz reddn '€tuopouoC) o apouoqdfs Jo uoqeuoz uBIuo^eCreddn 586I ){ H'euE'I pue'C'reddE[) OI'II3] snoreJluoqJB]-pllx ol uBluo a('luo{ uopnloaa luopouoJ uI selJdc 791 koLs [Ch. 10 References r63 Hd (Conodonta, Upper Ziegler, I,/. and Klapper, c. 1985. Stagesof the Zregter, W. and Sandberg, C. A. 1984. hooiferous). New.rle#ers _.D9von1anSystern. Epbodes 8 104-109. Palmaalepis-basedrevision ol upper part of stan- t:0. Zieglet, W., Klapper, G., and Johnson,L G. 1976. dard Late Devonianconodont zonation. In: D. L. k L Chlupdi, et al: The Redefinitionand subdivisionof the varcrr-Zone Clark (ed.\: Corutdontbiofacies and Drovinciali,sm. mdafv in the Lower (Conodonts, Middle-?Upper Devonian) i\r GeolagicaiSociety of Ame ca Speiial paper 196 urodian Area (Czecho- Europe and North Ameica. Gabgica et Palaeon- 179-194. t Gaio gischen Bun desan- tolagicalO 109-140, 4 pls. l Fk- rdonrs: those fascinating J o.i Paleonnlogy 59 lbcrc processesand global I l--:0. r.dmE for a natural D/C- d and M. Streel (eds.): ipous Boundary, Courier b*erg 67 24I-246. hdonteo der Eifel-Stufe d in benachbarten alvrgiana Lethaea 58 l - 19'6- The significance I rarios in limestonesftom E-os. In: C. R. Barnes rcy'.-- . Geo b gi ca I A sso c i a- Special Paper 15 4 s. 1979.Evolutionary Inian conodont genera t. Ga ogica et Palaeon- r|ooje and Phylogenie fuIen und ihre Strati- lHiatdl ungen des Hessis- Jtnbnchung 38,1-166, 1965).Eine Ver- neoeliederung-rprht an der fil Fottschritte in det d und Westfalen 9 anr stratigraphy of the E E-- C- Sweetand S. M. lsum on ConadantBio- J %ciety of Ameica lnrs and the Frasnian/ 4ial kciety of America, .15 no-1, '73. ,G- 1982. ^fhe disparalis troF.osed level for the r Boundary. In: W. q' feds.):. On Devonian toltsl of the Ardenno- t&red Devonianmatters. ra;r Senckenberg 55 11 Conodonts:the final fifty million years D. L. Clark ABSTRACT ous heights by the discussionof punctuated equilibria and what is called Darwinian gradu- A specieslevel analysisof the final 50 million alism, palaeontologists(or palaeobiologistsas yearsof the Conodontadefines tbree intervals, the new generation label themselves) have (1) a period of low diversity followed by, (2) a maintained considerable momentum by short interval of rapid diversification,and (3) involvement in the discussionof massextinc- another longer period of low diversification tion and extraterrestrial bodies and also that culminatesin extinction. through positive contributions to basic The Late Permian was a time of low diver- evolutionary theory. The Red Queen sification with no major extinction at the Hypothesis,coevolution, stasis, and periodicity Permo-Triassic boundary. Early Triassic of extinction have introduced concepts and evolutionaryrates were the highestof any time terms that have becomepart of the necessary during the final 50 million years of the Cono- working vocabularyof all respectablepalaeon- donta. Originations and extinctions show tologists,or at least,of palaeobiologists. some cyclicity during this final time of high While conodont palaeontologyhas not fig- diversity and rapid evolution. During the Mid- ured prominently in the vadouspublic debates dle Triassic, an interval twice as long as the related to theoretical considerationsof basic Early Triassic,origination rates of speciesand evolutionary principles, our sPeciality has generawere 50% lower than thoseof the Early shared at least a bit of the spotlight that has Triassic.Dudng the Late Triassic,only a single been captured by the profession as a whole new genusevolved. Th€ total of known Triassic becauseof the discoveryof 'whole' conodonts speciesis approximately100, distributed in at with indicationsof soft parts, fiftt in Scotland least19 senera. (Briggs et al . 1983) and now in Wisconsin (Mikulic el dl. 1985). Also, there is the poten- tial that proper interpretation of conodont element morphology may contribute to basic 11.1 INTRODUCTION evolutionarytheory. The eventssurrounding the closeofthe cono- Palaeontologycurrently enjoys a more lofty dont era are of concern here. The series of statusin both scientificcircles and the popular conodont originations, diversity changes,and press than it has at any time since the 19th extinctions have a distinct sequencethat is century 'dinosaur wars.' Propelled to glamor- sirnilar to those describedfor other bioloeical 3 Er.Il]uef pue u?rJqrusJ '986I uollea tllo4 pue ''86I IIrIurSpnui(eral pse qlesuels'f861 lleqtll) puBueurJJoH peqmued repun uo$nlo^e stslnqJlpougd lnq urnr.rqllnbete uopnlo,ra selPlolez snorftA eqt reuv tueuruon^ue Jo Jo |Ira:Forord, JI seq Iepour,,{reuorlets, eqJ s}ueuiuon^uepoq.rn ed DI puB /qrsle^P urnltq{Fbe le suorlBue^lq8qs IIuo qlr^r soteruollnlo^e tuelsuocseq Iepou uoeno peU oqJ suotlQuocleluellluor{ uo luereJJ:lplspun'salel Il-r-f L\o(oNoJ z'II uoqcuDxeluElsuocuou Jo lapoE ,,{reuopels,s^ salPlsuoDcsqxe luelsuoc }o lepoMeeno PoU-I-IT 8lJ luauro.r^u3 paqrnlsrO Allecrpouad -IIL-\Onlo^e luopouoJ Jo uorle.rnosarcaos uorleJnC sarcodS ftl: E qll,Ylur8aq ol Peeu 0t I eql 0t I Ercdur slePour Jo (n 9) | ireuollnloaa, EUIS -€qeJ 9. FEjrII :IE seIeI UOTlJUIIXE 6' I iLrqEll6lro 'uodar sql 0t a 0t .h r !L! rnq 'urauEd I"Jrp,{t s. m vea,{ uol11u s: lmd 09 (o llf,iuf,1-- uollnlo^e eql 001 oot ; -Io -' 'EUaAA I S.i p:]€IJossP sd€qJed uorlernOsarcadS uorlernOsarcadS 11 aq 01 perePIS l;:s.q.-ul I nt c -::..-6 0t 6 aqt Jo rnolA?qeq g) 4re tnaydruoc pcrSoloqd | -!:.sra-rrp tn es?alcul ue rp :cuereadde cpddrc (7) 0L ch ;r ;+-teJs uI eurlJap € (I) -sqJouud g freuorlnlo,ra a I !.E€ leql splouourlu? oor J ireuoun 1or'a ul sluela tr ..ar '(9961) asnoH f,q lUat!oirAuf luelsuoc sr.-rflar aruos e,req deru t I+\1o! l"uoqrpps uv ;o sserord anlua aql pacnper (y961) dneg srae-I).'eJ?ld eruzs aqt ur deal o1 'op uec 'uotruqxa (ueuop) '(I rI atc no{Euruunr eql p sel?t tr ' ', slrperd uaano ;e: ieoep lutlueuodxe '996I uolueg '?86I rIl|us Preu,(EIt pu€ pau aql s? 