The Conodonta: Morphology, Taxonomy, Paleoecology, and Evolutionary History of a Long-Extinct Animal Phylum (Oxford Monographs on Geology and Geophy)

THE CONODONTA G-

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Morphology,Taxonomy,Paleoecology,

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WALTERC.SWEEL TheOhioStateUniversity

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New York . Oxford

CLARENDON PRESS. OXFORD . 1988

PREFACE

Conodontsare common fossils.Almost anyone who dealson a regularbasiswith Paleozoicand Triassic marine rocks has probably seena few of them. Through the last 30 yedrs conodonts have come to be of exceptionalvalue in biostratigraphy, and they now have pride of place as index fossils in many parts of the geologic column. But conodonts are extinct and are unknowo to most neontologists.The sketchiness ofinformation about them in most texts on invertebrate paleontologymay result either fiom the fact that they are microfossils that haye only lately come to be important as stratigraphic tools, or from the fact that nothing quite like them exists today, so their zoologic relations axeuncertain. In this monograph, I provide a summary of information about a group with which I have worked all my adult life. Charts that name conodont-based biozones and show stratigraphic rangeofselected conodont speciesare included as Appendix B. However, I have purposely avoided a recap of conodont biostratigtaphy becauseit is constantly changing and cunent views are readily available in various other places.Instead, I focus here on the conodonts as a group of extinct animals, about which it is important to know as much as possible before assessing their distribution biostratigraphically. Of course, one takes considerablerisk in attempting such a summary, particularly of a group of animals known only from its fossil record, becausemost of what I think I know about conodontsasanimals is either conjecture or a highly personalinterpretation ofa still-irnperfect fossil record. So, with the caveal that whal followsis only one way of viewingan im-

portant group, I ofer my account of the Conodonta. For their "witting" or unwitting contributions to what I believe I know about the fascinating gtoup of extinct animals described on the following pages,I am gateful to a long list of my students and faculty colleaguesat The Ohio StateUniversity, especiallyStig M. Bergstrijm, and to members of the Pander Society, an intemational group of exceptionally goodnatured "conodontologists"that has met frequently and infomally thougl the last 20 years to share infomation about conodonts, argue conclusions,and correct the misapprehensionsof its seniormembers. Karen Tyler, faculty illustrator at The Ohio State University, drafted nea y all the figures from my very crude copy and assistedwith labeling others. Dr. Jerzy Dzik, of the Polish Acaderny of Sciences, Warszawa, provided about half the stippled drawings of conodonts that gace various figuresin Chapter 5- The anistry of tlrese two good friends is plainly evident in their work and is warmlY acknowledged. I am also grateful to Sue Shipley and David Little, of The Ohio State University, for their help in completing various parts of the manuscript and illustrations, and to Mark Klefner, who graciously cornpiled information on the ranges of Silurian conodonts and assembled the Silurian chart in Appendix B. The Department of Geology and Mineralology generously assumedmuch of the expenseof drafting and photogaphy. Columbus, Ohio February 1988

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CONTENTS l . Inhoduction

l.l History ofdiscovery and study 1.2 Achievements 1.3 Pending problems

2. Skeletal anatomy 2.1 2.2 2.3

2.4 2.5 2.6

J.

Composition of conodont elements Structure of skeletalelements Shapesof element crowns 2.3.1 Coniform crowns 2.3.2 Ramiform crowns 2.3.3 Rastratecrowns 2.3.4 Pectiniformcrowns Symmetry- and curvature-transitionseries Skeletalapparatuses Symmetry of elements,element pairs, and apparatuses

Whole-animal anatomy 3.1 3.2 3.3 3.4

The Scottish Carboniferousspecimens The Waukeshaspecimen Histology of demineralizedtissues Summary

4. Taxonomy 4.1 4.2

Form taxonomy Multielementtaxonomy 4.2.1 Multielementmethodology 4-3 Multielement classifications 4.4 A revised multielement classification 5. The major conodontgroups 5.1 5.2 5.3 5.4 5.5