'esneJaqJetleJ saler d:euoqnloaa FIt eqr Suunp raqurnu qlesuels 'A a) uollnlo^e 'aJueq Pue 'uollcuuxe Jr pe^I e^?q ,(aql 3uo1 ,roq 3o ssalpreEer p- 's:zed uolgtu 09 Io seleJ luelsuoJuou Sutplerd eurrl q8norql erurl ,{ue 1epuqxe euroJeqfeul saneds'snql -'t 'se1er I iJrrroDnloAa s.dneg olul suorleqrngad (ectsfqd) Izlutrrruorrluo pnbe 1esarSelerls a,rqdepe ,l,rauSur,ryo,ra r j:r$ds tuopouot JI ,{q pelcsge are pue rep8arrt ere setceds ere ruels,{socacrseq eql Jo u"d eJ?leql sarceds -(b - d) Jo saler uoEJurlxe lEql pue 'sursno3 pe^ll palelar ,{qecrEolocaraqlo pu? 'feJd 'sJol?paJd l 4. !ra-bo[ teqt Eutqdue -reuoqs Jreql usql uoncullxe ol luelslsal eJolrr Jo lsor| Jr.llesn€JJq ellllns ot 1sn[:lqrssod Ig :'soJ $lnsal JeIIuIIs ', 1ou ere sarcads ?ur,N,r:ns-ra8uol tBql aJuap se dqornb se sauozer'rldepe raueq olur ellole tlu:r uorurnue q8noqtlv -r,re se,{1uo,'ae-I s.uale1 ue6 ul pesn salJnt lsnur susruzSroteqt slseSSnsleqt srseqlodfH d:? ir[qe]s ro 'i(€teP d1qsro,r1,unsJo {tueaurl aql leJdrelul 'sreqlo ueon6 ped eq1 ,{q peureldxa lsaq eq f,eur t rts-r rno slurod dneg 't Euoue '(9961) Dauuex pu" red\ 're^e^\oH (9161 uapn uen) ,,rire1,sq1 '(9961 :gep) r.Erlttrnu sI 's pue '0 = aunl ',{1rsre,,upurnrrqgnbe le satcads alE-Js.ueP^ ue^ Jo lxeluoJ eql uI elqEpuels oS' d ,ro raqurnu sI 'seleJ e ;o aEuzr eql SulJnp es?eJcep to as€arJul -Jepun sr uoDnlo^3 luopouoJ l?ql eJuapha F.ads Jrs b pue d eraq,t lou seop uorlJuqxe ;o ,{lyrqeqo:4 'srusrue8ro se paraplsuoceq u?J saAJnJdrqsro,rt,rrns luop JOJSoIBJ UOrlCuqXa IUeISUOC arc',S$eqloddH, -ouor Jo ,{lueeuq p-raua8eql 'eldrusxaJoc .dt:S:='S .IIEA aq1 o1 Surprorc€ 'srql Jo II3 Jo slFseJ I E{rr$uxe puu uorlelceds eql (ssz19 3u1qoo7 aqt q8nong'11otrc3 s? 'saJueJeJlp luzcqruErs q1r,n 1nq sdnorE [€rI res II'qJ] sJBa,{uollllul fryg lBug aql :sluopouoC 99r lch. 11 Sec.11.31 The data t67 ? Looking G/ass). The speciationand extinction to a single equation, ancestralto the later 'euconodonts'then the Is- according to the Conodonta survived for a little less than 400 S,=Soe(p-q)' Ent extinction rates for million years. While this longevity is not 7 of extinction does not where p and q are speciationand extinction remarkablefor mostphyla, the rapid evolution, drring the range of a rates,So is number of speciespresent at some biostratigraphical utility, rich diversification, d\-ersity. time : 0, and S, is nurnberof speciesat time : widespreadgeographical distribution, and gen- Xennett (1983),among t. Raup points out that exponential growth, eral abundancefor th€ whole of the Palaeozoic hearity of survivorship decay, or stability depends upon (p - q). and Triassic,distinguish conodonts as a signif- hlen-s Law only as ev! Althoughexrinction results from an increasein icant group of fossils. rriring species are not 4. similarresults come from a decreasein p or While a complete understandingof cono- uion than their shorter- anything that lowers the numerical value of dont evolutionwill neverbe derivedfrom study l extinction rates of (p-q) of only the mineralizedconodont elements, the r and are affected by If conodont speciesnumbers are plugg€d morphologicaldiversification of theseelements tdcal) perturbations into Raup's evolutionaryformula for the final has left patternsthat can at leastbe described. E nonconstant rates of 50 millon years, (p - q) yields a negative Of particularinterest is the pattem of diversifi- ,6olution (e.g. Stenseth number during the Middle Triassic, and this cation prior to extinction. Was the Norian t 1984, Benton 1985, exponential decay results in Late Triassic demise of conodontspart of the major Trias- (Norian) extinction. sidJurassicextinction event (Raup and Sep- !d the entire processof An additional model for which conodonts koski 1984), precipitatedby extraterrestrialor may have some relevancehas been proposed whatever'cause, or was conodont extinction by House (1985), who reported a sequenceof only a predictable part of their evolutionary events in evolutionary patterns of Devonian history (Van Yalen 1.973)?Were extinction ammonoids that may illustrate some basic rates constant or variable? Conodont extinc- of the whole picture of I evolutionaryprinciples. His sequenceincludes tion is a rnicrocosm (1) a decline in speciesdiversity, followed by evolution and extinctionthat featuresso Prom- (2) cryptic appearanceof novel groups,and (3) inently in the literaturetoday. Even though the an increasein diversity,that leadsto (4) mor- data are limited, it is useful to determineif the phologicalcomplexity and normal evolutionary evolutionaryrates of conodontsare moresimi- J the group. The sequenceis con- lar to those predicted by the Red Queen 10 behaviour of sidered to be initiated by euxinic conditions, Hypothesis and biotic interactions or by the bn perhaps associatedwith global transgressive Stationary Hypothesis and abiotic factors events. (Fig. 11.1). While a few details that describe The evolutionof conodontsduring their final the rise and fall of conodontson the generic 3y 50 million years in part illustrates a similar and family level are available (Clark 1980a, t{ cyclicalpattern, but wjth somedifferences. For 1980b, 1983), this chapter addressesthe this report, origination, diversification, and evolutionarystatus of conodontspecies during extinctionrates are mergedinto the encompas- the final 50 million years of conodont history. sing 'evolutionaryrates' terminology.Are any J of the models important for conodonts?We l0 need to beginwith a look at the generalpattern U.3 TIIE DATA IN of conodont evolution. For a group of organismsthat hasbeen studied for a relatively short time, it is remarkablethat lmconstant extinction the taxonomy of Permian and Triassic cono- 11.2 CONODONT EYOLUTION I .rolurion rateswith donts is already so confusing.At the generic ! SaatioDary' model has a.d .n\ironment. After If the various'protoconodonts' of the late Pre- level there are fewer problems than at the .1935- cambrian and Cambrian are consideredto be specieslevel (as is the casewith most groupsof 'qcoda snor^oldruo4 sarradsre^oploq requnu earEuado :qroda uI SuqeurSo I a.s:nb aql sesEr sIqJ Jo - leqt sarreds Jo requrnu: eere peuialled srpe,{Eorllrur puIJ eqt Suunp ,tr$e^rp rlJrceds-€.Il .8rC E-farar JrssBr{ alpprtr l ^{eu 0S *:u; uaua8 yo sraqrunu s3t33ds -lvl_ol_ rrn --. 