Introduction Cavidonti and Conodonti The Proconodontida (Cavidonti) and its families (Fig. 5.1) The Belodellida and its families (Fig. 5.1) OrderProtopanderodontida, new 5.5.1 Family ProtopanderodontidaeLindstrtim, 1970 5.5.2 Family Clavohamulidae Lindstrdm, 1970 5.5.3 Family AcanthodontidaeLindstritm, 1970 5.5.4 Family DrepanoistodontidaeFihraeus and Nowlan, 1978

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6.3.3 Siluriancycles 6.3.4 Devonian and Carboniferouscycles 6.3.5 Permian and Triassic cycles 6.4 Extinction 6.5 Iterativeevolutionarypatterns 6.6 Evolutionary trends 6.6.1 Apparatuselaboration 6.6.2 Apparatusreduction 6.6.3 Elaboration of elementsin P positions 6.7 Developmentalslralegies 6.7.1 Recapitulation 6.7.2 Paedomorphosrs 6.8 Summary

7. Paleoecologyand paleobiogeography 7.1 Introduction 7.2 Mode of life, or habit, of conodonts 7.3 Ecologicmodels 1.3.1 Thedepth-stratificationmodel 1.3.2 The lateral-segegationmodel 7.4 Selectedstudiesof conodont ecology 7.4.1 Ordovician paleoecologyof Cincinnati Region 7.4.2 Mississippian paleoecology,westernUnited States 7.4.3 Paleoecologyof Pennsylvanianconodonts 7.5 Ecologicgeneralizations 7.5.1 Depth as a factor 7.5.2 Ternperutureas a factor 7.5.3 Nearshoreand ofshore faunas 7.5.4 Phyletic changesin ecologicaldistribution 7.6 Paleobiogeography 7.6.1 Late Cambrian and Ordovician paleobiogeography 7.6.2 l-ater Paleozoicand Triassic paleobiogeography

8, The phylum Conodonta 8.1 A personalbias 8.2 Summary of conodont characters 8.3 Conodonts as invertebrates 8.3.1 Arthropod and annelid connections 8.3.2 Molluscanconnections 8.3.3 Connectionswith other invertebrates 8.4 Conodonts as chordates 8.4.1 The opinions of Pander 8.4.2 Newberry, Hinde, Huxley and Myxine 8.4.3 Macfarlane and the nemertineanconnection

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1. INTRODUCTION

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With just a bit of preparation,almost any ma- opment. The literature on conodontsis now rine rock of Paleozoicor Triassicage,from al- rather large(and growingat the rate of about most anywhereon earth, will yield to the pa- 300new paperseachyear),soit will not be postient investigatoran assortmentof phosphatic sibleto exploreeveryaspectofconodontpaleomicrofossils termed conodonts. Although biology in great depth. But this book is not named and first describedin 1856,thesetiny intended as a comprehensivereview for spefossilswere paleontologiccuriositiesuntil just ciatists.It is an attempt to summarizecurrent a few yearsago.Little wasknown of their geo- knowledgeof conodontsfor the nonspecialist, graphic or stratigraphicdistribution, and they who may well have been wonderingwhy this were classifiedaccordingto a mechanicalsys- group of tiny fossilshas come to be so importem based entirely on shape. Furthermore, tant rn recent years. their relations to other, better-knownanimal groupswerehotly debated. 1.1 Historyof Discov€ryandStudy Nowadays, conodonts are mentioned in most reportson Paleozoicand Triassicbiostra- Sometimebetween1833and 1844,the Russian tigaphy, and their contribution is widely ac- paleontologistChristian Heinrich Panderdisknowtedged.Furthermore, enough has been covered tiny, lustrous, toothlike fossils in and washedresiduesof Lower Ordovician and Silearnedabouttheir anatomy,associations, patterns of developmentto make them the lurian clastic rocks from Estonia,and on the honestsubjectsofa wide variety oftruly paleo- surfaceof at least one slab of Carboniferous biologicstudies;Thirty yearsagoonly a hand- rock collectedwithin the presentcity limits of ful of paleontologistsclaimed more than a Moscow. However, these microscopicfossils passingacquaintancewith conodonts.Today, werenot illustratedor discussedin print until more than 200 personsdevotea major part of 1856,whenthey weredescribedas the remains their waking hours to the study of thesetiny of an otherwiseunknown group of Paleozoic fossils,and only a few of the world's major oil fishesthat Pander named Conodonten(concompaniesor geologicsurveysare without at odontsin English). In the yearsbetween1833and 1856,Pander least one conodontspecialist.The reasonsfor this paleontologicsuccessstory are numerous, evidently gave a lot of thought to the tiny fosbut they have to do primarily with the fact that sils he named conodonts.During part of that the conodontshave proved to be as successful time, he hadgeat troublewith his eyesandwas at solvingbiostratigraphicproblemsin the Pa- not ableto usehis microscope.However,what leozoic and Triassic as the foraminifers have he sawwhenhe wasableto studyhis specimens been in the interval from the Jurassicto the microscopicallyconvinced him that he was present. looking at the teeth and jaws of a previously In this monographI will introducethe con- unknowngroupoffishes,a groupwith no rnododonts;show how at least someof them may em analogue.Thus, in descriptionsof specibe reconstructedfrom the jumbled bags of mensin his collectionhe usedterminologyapbones by which they are commonly repre- propriate to the teeth and jaws of fishes, and sentedin the fossil record;and look at major some of that terminology survives today. Pander's1856monographnot only provided features of their long geologichistory, with an somelhingusefulabout their the first descriptions of the hard parts of a preeye to suggesting relationsto other animalsand about their pat- viously unknowngroupof animalsand a name tems of deploymentand evolutionary devel- for the group itself, but it also began debates

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INTRODUCTION J sr eral different rL< out to have rSme today agrees h :ased his conqrs.nrs bits and tra Jilferent spe! r!g:L bur for the h:.-iogists E. O. trnaal manv new presented a h.= r- .::d suggested bn-.:g:r be of cond[i< rtre srratigrrs s.rch as the he-.:rcga Shale. i rhat conl' --:.-i of priml::s ,d[ :l lhe same ,-- =E: rhe prevat :i :lese teeth I cr :ires in the bv the --::d rmly on b-E:mt:--r of zoolk ltaving Ul--a :-rib$antial f Bassler wrote ta= ir is clear :afer |!c -\meriql --:e srud] of lE:Jnrial were at l*itr:ologrsts g- 6on.on b= ll:,;. -: Studies, lllcl-:\hed b-y the i : : ll a n d 1 9 34, tirr s:.d;- of conr making hit: rcpresen-+: qosn than G l* a rc:e -oduced and deal group of r:r:: J C:.,:itnt Stud: lftir: --. . nearly G classed k- -re *e con-i

In 1941, Samuel Ellison, a student of the cur in that classification."They regardedthe evidencefor this statement,and for their fur- Missouri school,and Roy Graves,one of his ther conyiction that conodontsare polyphy- students at the Missouri School of Mines, letic, as conclusive,but they put off describing jointly reportedon a group of Pennsylvanian that evidenceuntil a later time. Unfortunately, conodont elementsthey had collected from that time never came. However, they did al- samplesof the Dimple Limestone in Texas, lude to the fact that their evid€nceof the fish which they had dissolvedin dilute aceticacid. natureofconodontsis the fact that somespec- Ellisonand Gravesdo not describethe "acetic imens are attachedto a substance". . . that ap- acid method" they used, which suggeststhat pearsbony but doesnot have the structureof they did not regardit as novel. However,they were evidently the frrst to usea laboratory proordinarybone." Bransonand Mehl also joined Pander,Ul- cedurethat, a decadelater,had becomeroutine rich and Bassler,and all previous studentsof in researchinstitutionsaround the world. The conodontsexceptHinde in usingthe shapeof significance ofthis is that prior to 1941,and for individual "teeth" as the guide to their classi about a decadeafter, most collectionsof conthey odont elements had been assembledfrom fication. That is, like their predecessors, rocks, adoptedform taxonomy. However, like Ulrich shalesand other readily disaggregated and Bassler,they thought it likety that speci and carbonaterocks were generallyignored. mens of different shapefunctioned in diferent However,as Ellison and Gravesdemonstrated ways-some as teeth and jaws, othersas body in 1941,asBransonand Mehl reportedin 1944, scales,and additional ones as denticles on and as nearly everyoneknows today, much larger collectionsof well-preservedspecimens sprnes. Between1933and 1950,the Missouri school can be isolatedreadilyfrom carbonatesby disflourishedunder the leadershipofBransonand solvingthem slowly in l0 to 15 percentacetic Mehl, and the predictionof Ulrich and Bassler or formic acid.This, in turn, meansthat, unlike that conodontsmight one day be of greatuse many other types of microfossils,conodonts stratigraphicallyencouragedother American may be collectedeasilyand relativelyinexpenstudentsto collect and describethese previ sively from both carbonateand siliciclastic the rocks. Thus their distribution in stratigaphic ously enigmaticfossils.As a consequence, literatureon conodonts-a mere200articlesin sectionsof mixed lithologic type may be deterthe late 1920s-more than tripled between mined wilh considerableprecision. 1930and 1950.It is not easyto singleout any The increasedsize of collectionsmade posof theseconributions as more important than sible by more widespreaduseof organicacids the others,but it is not diftcult to identify those encouragedmicropaleontologists interestedin that introducedsignificantnew trends. conodonts to broaden their studies through In 1934,for example,Hermann Schmidtin considerationof sectionsin which there were Germany and Harold Scott in the United no shales or mechanicallyreducible clastic Statesindependentlyreportedthe discoveryof rocks. By 1959,conodontshad beencollected clustersof morphologicaltydifferentconodont in some variety from rocks that rangein age elements on the surfacesof Carboniferous from Late Cambrian to late Triassic, and there black shale stabs. Like Hinde before them, was one report of distinctive elementsfrom Schmidt and Scott regarded these natural as- Upper Cretaceousrocks in west Africa. Fursemblagesas the more or lesscompleteappa- thermore,conodontswereknown from marine ratusesof individualconodonts-an opinion rocksin this time interval from six ofthe seyen that wasroundly criticizedby Bransonand ev- continents. The stratigraphicrange of conidently acceptedwith great reservationsby odonts had been considerably broadened other studentsof conodonts.In more recent through improvements in laboratory techyearsnatural assemblages have playedan im- niques,and geographicdistribution had been portant role in the developmentof conodont extendedand collectionsgreatly increasedin taxonomy. size.No longerwereinyestigatorssatisfiedwith