19salur luep,rmba 09 0r oz I L-: rereduoJ eJaua? l sepads 91 d1e1eu Nvtv{i|]d I -r =:urpe.1-? o1 uerqldcg) fIV"l eql 3uunq 'arur1 lK{r: NVIHJ.ACS r E:EaS rqira eq1 ;o o,t1 k-,- pleuiuo uetruro4 NVISINV r =L-ads aqt Jo lsow :-_r--. rqire i laterurxordde m NVtNt0v-l r i5zqord oraql lPAJetur 8t ,LL L=q '1!'11 'z'I I sEIC) n- :Sra ur satceds NVINUVY 8t ,LJ t ,i-=Jr uorlllu €I euos -ed JreT aql Suunq NV IUON -r-- 9l al \oIIn'IoAs EI OI -\ATAIUUiI Ar-rsuSArcsSrclds ''II el€JncJe uB aq ol pareprsuoo eJe pesn el? a^rsuelxe uu dq peradurel eJntejalrl luJr leql sJequmu aql'al€rncc? aq ol lou paalue.rzn8 -Jnc eql Jo uorlelnq€l ar'qcefqns e eJ€ uoder \-:::rra4 etB'I Jo sturel elqd\ 'orsspr{ ar{l roJ ereue8 61 pue serceds srql Jo elep eql pu? 'paqslqnd uaeq 1ou a,req 'r F qI i:6I lee.tr5) ,ra1 gg1 dleleurrxordd" s1elot srql pue 'ereue8 sqderSouour osaqa suorlzlardralur sarcads lr !"- l:lJ3seun ,{le^neler pue selcadsyo spodar 1p lsoup peldeJc€ al?q ur saJuereJJrp eAIosaJ IIur leql sqderSouour pqt t T;f\: ol asr: ue,lt3 I 'sluopouoc rrssErJl-orrJJad r{ll.,\,\eJuauedxe Jrruouox?l roJ aruq fpearp sr 111nq '(sursruzSro ; ura- lla.\oPloq a^g ala,{r 'ucode Fl;': sn1r Suunp paleur snor^erd tuoJJ ereue8 re^oploq Jo requrnu = eare uado :qcode lpql ur 3u$eur8rro eroua8 Arau lequnu c--\ rI?qL\JS ls?el lE Jo 0t : Eere peurelled /ftolsrq luopouoJ Jo slEaf uortlu 09 lBurJ eql Suunp,{1rsra, p cuoueo-Z II SrC | *c.:r:rd uro4 sJa^oploq vu3N39 -tvrol_ I .r.:c u:qr ;o aAIJ'EJeueS ':ierqric5 eqt Sulnq ot 89V ftr.--q. :usardar o1 lqEnoqt I I i.=f rql s:eef uoqpru NVry'tu3d3t v-l s Eja:J ueruued el?'I eql [:;i;r::.2 E. :,1!. e uo (suorlcuqxe NVIHJ-A3S tr rr uESSs3euq 1I 'u?lur ;d:.l qcrq,r Jo or'rl ,{1uo NVISINV I F: :-raua8 ueruuad ele I m I E. 18 Pue 'SpOUedsnor^ NVtNtOV-l :, I 1x lqeqord 'Jrsseul eql Il rz :Jar erarll uounlo^a NVINUVY ry-E 0s Iesg eqt roJ elBp tt inqs: pue II 711sEtg NVIUON (63;1 t raup4 pue (286I) v ] ;r fu.-er: aurg luecer eql uo p puz =3y uoqeurrxordde l_tsuSArccrufNf9 [r'I I 'Jrs II'qJ] srBa^ uollllur ,i$U lBuU aql :sluopouoJ 89I lch. 11 Sec.11.41 Permian to Early Triassic evolution 169 approximation.Ages and datesused are based Permian was an anomalouslylow interval of on the recenttime scalingdata of Harland.etal. diversificationor whether the Early and Mid- (1982)and Palmer(1983). dle Triassicwere anomalouslyhigh intervalsof Figs 11.2 and 11.3 show genericand specific diversification(at least as comparedwitb the data for the final 50 million yearsof conodont Permian).The former is apparentlycorrect. evolution.There wer€ at least 19 generaduring At leastfive Late Permianspecies are found the Triassic,probably six 'holdovers'from pre- in the earliest Triassic. The genus Neo- vious periods,and at least 13 new originations. spathodusis reportedin both the Late Permian Late Permian generanumbered at least eight, and the Early Triassic,but I am not awareof an only two of which originated in the Late Per- individual speciesof Neospathodus lhat ocq)rs mian. It is necessaryto view originations(and in both the youngestPermian and the Early HIAN extinctions)on a'true time' basis(Fig. 11.a). Triassic (Permain specieshave been assigned The Late Permianrepresents approximately 13 to a different genus,Merrillina). The normal UIAN million years.The Early Triassic (Scythian)is diversityof Late Permianconodont species was thought to represent only five million years. approximately five to six species, so the During the Scythian,there were at least nine occurrenceof this number on either side of the genera,five of them new (the other four being Permian/Triassicboundary confirms Sweet's holdoversfrom previousperiods). There were (1973)conclusions (Figs 11.4, 11.5). rt. FaEemed area = at least 40 Scythianspecies, 35 of which orig- !e.a foom Previous The size and diversity of Permian conodont inated during this epoch. The fact that there faunaswere constrainedby an Early Permian were five holdoversfrom the Late Permianhas extinction event (Clark 1972, Ntter 1985a,b). -Triassic conodonts.I given rise to the idea that conodonts were Late Early and later Permian faunal diversity is I reports of speciesand relatively unaffectedby the 'notorious P/T fil- a reflection of an adjustmentto this diversity |s approximately 100 ter' (Sweet 1973). This idea is easilytested in reduction. Probably there were severalminor lr rhe Triassic.While terms of Late Permian-Scythiandiversity. expansion-reduction events during the [ate. the numbersthat remainderof the Permian,and the pedod cul- :d to be an accurate I1.4 PERMIAN TO EARLY TRIASSIC EVOLUTION During the Late Permian (taken to represent 9co some 13 million years) there were at least 50 o30 species in eight genera of conodonts (Figs.11.2, 11.3).During this 13 million year interval there probably were never more than approximatelyeight speciesliving at one time. c to Most of the speciesliving during the Late Permian originated during the Late Permian; 10 two of the eight generawere new during that time. During the succeeding15 million years -0/ 6 \,'"a 6 T (Scythian to Ladinian), there were approxi- o) IIAN mately 70 speciesthat originated and 12 new genera. Fig. 11.4-Species and genericdiversity, Late Permianto Compared to the previous almost Norian, on true time plot. Upper pair of plots : specific equivalentinterval of time (Late Permian),the diversity,solid line = total speciesliving during thatepoch, numbersof generaand speciesof the Early and dashed line = total new speclesoriginating during that epoch. Lower pai: gene c diversiry, solid line : total of new species Middle Triassic represent a 100% increase. generaliving during that epoch, dashedline = total new Fcn.