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searchingthat multielement taxonomy was lison have continued with the published biblimore desirablethan form taxonomyfor cono- ographies of the conodont literature that are donts and was also attainableeven in the ab- listed among the referencesat the end of this senceof natural assemblages to supportevery chapter. In more recent years, the bibliography diagnosedspecies.A complete multielement of conodont literature has been updated antaxonomy is still not in place, however,and nually in a supplement to The Pander Society's thereare a number of workerswho still prefer newsletter. At the end of 1987,my file included its much simpler predecessor. However,a re- the titles of more than 7000 books, articles, and vised versionof Volume W of the now-vener- papers on conodonts. able Treatise on Inwrtebrate Paleontology (Clark et al., l98l) issuedin 1981(but written 1.2 Achievements largelybefore1976)is couchedmostlyin terms of multielementtaxonomy,and a majority of The study olconodonts is very different in the current reportson conodontsat leastgive lip 1980sfrom what it was in 1950,when I began. service to this more sophisticatedmode of For example, in the last 30 years interest in classification. conodonts has again become international, For In 1967studentsof conodontsattendingan much of the time between 1926 and 1950, internationalsymposiumon the DevonianSys- study of conodonts was largely an American tem in Calgary,Alberta, were impressedwith endeavor; now there is active and increasingly the fact that data provided by conodontscon- well-informed interest in nearly every part of tributed to a very largenumber of the reports the globe. At a meeting of The Pander Society presentedand with the difficulty they experi- in 1985, the approximately 150 participants encedin keepingabreastofthe burgeoninglit- represented 31 different countries in Africa, eratureand researchinterestsof a rapidly in- Asia, Australia, Europe, and North and South creasinggroup of conodontstudents.To solve America. Study has become international theseproblems,the groupfoundedan informal again, and there is also close coordination and organization,ThePanderSociery,opento any- an unusual degree of cooperation among stuoneinterestedin conodontsand with the single dents of conodonts. These achievements, perpurposeofsharinginformationon currentcon- haps more than any others, have enabled the odont research.Subsequently, The PanderSo- rapid growth ofknowledge about conodonts in grown in membershipto more than recent decades. ciety has 250; has becomethe official working group on As noted earlier, assessmentof large, straticonodontsof the International Palaeontologi- graphically comprehensive collections on an cal Association:and has distributedan annual intemational scale has also caused a shift in newsletterthat includesreportson the research taxonomic base from the morphology of single activities of members,addressesof conodont skeletal elemenls to the composition and relaworkersand, in recentyears,current bibliog- tionships within recurrent groups of skeletal raphiesofthe conodontliterature.The Pander elements. This has resulted in a taxonomy for Societymeetsannually in North America and conodonts that, while admittedly more comat 3- to s-year intervals at various sites in plex than its predecessor,is probably closer to Europe. biologic "truth" than the form taxonomy of postof the In sortingout significantevents Pander, Ulrich and Bassler, and Branson and 1950era in the history of conodontresearch,I Mehl. Such a taxonomy is obviously a collecshould mention that students of conodonts tive effort and, like any other, will always be have been blessedmore or lessregularlywith ripe for modification and the subject for debibliographers,who have kept track of the lit- bate. Nevertheless, students ofconodonts now eratureabout conodontsand haveperiodically have the framework within which to make assembledand published lists of it. Grace biologically meaningful statements about conHolmesbeganthis servicein 1928;RobertFay odont paleoecology,biogeography, and evolucontinuedit through 1948,with a usefulcatalog lion. and that musl be regardedas an imporpublishedin 1952;and Sidneyesh and SamEl- tant achievement.

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!tr,i...:..,.3mbled ts a.;:!:c devel_:\onomy EI[::] Ej:,r j lbr the i l" : . : : -: b r tn a 4|:r,. = :o e\al-: l*ul -: : ::--. mode re. a: ---,.re has f :--oul --:i:::]:te IE - : ,:: i o sln!CL:: ::: most -::s may &u . .:r . ::t-::nl1 a . {[: _--: ;e still fil ._:: :;3stton, r c- :- I : se e ms J +=-: l'.-:-usi\ ely r =-;: sied rhat ,tr:,:::-.in dgell!f ::a:- of lhese ! r: :: lIOpOSed mg ::ce of life m: :ecOrd. I E[ -,]a-:ience Of tc -an rndi- g a s petr : : r-: { :: -: ::re more m.,--. -: e lurther rF--: ::.ltrile and E - - : i:1 e r this

may not apply to all conodonts,and the elongate, prominently "tailed" hagfish (or slime eel),which many hold to be the nearestliving relative of conodonts,is a largely sedentary creaturethat spendsmost ofits life in burrows in the muddy substrateof relativelydeepmarine water. Clearly, we need to addressthe mode-oflife question intensively before we delve further into headysubjectssuchas conodont ecologyor biogeography. On a more mundanelevel, but nevertheless of signalimportance,is the businessoffilling in a number of gapsin the current stratigraphic record of the Conodonta.Lower Ordovician speciesare well known in the high-latitude faunascharacteristic of Europeanlocalities,but in North America there have beenonly a few reports,and theseleavea numcomprehensive ber of important taxonomic and phylogenetic questionsunanswered.The Silurian is also a problem, particularlythe part of it above the Pterospathodus amorphognat hoidesZone.This part of the Silurian was evidently a time of with widespreadshalmajor marineregression, low-waterenvironments,evaporites,and conditions hostile to developmentof rocks from which conodontsmight be extractedeasilyand in abundance.Nevertheless, a number of very important eventsin the evolutionaryhistory of the Conodontatook placeduring the late Silurian, and we badly needwell-documentedcollectionsas the basisfor frndingout what happened. Taxonomy of Carboniferous and Permianconodontsis in lessrobusthealththan is that of earlierforms or that of most Triassic lineages.Only a few studies have addressed multielementtaxonomy of Mississippianconodontswith conviction,and a few excellentrecent studiessuggest that thereare major surprises in store for anyone who undertakes detailed studiesof Permian conodonts.I recommendthesestratigraphicstudiesstrongly. Finally, we have probablyreachedthe stage at which monographicstudiesofthe taxonomy and phylogenyof major lineagesare in order. To be sure,a numberofsuch studieshavebeen completed,and my debt to them is plainly in evidencein Chapter 5. We need still more of them, and existingcollectionswill probablybe adequateasthe basisfor many.