-her This raises the question of whether the Late generaorigmating during that epoch. ler reqt qrv'r s1c,b ree,( aql Jo uo[€rrJrsJe^rp lselea$ aql lrqrqxe 'uersruy) s1e,rraluratuu pnbe pue Surpaac r or :Iqrssod readde 1ou sluopouocuprqlurs 'slaleJuzrlpeqseug o1flrs -cns Suunq'peJJncJo uoq?trJlsJe^IpJo Je^el Rfr uoueJ rlrsJe^rp .srsuc -IaAIpluopouos uErrouer(I eql pSurnlersuorl?ur sql leql eu41Jo le^Jelln ls€l aql s€/t\uelql{ts pue I q relJe s:eaf uoqpl -Erro sercads/aeu 'saneds Jo lJols reao eql u€rJoNegl olur po^l,rrnseuo puz uerq1,{cg rco eraua8 pue serceds -ploq uerreuarq rellPurs€ ur psllnssJ uerqJEq aql pe^r^Jnso,tt'1 ,{po 'eJeue8JeAoPIoq JnoJ (lrsra,rtp 'erauaS Er!.s PJPnlJ lEql uorlsc -serJg eql po^\olloJ l?ql uotpnper eql 19 rar'oplorlJnoJ eql qlyr\ reqle8ol qI -nuopouoJ Jrss?HJ arll '(9 II'3rg) sar.repunoqa3e uerq{cg aq1 perrnccosuorleurEuo crrauaS e,lrg 'plratur srql rsDal srql'ssalaqleuoN Jo qJeel" uoucnpe.rdlrs.ra,up ogrcads s€A{ aJaql ;o ged lsaqreeaq1 Surrnp serceds re^oploq a^rJ uerql,bg pue crlcdcser'r uerqlf,Js aql Euunp uoqnlo^A eql Jo euJosls€el 1€ qlrfi\ SurArT'pa^lo,ra sercads 'aurq r- p*rrudun-iersqns u€ uo Paseq crssepadpeg ,nau96 ,{lalerurxorddep,rralut srql;o srea,{uotl r .:i uorlru gZ'I al€ur Jo sJea,{uoqlu eArJur pe^erq3ese^{ l"qA{ 1ru e^rJeql Euunp pue 'crss"uJ,,(lJeg eql uI rf iEu sraqurnu serJads op 01uoqnloae rlss?rJl elppq l Jo sreef uoqlur prder se,,,ruo4?tr1rsrelrq (tr'1 y'Etg) uetureg h 4'ri1 srqt uaql 'sfte,{ 0I loor tI '.(t'tt 'Z'tl's8rg) 3uo1 se ect,t.l ele'I eql ur palstxese sorcedspup ereue8,{ueur E-Ea-r uErq JS eql JI lsee11e flluapr,ra se,tr3rsser{ elppq l oqt tnq se qln ue8eqsluopouoc Jo uorlnlole uelqlf,cg Ssvul appgal) uelsruy 'uelqlfcg eqt Surrnpse reqrxnu aru?s eql lsourlz l{gqurs eJe^{ pu? slaael 'suorlzur?rro sarcads gg {yalerurxordde pue NOIIN1OAS CISSVIUI ATf,VS S'II E:::: Sraqwnu sarlads eraue8aleu ua,res eraar e-req1(crsserrl alpplW:) | t.i? rJqlo Jo asoql s3 uelulpu'I pue u€lsruv paurquJoJJql ?uunp tqF1}. ursre^rp luopouoc 'p€J uI 'u€rqtfcs eql Jo t?rll Jpq euo ,{po era,lt 'rrsserJl aql olul pear^lnsel\IJ ls?el ;-:: uerqred5-uery1rurg uorl?rrysJe^rp 1o seler (ueture) 'uerurp"'I t? qcrq^\Jo sarcedsuezop Jlpq E qlr^\ petEurur IJ Frrt;--u3r-Uoqs[re^ ' 'tou rfua iurpnpm 'ueq1,{cg op sleror esonnr puE 'spodorqJerqpltrnpord 'suEozo,{rqetellselle] 'splotselqsnoauerodueluor ellq^r pepeJJEun ,{lo^oelar uorlauqxeJo atuDrofeu slql e^l^rnssluopouoJ tue^e uollcurlxepue ,{repunoqcrsseya-ouired-9 I I 3u -suorlrpuocopl{r llrl :u_.'::_:iuEIqt,i.S q?ln PUE \ I rF-.: ,::i sorTlrurZ I ,{Ialelu r G ri--EI3rlrlarrp Jo olJ^c sql lEr:-r1- tLl lje S33eUeql,{cs ffi :-S puE rfiepunoq e8e mffi ffi IEEL- :Tar\ '_\lrsr:^rpe^IleJer ts- -,qig"l) ueql,ics oql 3ur trn- ;:6351srrlr,{J-9 I I 8rd ffi- # ffi .*cg J^ R_. ) wRI w w d @ --_-/6)" A @ tu @ ctssvIUI NVtnu3d [9 II ces I I 'qJ] sJeaf, uolllJrll ,{ g lBuU aql :stuopouoC OLI lch. 11 Sec.11.61 Middle to Late Triassicevolution 777 Anisian Sepkoski(1984) and used as the basisfor the TRIASSIC various Earth impact models, but the unique speciesproduced at the beginning and end of Spat hian this conodontcycle are of considerablesignifi- cance in conodont biostratigraphy (Ritter 1985a). Smithian The problem with all of these plots is the crudenessof the time scale. Exactly during what part bf the Scythianages did the species Dienerian become extinct? If all of the speciesbecame extinct at the same time, a different interpre- tation could be expectedthan if the extinction Griesbachian rates were random during an age.Similarly, if extinctions are concentratedin a part of the age (somethingto be expectedif there were a Late Permian sudden environmentally related cause),rates Fig. 11.6-Cyclic speciesdivenification and reductiondur- of extinctionper million yearslose their mean- ing the Scythian(Early Triassic).Width of figure indicates ing (Raup 1984). Also, the precise time of relative dive^ity. Notice divercificationof speciesat esch same age boundary and high Smithian diversification.If these originationsis obviously affectedby this Scythianages are of approximatelyequal duration, then consideration,and if all speciesevolved within this cycle of diversificationto reduclion covers approxi' a small time interval, the rate of evolution per mately 1.2 million yeals.Data basedprimarily or Svalbard yearsis meaningless. and Utah Scythiansections but this approximatesocean- million wide conditions. lir time of extinction dil t'rachiopods,and 11.6 MIDDLE TO LATE TRIASSIC Scythian,including many novel speciesthat are EVOLUTION very short-ranging (Can etal. 1984). The les of diversification Smithian-Spathianreduction left the Spathian Following the Early Triassicdiversity increase, d tle Sclthian. In fact, conodontdiversity levels at the samethreshold there was an abrupt decreasein the early Mid- Anisian and Ladinian as those of other age boundaries. Spathian dle Triassic.During the Anisian approximately Erere sevennewgenera speciesnumbers returned to early Scythian two-thirdsas many specieswere presentas had I species originations, levels and were similarly reducedto begin the existedduring the Scythian,and approximately 15 duringthe Scythian, Anisian (Middle Triassic). one-third of these were holdovers from the Bras evidently at least If the Scythianrepresents only five million Scythian.Species originations were reducedby lf. 11.3). It took 10 years, then this cyclic expansion-reductionof 50Vo even though generic originations Triassic evolution to do speciesnumbers may haveoccurred at approxi- remained almost constant. Ladinian species r fre million years of mate 7.25 million year intervals, but this is increased, but only partly because of new based on an unproved assumptionthat the originations; more than one-half of all Ladi- Sclrhian was cyclic and Scythian substages are of equal length. nian species(about 18) were holdovers from iy reductionat eachof Nonetheless,this pattern appearsunique for the Anisian. As earlier suggested,total Middle daries (Fig. 11.6). The Triassic conodonts. The Smithian diversifi- Triassicevolutionary rates were approximately I followed the Gries- cationthat includedseveral short-ranging novel one-half of those of the Ea y Triassic. The raller Dienerian hold- speciesand generaoccuned approximately26 cornbinedAnisian-Ladinian evolutionarypat- md new speciesorig- million years after the Early Permian 'post- tem is one of leveling out, a sort of statusquo irrerian mnodont diver- crisis' diversification (Ritter 1985a). It does for conodonts, down frorn the levels of & Smithianconodonts not appear possibleto relate this 26 million Scythian activity, and poised for the final cur- rft-enification of the year cycle with that reported by Raup and tain. No sisnificant Anisian-Ladinian bound- ,(pq311spa,n,r:ns pqr Jo sr u"ql/bs eqt ,(lelerurxorddeser'r uralled usruIPEI eql (?) 