References Aldridge, R. J., Briggs,D. E. G., Clarkson, E. N. K., and Smith, M. P. (1986).The afrnities of conodonts-new evidence from the Carboniferous of Edinburgh, Scotland. Lethqia l9(4), 219-29t. Ash, S. R. (1961).Bibliographyand index ofconodonts, 1949-1958.Micropaleontology7, 213244. Bergstrtim,S. M., and Sweet,W. C. (1966).Conodonts from the Lexington Limestone (Middle Ordovician) of Kentucky and its lateral equivalents in Ohio and Indiana. Bull. Am. Paleont. s0(229),27t-44t. Branson,E. B., and Mehl, M. G. (1933-1934). Conodont Studies.Unlv. Missouri StudiesS.l300. Briggs,D. E. G., Clarkson,E. N. K., and Aldridge, R. J. (1983).The conodontanimal.Lethata16, 1- 14. Clark, D. L., Sweet, W. C., Bergstriim, S. M., Klapper, G., Austin, R. L., Rhodes, F. H. T., Miiller, K. 1., Ziegler, W., Lindstrtim, M., Miller, J. F., and Harris, A. G. (1981).Conodonta. I\ T rcqtise on I nvertebrate Pqleontology (ed. R. A. Robison),Pt. w, Suppl.2, wlW202. Geol. Soc. America and Univ. Kansas, 202 pp. Ellison, S. P., Jr. (1962).Annotated bibliography, and index. of conodonts. Texas Univ. Publ. 6210,128pp. (1963). Supplementto annotated bibliography, and index, of conodonts. Texqs J. Sci. 15,50-67. and Graves,R. W., Jr. (1941). I-ower -, Pennsylvanian(Dimple Limestone)conodonts of the Marathon region, Texas. Univ. Missouri School of Mines and Metallurgy Bull., Tech. ser.r4(3), l -21. Epstein, A. G., Epstein, J. B., and Harris, L. D. (1977). Conodont color alteration-An index to organic metamorphism. U. S. Geol. Surv. Prof. Paper 995, 27 pp. Fay, R. O. (1952).Catalogueof conodonts. Univ. Kansas PqleonL Contr. (Vertebrata) Aft. 3, 206 pp. Hinde, G. J. (1879). On conodontsfrom the Chazy and Cincinnati group ofthe Cambro-Silurian, and from the Hamilton and Geneseeshaledivisions ofthe Devonian in Canadaand the United Statas. QuqrL J. Geol. Soc. London

35,35r-369.

Holm€s, G. B. (1928).A biblioeraphy ofthe conodonts with descriptionsof early Mississippian species.Proc. U. S. Nqt. Mus.72, Art. 5,38 pp. Huckriede, R. (1958).Die Conodonten der Mediterranen Trias und ihr stratigraphischerWert. Palaod. Z. 32, l4l-l'75. Kohut, J. J. (1969). Determination, statistical

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2. SKELETALANATOMY

For reasonslisted towardthe end ofchapter l, it is now clear that conodonts were primarily soft-bodied animals. Their only mineralized parts formed a cephalic apparatus, at least partly of epidermal origin, that probably functioned to grasp prey and aid in the intake of food. In most, if not all, conodonts, components ofthe exoskeletalapparatuswerediscrete objectsthat becamedissociatedfrom others on death of the animal. Although collectionsat various places in the world house several hundred more or lesscompletely preservedapparatuses,discrete specimensare the ones on which most of our information about the Conodontais based. Pander diagnosed lhe Conodontenin ieffis of the physical characters of discrete specimens. However, everywherein his monograph excepton the pagebearing the diagnosis,he referred to the "teeth," "jaws," or "rernains" of conodonts and thus usedthat word both for the animals as a whole and for their hard parts. I am not aware that this dual usageof conodont has ever puzzled anyone, but tlre potential for confusionexists.In the rest of this book I will use the word conodont only for an entire individual ofthe Conodonta.The term "conodont element" (or "skeletalelement," or just plain "element") will be used for discrete components of the cephalic apparatus.