1sele1,{:a,r eql otur) ra?uol ',{ur euos sI k-Es aq1 1e^eltuereJJlp rad serceds e,req feur sluopouoJ l?ql uoqsaEEns 3ur,r11serceds JnoJ lf :r"-I aql 111unpel€ader g'g pue ,(rured e-reua8g'g iqcodesnonard aJaql eurrl leql Euunp ls"el (€) qEnoqlp raIIrBe lfc eqr Eur-rnp satceds eql Jo leql Jleq-euose^\ uollnlo^a u?rsruV le erar\ ereql '(,{Eoloulrrral ar-::E:dde aq1',{1r,,r.r1ce 'uoqnlo^a luopouoc Jo ueqa?qg) uelroN lselel aql e^I^lns lou r uo pue (9 11'8tg) ;o aErnsroferu lsq eql s"ra srql ,(ru red prp sluopouoC (ggOt rt.ntC) aleq,t\eslapsqslT " rT q pelalduror se,t suoqeurErroserceds 4 pue crlaueS,tau -qnd uaaq e^€q pu? eutsnv ur suoq '-l 9 "pe^aN Frl aql uI rnq :(986I euo 'prder aJonr s€at uoqnlole uelqltcg (7) -ces eurJer[ JIss?JnIFrsseuJJo serpnls u.IorJ b ::qro ur u/aoqs uaaq 'tsul pe^rrapsluopouot Jo uoutullxe eql Jo slrclao rp sqrqad ,{qpageuruual -unrd crurouoxeluerrura4 fpeg eql ruo{ NOIJJNIJXS p sqogad ,t6.oqs,{lrcelt pallnsar ,{1rsre,rlpa\ol Jo I"^felu sIqI 'dur pue ''U ; =trrnrd ol ,fuzssacau rad sarcads2'7 [1a1eu1xotdde sJea,{ F+1. se aq ol luElcnlgI uoqgrrJ xrs red uorleur8yo cueuaE,lrou dla^peleJ eJe,rr 'crsssrJI tp -,!a:-"3raldrurs sql ro; auo flelzurxordd? '^rol r r,:rrrri raqto ,(q u,troqs sele-rdreuoqnlo,re luopouoc uErtuledal?T (I) ele'I eqt Surrnp pacnper dllearE ere ,{ur red suorleur8uoeraue8 pue seIJedSetu€s eql F: \JEqOIlnlOAe le^al :apEurJre suorsnJruorEurrtrol y ,(leleulxordde sr sleed uoq1lu eqt f.-I :o uosueduroc :ql 'aJnlEralII JIssEItI ,\tJt el e^tlJa[qns 0g I?ug Jo loJ Jo ged rrsseul eql ,{po ro; ernSr; aq1 ' f,wfercue? e ,(q paurelqozlep cuaua?pue cgrcadsaq1 1o suorleur8rrocuaua8 pue,(ru/sarcads .rEa i uolJl[ur 9I pue sa8eclsszul eql ,{cemcceeq1 Sulunssy €'0 Jo ;o uogznads ueelu peq sluopouoc f:-- ;zq satoeds,,,reu 8'Z Jo seleJ sreeduo11Jru.r purl eql Eul t +.+E 'lsrea,i uollrur SNOISN'ICNOC(INV AUYIAMINS 8'II 'uounloAaJreqt Jo 6g ,nau etuesqpse,{ G reu I pue snue6 -lnq suopeur8rrocueue8 1o T. :f{!- .\rBpunoq crsseua i(1.regeqt uro+ sure ed ,fteuo4nlo^e e8ueqc aql ',{gza13 lf :(:r rE peuncJo ur luecr;ru8rs lsoru '(9961 poors '(986I pue4 srql ernl?u reeull 4: ;sl ]o ploqserqr 1re13) {r€lJ) Jo 3s so'u aqr sd€qred (r) -Jepun sr r{Jrqa{Jo auou 'suoseat lueJeJJIp[te^ eql elsJtsnllrslPp flure; pue cr-reue8roJ sa^lnt eu?Jaq cueue8 g1 pue E:l :qr Jo asol3 eql JOJarXrl au?s eql lnoqs 1€lsurlxe Euop drqsro,u,rrng suorleur8uo sql sdeqre4 (uerurpel pu€'u?rsruV { F ?ue 'seas u?rloN -ouoJ Jo suon"Fdod leurs rno; sarceds99 aJeA\eJeql rc ro satoads e,rr; 'sluopouoc ,{q palpueq 3le1r{JIss"I{ pu? 'uerq1,{cg)s-reed uolgrru 91 snor,,r.erdaql ul tnq :1 'ueqric5 aqt ut crozoeel?d aql Surrnp sa8ueqc^IS?e le11ulg 'apnltu 'uoqpur8uosnue8 auo pue suoqeut8uosetceds plueuruoJllue uB eJ3.jAaJeqt uorl[ldJ = uelloN !s rr:aua;- fueur se I -8eur lze-r8 ;o e8ueqc lou LZ '(srBed ZZ rl .\-Eil ?sE srqt SuunP s?^\ srql'sluopouoc ou pu? eun€J Jrqlueq pue uerule;) f,rsser{ ale'I eql Surrnp 'i(zar € qlr^{ suouPuoc ,(3raue-reqSrq 'reA\olpqs reqtoue (paulqurocu€IuftX pue'uelulpe'I -s-r. iue se 3uo1 se tn4 peceldar pue peSueqc (sluopouoc 8ur Surpacardeql seSuoJ su) Jee,{ [ :rJ -iu] :ed setceds qtr,u se,r 'uersruy 1zlrelur eune; crSelede qlIA) suotllPuoJ uol rru z Suunp poleurEuosnueS,tau auo tT -fooeu6uo snue8 -pnlcur 'Ieurseq lI seeJpo^il eqt puz nau ua1f,po lpuert slql panu[uoJ 4. -l JrssEuJ ueuoN sdeqrad 'd8raua-,nol teql etErlpul serceds 1 s o-uol se lsorule) ur sloor eleuoqJ?J eqa a8ueqc Flueuluolrlue (uerleeqg Surpnlcur)u?IJoN eqJ suoDeutSrro \-rq! Suunq 'uor1 ue se elxll aul?s eql l3 Po ncco pu€ cuauaS ,teu ou ere,n ,(1luapr,reeJaql '(sreed r _: "p?AeN : q--.od3 uerroN aqJ, (9) €rJlsnY uI eluopouoJ eql Jo uollcuDxe eql uolgur g) orult ]o s1e^relurluele,tnbe puz 3u1 .(€86I psleurElro G ;-a 'ploq UBIUIP?'I IIPIC) -pacardaql ur sz uerule;4aql ?uunp aqt '1aa.a1serceds suorlelndod leurs ,{ra,r ut serceds rno; {po serceds;o Jequrnu eures oqt dlal?ulxoJdde palsrsuoJ pun€J uz,{q1a1 aq1 elrq1\'Jrss€lJI ete'I eqt Suunp per:ncco ;r5-n iq peurElul€tu Jo tuopouoc leug pecnpe-r ,{11eer8 ur e8ueqcpunogord aq1 Io\f -srea,( uoqlrur pu€ 'u€rroN tselel eqt Suunp sal?JuopnloAe lsour E-Ll truaua8 ur e8ueqf, sea\ ecuEpunqv slueu[uoc uee,rqaq sarJe 'aurg aruesaq1 1e 3ur,r11 sarcads 1q3ra lnoqe uerll puP saues uouelauoJ rood eJorl uaeq e^€q Jeaeu,{"ur aJeql sJ?add? F uoonlo^a u"rule) (9) -punoq Jrrlolsds ]o lBql h .IrJ sarcedsp g Puz Jo peJqr€ ue aq feru sql lnq '?clJeurv quoN 11 tuaredde ere (9'11 3lg) se8e ueqlb5 'qr leql sEarl€s eql ur prp,{eqt uer{l eere uedqleJ eqt ut (ueuo51 eql Jo esoql sE qf,ns sesllJ ,(1rs::,rtp ,{:e -lo II'qJ] sJ?ef, uollllur &Jg tBug aql :sluopouo] zLl lch. 11 Acknowledgments 173 aEa than they did in the sameas that of the Anisian; 0.6 genera the late Early Permian, and these two events r raI be an artifact of and 3.4 speciesper my. are separatedby an interval of approximately r and systemicbound- (5) Karnian evolution shows a remarkable 26 rnillion years. G- Abundance was changein genericoriginations, none in five Overall, conodont generic and family level ft latest Norian, and million years. Evolution could not be evolutionary pattems are consistent with the ht fauna consistedof maintained by slowing down. At the model exhibitedby the Red QueenHypothesis lr small populations specieslevel, the samerat€ as that of the (Clark 1983), althoughthe data are so few that Ladinian held, 3.4 speciesper my. a cornparison is quite subjective. On the rconodontain Austria (6) The Norian Epoch was a time of extinc- species level, there is less consistency in t tt€ same time as an tion. During this 17 million year interval evolutionaryrates, however. The Late Permian lle carbonate rocks in (almost as long as the combined pre- evolutionary rates are greatly altered during I lo*-energy, perhaps Norian Triassic), there was a single new the Scythian.The following Anisian to Kamian e pelagic fauna includ- genus origination, 0.05 per my, and 0.8 interval has consistent rates, but these are ad wasreplaced with speciesper my. The Norian is three times much lower than those of the Scythian.Thus, y conditions with a as long as any other Triassic epoch, but the perturbations of the subjective specific Gmodonts. This was during this time only ] asrnany species and evolutionary rates may be interpreted to sup- lange of great mag- ] asmany genericoriginations occurred as port the nonconstantmodel of evolution (Sten- the Palaeozoic in the Scythian.There were approximately seth and Maynard Smith 1984, Wei and Ken- bndled-ring by conodonts. five species of conodonts living in the nett 1983), while the overall picture derived lopulations of cono- Norian seas,and all becameextinct before from a generic level analysis(which may be lout the sametime for the closeof the Triassic. more accurate),could be interpreted to sup- me of which is under- (7) Perhapsthe most significantevolutionary port the Red Queen Hypothesis(Fig. 11.1). threshold of the final 50 million years occurred at the Middle/Late Triassic boundary. After this time, only one new ACKNOWLEDGMENTS genusand 27 new speciesoriginated (in 22 CD\CLUSIONS million years),while 12 new generaand 63 I have reviewed almost 500 Triassic and Per- new specieshad originatedin the previous mian publicationsfor this compilation.This is f the Triassicages and 15 rnillion years essentially all of the Late Permian-Triassic tfo data obtained by a literature published,almost all of it in the past xic literature,the fol- A comparisonof Triassicconodont species- 30 years.I de: acknowledgeall of the authorswho level evolutionary patterns with patterns have described these species.This includes ht evolutionaryrates shown by other groups is not really possible, graduatestudents who produced15 thesesthat r, approximately one for the simple reason that most studentsare have been of great assistance to my tion per six million reluctant to be as 'objectively subjective'as is Permo-Triassic research during the past 20 2.7 speciesper necessaryto produce the data. Conodonts years. These students include L. Cameron -E\-b*- diversity resulted clearly show periods of rapid diversification Mosher,Fred H. Behnken,Laurel C. Babcock, taxonomic prun- terminatedby periodsof extinction,just ashas Mark A. Solien, John A. Larson, William A. -rn been shown in other organisms(e.g. House Morgan, RachelK. Paull,Tim R. Carr, Eric W. fas more rapid, one 1985); but in the Triassic,this cycleapparently Hatleberg, Reed H. Meek, Ronald R. Char- 5 species originations was completed in less than two million years pentier, StephenP. Carey,and ScottM. Ritter. E last major suge of (Fig. 11.6) and on a much reduced level of The National Science Foundation has sup- actiyity. The appearanceof novel genera and ported most of our work, and this includes G one-halfthat of the speciesduring the middle Scythian was not current fundins from EAR-8205675. generaper my and 3.6 repeateduntil the Late Triassic,and then on a different level. The morphological variety of n Fas approximately the Scythianis of the samemagnitude as that of 69I ' snwouSortcog G:: _S:':epnuopoqlsu€o4ceg 'Jlopuessenltpue "c g c's33ug'c ('clln{lhtr '0zz-8lz soe atntDN'etellurwetol 'ZZ-LI E11EarrirN suoqeqrnlredf,:eluerutpes .ruollueld .rueSoeN Jo selsr uorl.uDxr pqol8 qfl^! slue^e fueuopnlo^e ptououjllle luelsuocuoN €g6I d f ']leuue>I puP I-I 'lend. Jrozoeeled-pruJo uorlelel.loJ986T U 'aSnOH 'OE-I I KrcaLUtQouoqnlo|Z '{ '€€-6 'uoqnlo,re 'Lrel rfieuorlnlo^. ,t\es V €16I -'I 'uel€A UEA 61 ,t?o1otqoa1o4 satcedsqpruJo slepotu '9n9-089 Z tlotuaw gstdoloaDutnap) ;o 1se1crSolorqoayed e :uets,{sc4{ueld clSelede -tad lo ttapog uDtpDuoJt(spe) sllrH 'A V pue ur uorlnlo^e tg6l V f llaqJlr) puP V'ueuJJoH ue8o.I'V :uI ,{lepunoqlenlnul neql pu€ suelsds 'dd I€I 'sserd,{ltsre^lun e8puqtueC'seuas crssPul puP uerurrad eql ur 'sEungJluopouoc ecuercsqu€a e8puqu.reJ'aPJs arau )8opa3 v crsserJlflreg puPuerured ete'I g/6I J l[ 'lee^rs '286I d 'srelpi[ puE' o'v'qllurs' c v J'uol 'LL8-018 8E uotnp^g Lslse$rc -{rld' c 'd 'uffle1riel-I'A v'xoc ' g /ld 'puelreH uoqnlo^a ueeno peu :suelsr(soceuI uoqnlo^aoc 'Igg-Z9g LS [SolotuoaPdlo 't86I f 'qlFus prEudEr{ pue 'J N 'qlesuels punol'sluopouoa lo uorlcullxg €861 1 C'{rEI3 '/z 'uoldujeqlnos Jo ,{lrsle^ 's6I-t6I S€ ualL llDlsuosap rLqcl s:ee,{ uoqrur ,{grg pug aqt :sluopouoJ vLt lch. 11 lpdied brota. Science2X D$ade of North Ameri- !r Time Scale.Geology lkrnary radiations and fran and A. F. Trendall 4c ia Earth evolutidn, IBJ- Springer-Verlag, i,I- J- 1984.Periodicity of jcpst. Proceedingsof the irz El 801-805. rilart' divenification of i Sn-eetognathra ir the TaxonomicIndex *ed States. Geological ftrt 17 2. frnogpic diversification i.ge: a biostratigraphic Ebser Permian. In: R. J. "ll'7, rd \1. P. Smith(eds.): Acanthoduslineo tus, 1l2, 116, I20, 1,23 Bellefuntia chambe aini, 117 ?Acantlbd.w Cood>nt Symposium uncinatus,112, 116, 11,1,120 Belkfontia sp.,117 Acodlu, 116 .I :- Abstracts,Uni- Belodello n. sp. a, 131 Acodusdeltatus,l,16 Belodellasp.,50 .n- 'A@dus'oneotensis, 112, 117, 119, l2O Bebdina,9l, 100 trard Smith, J. 1984. Acodu,s sp., 11'7 Bessebdus,91, 101, 102, 103 E Red Queenevolution 'Acontiodus' iov)ensk, 112 B esselo&u arcticus, 91, 100 ll)-s17. 