maintain that conodont elements are composed of carbonate of lime. He cited this evi denceto counter conclusionsthat the little fossils might representa group relatedto annelids or "naked mollusks," whosejaws or radular elementsare of purely organiccomposition. In 1926 P. v. Roundy reported that conodont elements". . . appearto be a phosphatic carbonateof lime." Six yearslater Staufferand Plummer asserted,without citing further evi dence, that "conodont teeth are probably chieflycalciumphosphate."Bransonand Mehl seem not to have worried much about the chemical composition of conodont elements but, in 1944, one of their doctoral students, Samuel Ellison, demonstrated from X-ray analysesthat the substanc€inyolved is a member of the apatite isomorphous group. Because ofthis, conodontelementsare relativelyheavy (2.84 to 3.10) and are less soluble in acetic, formic, and citric acids than the matrix of carbonate rocks enclosingthem. This is imporit meansthat conodont tant. ofcourse.because elements may be extracted from carbonate rocks by prolonged soaking in such acids-a capacitythat setsthem asidefrom many other fossils-and they may also be separatedfrom large acid-insoluble residues by use of healry liquids or other types of gravity separation techniques. The most definitive, and also the most recent, contribution to the subject of conodont2.1 Compositionof ConodontElements elementcompositionis a monographby PietzFrom what appear to have been rather primi- ner et al. (1968),whosestudiesled them to the tive wet-chemicalanalyses,Panderconcluded following formula for the mineral matter of that the skeletal elements of conodonts are conodontelements, composed entirely of calcium carbonale(reinCa, Naoto (POa)3.qt(COj)0.r6 Fo', (HrO)'.s5 em kohlensaurem Kalk). L few years later, however, Harley ( I 86I ) wrote that ". . .they are They interpreted the mineral to be francolite, a composedofphosphate and carbonateof lime, carbonate apatite in which the OH and CO, the former, contrary to my expectations, the ions substitute for phosphate and do not ocmore abundant constituent." Hinde (1879), cupy lattice positions as they do in hydroxylike mostothers,felt that mostof Harley'sspec- apatites.Tracesof at least 39 other chemical imens were not conodontsand continued to elementshave beenidentified in various nlaces

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001

THE MAJOR CONODONT GROUPS

end ofthe element.Thus the Pa elementsofP cooperi are transitional in rnorphology between those typical ofthe linguiformis and Ihe robuslicoslatus stocks. In the early Middle Devonian,Polygnathus cooperi apparentlygaverise to forms with subsymmetricalPa elementsthat lack a posterior "tongue" and are distinguishedby a carinathat extendsto the posteriortip. In speciesof the zieglerianusbranch(Fig. 5.45.1, 5.45.2,5.45.3, and 5.45.41),platform segments ofPa elements becomereducedin widlh and the carina iuts

r01

out beyondthem for slight to considerabte distancesposteriorly.Pa elementsofthe morphologically conservative robusticostatus brarrch (Fig. 5.45.33),on the other hand, are symmetrical, retain both platform segments,and differ from those of ances1ralP- cooperi only in lacking any traceofa posterior"tongue." Symmetrical Pa elementsof the trigonicus branch (Fig. 5.45.16,5.45.18,5.45.36,and 5.45.37)are blunt, robust,and havewide adcarinaltroughs that extendfar to the posterior. Pa elementstypical of the Polygnathuscos-

Fig. 5.45. Lower and Middle Devonian lineages it Polygnathus- Speciesrepresented by the el€ments figured are identified in the text. Modified, with additions, from Weddige(1977)and Weddigeand Ziegler(1979).