'Aarntiodu*' iowensk-propinquu,r, 117, 120 Bispatladur , 158 lrnian and Early Triassic Acrctrcta sp.,ll,'l Btunchiosktma,24 : PerDian and Triassic Ade tognathus lau tus, 160 lbomdary. In: A. Logan Adzlognathus spoIhus , 160 Camioduscamulw,131 Ade tognothus f-adun Society of Pet- unicor nis, 159 Cavusgnathus, 160 Amy dro taxis jo hnsoni, 152 Cavusgnathusahus,159 a 2 630-646. Amy d ro taxis seiden tata, | 52 . EE evolutionary law. Cavusgnothus naviculus, 159 Anaspida,28 Cavusgnathus unico rnh, | 59 -3{}- Ancoradella,l32 Cephalochordata,88, 89 L P- 1983. Nonconstant Ancoradella ploeckensis, 130 Chaetognatha,35 \eogenic planktonic AncyrcdelLa,152, 1,55,156.157 Chosonidina herfurthi, 112, 12O, 123 i2rE-220. Ancy rcdclln binodosa, 15 5 Clawhamulusdensus, 112, 117, 119, 120, 123 Anc y rodella rc tundilo ba, I 5 5 Aebndio utahensis, 111 Anc!tudelbides, 152 Cloghergnathus globens kii, 159 Ancy tudelloides kutschei, 152 Clydagnathu.s, l5E Ancy rodelb id.esomus, 152 Clydagna thut covusfurmis, 159 Ancy rodelloides tigonicus, 152 Clydagnathus1cf. cavusftmis, 23, 25 Ancytutnathus, 155, 156, 157 Clydagnathus gilwernensis, I59 Anq rctnathu.sanclrognathoideus, 1,55 Coleodontidae,102 Ancy rolepis wallisei, | 54 Cola us,102 Anbgnathus,l5S Conodonta,23, 24, 28,79, 1O2,105, 165, t61, 1jO Apsidogna thus I ugino sus, 131 Conodontochordata,22 Apsidognathustuberculana, 131 Codybdus, 40, 42, 44, 91, 100, 102 Apsidognathw walmsleyI, 131, 1,32 C,otdybdusangulatus, 172, 11'1,119, 120, 1,23 [email protected], 102 C-ordylodussp., 40, 42 Asp elundia copehsis, 131 As trcpentagnathusirregularis, 131 Dapsilodus? sp., 50 Aula@gnathus, 132 Dapsilad.usobliquicos tatut, 96, 131, Aulacogno thus bullalus , 131 Dapsilod.us p r aecipuus, 131 Aulacognathus kuehni, 131 Dapsilodu.s spa rs us, 131 Aula@ghathuslatus,131 Dechnaqhathodusnodulife tus, 160 Decoiconus ltagilis , 131 Bactrognathodontidae,150, 159 Dentaadina? sp.,96 Bacttugnathus, 159 Diaphomdu"sdelicatus , 112 'xe4utls snPot@uo ItUaOolOrSlrpOlJ tI 9aI 'snqlouSortDd OZI ' LII L9|' 'zII ' stsu2uoslDpsnpoloeuo 'xttlDl t/lpoltx( _.-. d\ sltpdoqtaldolDd 0zl z9I ,0zI .9It,snpot@uo ' ' snsuodra snpourl a- - t4pdowaldDrDa ezl Z8 18 'sJu{llP snPolltl _--' :-:.,.]23 ---:.126 sUpdowaldotDa 0ZI' L'gll'ZIl'suofl8uo14,snpotslo, tSI 7lI 'srunrq s[rlDlDd tZ 'tZ 'snpuo erqdtlSotuoPo III OpnOJlAJnaSnpOU)J snpolnl t'|ioljou mpoqlDuS,roa EZ 'snl&)Soruopo vgl 'ra3lutor - ;' mpou)I \ :,--:.,- :itrioq tnpoqlouS1tod t9I'aolpnolr snpoqrDuSoqt DdsoaN snuo4lsoxsnpou)I ----!u@ mpoqlouSotod ItI '!u!dttl.!r z9I *;- st^atq mpoltcl \9I snpovouSDtDd 69I 'snpowodsoaN lsl ' snpourl snporepuDdotod O9I'snxu P1utuKs snP oqt ou8 oaN ZSI'sryDtuaplqsaplotsnSuD I6 snpott.I -:-. .-a do DuqqulropuoJ 8II 'LII 'Dtots@lFu s!41to11bN z9I 'aDlotlD gII I s!LtltouDN '1.9I'9SI 'ngr 'Z9r 'r0r 'r8 '08 '09 'LZ 'snpqt.I rc --: .'];rD oullaultopuod Lll 'slsaafunEuoq 8ST -- '18 'Lz 'aepquopoutl a;I oul auuopuoa zEl 'leapuos :'4 -;'.:--.: tor)un snpolapuod 8Z 'Peproulx^I^l Itl Lonapouzl - gz'aulxllI suo.ul qlaPouzl _: iI i ds snpdapltDd IEI'Suot d svdotSouow --: i q ds snpotaPu,d 6eI 'uoPo Ir1loaull ttustl4 sryd'tSoltow Lll ' ds snrfi ttstrH rtt + -:..rn) snpo/apuDd 8eI gEI'LtIt nual m)Mlrqoq s4 alorSouol\l 'mtolou,S4tDdsntn uIsf,H i { .--i'w!\8tnl mpoJ2puDJ LII 'nnro18 'stl ua4oq sw dotSottol/! gl| ' LII 'sntqouaSsntrculs(H - :: mpatapuoA LEI : ':.:.ar!nba mporapuDd 69|' u h4lq snqtouSo$a I 1 8Il 'Sfi1nrulslH DUBoJ saw ds rnPoePtuH 7- :- snpotaPuod 69I' tuuDuq)aq snll 6SI|snln - ggl t ds v slmr osall i : -iardut@:-16 Ba snPolaPuod 691'snrytlcs woePulH 1ta14U1{SOS|XO|OSaW snql Dsauad snPoaPu.t -.__ :.1 ualstDq,snpollDd, 99I 'Sn 69I' H -:l :-,.,,;!ou1 sldaptoulod 9SI'stxDlos2W o9I 'ffi|nutut mp@pulH _. s11pozpu1H a :,.:lstDrt sldapl Du.4oa 691'bu\luraw 69I 'sqqsux - ,0zl snPotaxol owpoaputH | .; t:. :1.anuzt std2loloutDd EZl zII'ao11Dlp I sol'suanl[u@ ' ollzpoaPutg : .::: l adqns sldeplDrullrd 99 mpoxol 'earellePleqqrH .:Drqns sdaploutDd EZI 'OZl '6lI ' LII 'ZlI 'tuosuuq ,z (rCeJlSOrelaH -:- 'q ds oulpouo3lI _: .:.cq|nb edzlotoltlDd z€l 'I€I 8Z .:nutw edaptolulDd ZtI '|tI 'ol1ladoulpouo8tj _.. gz (EleuotsoqlEu9 - :--, a .'6ull ndaplDlulod IEI 'Ia8an{ DmPouoSl'I -: s$uablTfttu^l p DulpottoSl'I 'snuoxatsnpoqtouc _. \'a3'3 sldapt'apd Ial ' 'I 6SI - lu 13e$oD'7 e qq8nuas opnas d snpoqt out _ a'. :'-:. r, lap sldaptoll\Dd 8II ' LII 'Slsaanolluolu 69I' -.1 OgI' 1su133rqsnpo4t ou t : -,: .;_ 08 ' sldalotDlt|od : -- .:lrLod snptb@|Dd 6tI 'lefl DiAD\apltot O9I 'xaldutls 1&!8 snpoqlbuD z€l '1tl sllqutoA o rpltoN 6Sl'!{tl!3 snpovoug o aP4ro)t 'ZB 'slsuaxopousntoeullq snpoqtou) :r-, !t3,.rsDulpo4tozo ZeI 'I€l 'stutt4\nu 98 : . :'-: r otl u Dl s DulpolDZO 6EI'ZEI'Dlryod o aP)PoX 69I '89 't9 '89 'snl'aulflq snpoqtou{) - D ePl)ox .69I ,18 '08.sflpolt|Du9 :: ! \ou ds DulpoqDzo 6El 'Dtoplsqo 09I DuLaddolx '9II 'ZIl 'mtDld ponb snuaqdtlC ::. P ds oulpolr'zo 991 'sq'^o tZI 'OZI -:I odslP ouu ed dolx ,IEI ,62I,.^ou dste ue8 :- : : up[ous DutpalDzo I I l'slqt ?El buuaddolx .;1 qlas oulpolr'zo ggl' DtotodstP g De DdstP DuIl 2 dd,ll 'LE' ds o l\slutnl - --t.:.DuuDs oulpoltDzo I l' 8€ - ^Pt ggl'tluueddDlx 'o|q|t t ::- ,auada! oulpotlozo 8t lqslutnl DulpoltDzo LII ' ds DIeulDX Le 'Dulltsluni :.':):L'lPswat aqqwtol + .-.'r|t)u\od Dulpolr,zo 8ll'1s3u1 lq x'!. o auPx 6Z'D -. .:.::tzDDnDd DutpOIrDzO -a .1oPuDd DulPotlDto IEI 'DtW 4 snl'ou8oullot 8Z 'EprdsEele8ng - -:-. snqou8oul|ot '9II 'Zll ' sqa otbd snpdolltllg : 't6 DrpDl.lDulPoltDzo zel ' EZI'O?l'Lll . - .1.ilan1n8 oulPollDzo 691'snr\dotq! mlonuls s?p|o\tou3olpl Isvtcltls snpo1lt'u8o4 : t; ?lt tDxxz oulpoltozo ogl ' zsl 'qpuq snpoqt'u8s! ; -1- -!-, r'4 ou uDl soa oulpollozo 09I'eEp0uoPoqleu8olPl l9I a ::rli- f.tDtsoa p oulpoltozo zgl 'u plul.lrso$ !1plluqlsotA snpolltl v9l 'snpo4tt:tu8oJ -li!)qdao7 Dulpo|ozo z9l ' qprutqrso^tsodupluqzsart srlPol!4 pp'gy ' ds ntuoPouocog - I snpolJcl iI l: -: !: slotq oulpalozo ZSI snuadsal.!|prluqrso^ -: snpourl ds snpotslou'd 89I'iSI z9I' !1p11u4rso'a LII' O 'uruDaut^ snpou)I '16 9 jj _t '6r'91'DulPoltDzo z9I 9lI 'snpo$touDdale -, snPollxl 'aou{lloe 96 snpqno vgl 'zgl 'esuaqcttula|s 69I snpolnl 'm\lou8oloQ 96 11a3an["nluad LSnPqnO zgI 'lotolnuf 691 '.a snrotnSltDsnpqno zsl' suDpsuq )ar snpou )J I€I's2plovtouSotnots elpoutotsl.l gz nErlsoolso z9I '!?ztolsnPollJl I€I ' snsopou mPauotsl(l suarsatllt4tl snpo rl zEI mpou]olslo : _ .l ;I I ' ds snPoloauol l9I ' xepul JlrlouoxBl 9LI Taxonomic Index 2on@bdus sp., 112, 117, 120 Pedavis bieto ramus, 152 Osteostraci,28 Pedavis mariannae, 154 Oulodusangulatut , 2l PeA)vispesavis,l52 Oulodus'l fiuegeli, 96 Pelekysgna thus ser ra tus, 154 Er Oulo&rs petilus , 96 Pekkysgnathui ser tutus ebnga tu.s, 152 Ozarkodina,16, 49, 51, 55,56,60, 63, 80, 81, r05, 152, Petromyzontida,28 154,158 Polygnathacea,16, 63 ., li: Ozarkodinabrevis,55, 81, 82, 154 Polygnathodontidae,1 50 Ozar kodina campbelli, 159 Polygnathoides, 132 Ozarkodinacf. eosteinlbmehsis,55, 56 Polygnathoides 130, 141 -,1a.152t- 15: siluicr.6, Ozatkodinaeos teinho mensis , 49, 50, 5I, 52, 55, 56, 81, 82 Polygnathu:,51, 56,80, 81, 86, 150,154, 155, 158 Ozar kodina excavata, 55, 131 Polygna thur angustidiscus, 15 5 Ozarkodiha gulletensis, 131 Polygnathus ansan6, 154 Ozarkodinahodra, 96, 131 Poly gnathw asymmet ricus, 155 Ozarkod.inopando rc, 152 Poly tna thus b rcvilo minus, I58 Oza I kodinapoucidenta ta, 152 Polytnathus b rcvis, l,5 5 Ozarkod.inopolinclinala, 96, 131, 132 Poly gnathus @mmunis, 158 Ozarkodina remscheidensis, 152 Pol! gnath6 costa tus p ar titus, 154 Oza r kodino rcpeti tor, 1,52 Polygnathu:cisto h.s, 155 Oza r kodina sannemonni, 1,5 5 Polygna thur decoro sus, 155 Oza*odina se|fi,154 Poly gnathus d.ubius, 1 5 uzafKoatnasnalaln, 5|J,5I, Jl Polygnathu"sglaber, 159 Ozarkod.inasp. a, 131, 132 Polyqnathuslatihssatus, 155, 159 Oza @d.inasp. nov., 50, 52 PoIygna thus limi tais, 15 5 Ozarkodina steinhomensis, 55 Polygnalhus lingufu mis, | 54 , Lr: Oza ttl ,0leqsuoung, :i -jorl?uuoJ eSueu lrng roFqS EZ ssoarng 0rI'suEozofuq i1 : -r_ur:l\ aSpug uopuEra -:: -60I 'gZ I - ! -: 'sPodolqrerq 0lI sprorsPlq ::::rqjslesary-1jlnirrlrs?g -: uollezrlerounuolq 6- -: !,rT til III'sereuorq IF .- :-: uratrEdprBpuets _r.: :g t ::'-:uEd relturpuedred :-:-ag uraued a! -- IelleEd ari ,:- :i::1?d pjepuelslosJJo c_ _ .!: _g rualled z reeurl ;_ _ !.r '_9 -u.iollsd Eeurl _ag I t _- -': t urallEdreaurl F:-:: ::::':rassp euEldSuppeq :: - -:.otsaurl qrlnc rea8 ; ..ilrqEulroJeSpuquEg L _- :- ;! rl: 9I'sllPJ ^alrEg II saqlullrrleqtse ,spodorqtre II dnorg ePlcnqrv I -:: ::: til :I ;I II spqeuue E- )-t 7.\1a\raorlo eptu@uv - !! _j tz 'snxorqdue _-: 91 85I 'sprouolllllle ,AE3IE II I€I ' ds snpdas\Pla O€I 'snDtuDqJq'ulfiu snpolqDuSoqrods, ItI', u Tolcu)u0s snpotaslno[4 gsl'D\orlns o aPouoqalls ItI 'Sut nJ SnpOJa9DlA 691 89| DtD4nszDldD apouotldtl ztl 'snporaslDta 89I 'rtl 'oppouoqdrs 0ZI 'snPoqucbDluasl rapul Ierauec zll ' sltuaulswns1^oulpotplrln ZII ' sntnlns ,snpodopxs, €ZI 'OZt ' dds ourpoL!41\n tzl '7II 'xat snpodqors I ZII slutolltflu@ snpodo(ns IEI' aouun al uDqt oq DIVDIaxntr 16'snpodqors tgl ' snPottoJ 'mqlou3olp1utq,s xapul JlurouoxBl 8Ll Ftirutus,132 l1- tl llt 11i9 tr t: General Index algae,11 collagen,105, 106, 108, 109 ammonoids,158, 160, 167 conodontochordates,22 amphioxus,24, 77, 88, 89 conodontophages,22 Ancoradclla pbeckensis Zone, 129 133 conulariids,11 annelids,11, 12, 13 Cool Creek Formation,ll2, 1,18,119, 120,121 Arbuckle Group, 1I4, 11,8,121, 124 copulatorystructures, 13 arthropods,11 cxaniates,25, 28, 102 aschelminthes,11 crinoids,83 crustaceans,30 BaileyFalls, 1,6,20, 65, 66, 61,70,7I crustacealspines, 12 BainbridgeFormation, 96 cuttle-fish dentition, 12 Bear Gulch Limestone,21, 22 cycles,147 -161 , 167, 170, 171, 173 beddingplane assembl^ges, 15-23, 55, 63-7 4,91,92, 98 late Siludan to Lochkovian,147, 152 linearpattem, 63, 64,66,61,68,70 Pragianto mid-GivetiaD,147, 152, 154 linear1 pattern,67,68,70,72 mid-Givetian to Frasnian,147, 150, 151, 154-156 linear2 pattem,61,68,70,73 Famennian,14'l , 156-158 offset standardpattem, 65, 66, 70 Touhaisian, 147, 158, 159 parallelpattem, 63, 64, 66,10,71 Viseanto early Namurian, 147, 159, 160 perpendicularpattern, 63, 64, 66, 70,12 post-mid-Carboniferousboundary, 147, 160 standardpattern, 63, 64, 65, 68,70,71, biomeres, 111,124 echiuroids,13 biomineralization, 105, 106, 109 Eke ma(I, 133 bisulca h.m-Kieselschrefer, 67 elasmobraflchs,102 blastoids,170 Estill Shale,135 bnchiopods, 23, 109, 115, 118, 125, 138, 760, 170 euconodorts,12, 14,28,29,35,38, 40-46, Brandon Bridge Membe\ 102,167 91, 92, 96, 135 Eureka 116 bryozoans,170 Quartzit€, BurgessShale, 23 Burt RangeFormation, 86 FaunaC, 111, 112, 114, 116, 118,119,120, 123, 125 'Button Shale', 134 FaunaD, 111, 112, 114, 116, 1,18,120, 123 Fillmore Formation, 111, ll2, 114-118, I21,124 Cambrianconodonts, 12, 23, 28, 35-38, 46 fish,12, 13, 19,20,21,22,101, cartilage,101, 102 foraminifers,160 chaetograths,1,1,, 20, 23-26,28, 35, 37, 38, 44, 46, 88 chordates,11, 23,24,26, 46,7I, 109 GasconadeFormation, 123 ciliatedtissue, 20,26, a4,86, a8,89 gastropods,20 clusters,16-18, 20, 21,28, 46, 49-61,,63,73, 78, 80-83, GeneseeShale, 15 86, 89, 91, 92, 95, 96, 100,101 gnathostomes,24 chaotic,51,52 gnathostomulids,11 openedout, 51, 52, 55 Grantonconodont animals,23,24,25,27, 44, 63-65,68, piledup, 51,52,55 11, 74, 78, a6, 88, 89, 93, 102 clymenoids,158 GEnton Sandstones,18 elEqs ele^r$otlr^{ .spBuozfruortod ,uoteuuoCs€I Z0I ZII Suuds lso^\ .uoseturoJ .leurrue {eetC 98 elleal z0l-L6 't6-I6,82 tuopouoceqser{nE^\ 9t 'nb 'Zn '8848'62'82 'ZI 'sluopouocered 9€I eqs re{cea, t8 'uoqeulroc sllIl{ u€rr^ rselerqauo^ ".SuraporBrlso 60I'6L'nZ "€I'II 60I ggl .^re.I sluel"A (requal,i{ ue^ t€I poo8so .l€I ,{p{uerocoquo 8€I .speg ,tqsr^ ,/rI r{le{re:{co 0€I reddl-l 8€I .uOU?rurOJ {uorteurod III eloauo €9 rel{J rslueuSes (proqcolou zI elrqoF-! tz ,uonerutoC .SZI .VZI ,ZZI 89r Zg rerdEN 'IZI 'AII 'LII '9I I 'SII 'III 'SOIIqOIq rZI 0z '.Euopoleqt Eol 'Z|t 'tL'92'nZ'OZ 'sprourxfur ,tZ ,setuoto,(ur II'seleFceluet 6g 88,tZ,ZZ,OZ,rI,seperuel 60l tI zI 'II'srsnllour !99I {sa^rnr €.zI Z4I drqsro^r^..rns 'zzl ,8lI ,uouErrroJ ,requjew 'Izl '0zI '6II 'UI IIrHerzue)cI I srl punupg .ls xapul lErauac 08I lLt