b lr&i

Pelmatolepis

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21

*midtognathus, Erated in Fig. 5.14), together mmmonly been

Fig.5.4?, Elementstypical ofmajor speciesofthe Palmatolepidae.Modified, with additions and omissions,from Helms and Ziegler, in Clark et al. (1981). (l) "Polygnathuf' lalilossalusWirth, (2) "Polygnathus" limitaris, (3) " Polygnathus" cristat s,(4\ Schmidlognalhus wittekindti,(5) Klapperina disparilis;(6) K. dispatalvea,(7) Mesotatis asymmelncusi(E\ Palmalolepistrunsilans,(9')P. p nclata; (10) P. proversa,(ll) P. hassii(12\ P. nicornis,(13) P. subtecla;(14) P. gigas;(15) P. lmguifurmis; (16) P. t ahgulaisi (17) P. pe obataperlobata;(18) P. uepida; (19\ P. termini; (20, 23) P. pe obata schindewolj; (21) P. perlobata moximai (22) P. perlobata helmsi; (24) P. pe obata postera, (25) P. pe obata sigmoideai (26\ P. rugosa ampla; (27\ P. pe obata grossi; (28,29) P. minuta minuta; (30) P. minulo schleizia;(31) P. grac is gacilis;(32) P. gtucilis gohioclymehiae,(33\ P. grucilis ma ca; (34) P. quadruntinodosa nllexoidea; (35\ P. quadrantinodosa iaflexa; (36, 31\ P- marginifera; (38\ P. ntgosa tachyteru; (39) P. rugosarugosa:(40) P. quadrantinodosalobata;(4l)P. subpe obata:(42)P. ten ipunctata;(43)P. gtabraprima,(44) P. klappen: (45\ P. glabra acutai (46\ P. glabra lepta; (4'7) P. glabra pectinalat and (48\ P. glabru distorta.

103

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t0r

THE MAJORCONODONTGROUPS

Erses of species Fina hzve not ibed nor have nl specieswith E rclationshipof d ro the betterS Palmatolepis tarures of their r duparilis (Fig. JI-6r. and.K. disdose. and carh bladesand a ir of midlength Each the postete under side, hel- is a distinct I$inctly to one rtasal pit seems fupperina from ruhus cristatus I bsal pit tends im morphologSnatolepis(Fig. ronll have no rr5 built by spe. 5--11.4)have a r-like basal pit llh of the platI On the other rrlin iplanatePa pc-rarru" Wirth lrc be the anceshrared near the ioped platform. rdantly reprerl from many lied intensively * for the high'[per Devonian P. | 971)and varE publications. 4- of Palmatolelosn schemati;ed largely from m preparedfor $' Helms and I and Zieg.ler(in r number of dis-

105

tinct branches,most Out not all) of which are rugosa.P. rugosa ampla (sketch 26) is thought shown schematicallyin Fig. 5.47.Initial Fras- to be a derivative of the P. (Palmatolepis) nian radiationis represented in Fig. 5.47by Pa stock,which is whereit is placedin Fig. 5.47. elementsofthe groupofspeciesnumbered8 to However,Sandbergand Ziegler(1973)postu15.This speciesgroup is identifiedby someau- late development of P. rugosa trachttera thors as the subgenusManlicolepis. Note that (sketch 38) and P. rugosa rzgosa (sketch 39) Pa elements of theseearly speciesof Palmato- from a speciesin the Conditolepis stock that is lepishave a prominent lobe on the outer side; relatedto the one u/ith Pa elementslike those a straightor only slightlysinuouscarina;an un- shown in sketches36 and 37. If the phylogedistinguishedcentral node; a downwardlyde- netic arrangementof thesethree subspecies is flectedposteriortip; and no parapetalong the as shownin Fig. 5.47(and we may alwayshave inner margin. to guessabout that), some nomenclaturaladThe severalFamennianlineagesrecognized justmentswill obviouslyhaveto be made.The vtithin Palmatoleprs diverge from P. triangu- more important lesson,however,is that gross laris (Fig.5.47.16),whosePa elementsare re- morphologyofsingle elementsmay not always latedin rnanymorphologicparticularsto those be the best guide to relationship.All of these of the Frasnian"manticolepids"but differ in "subspecies"are said to be connectedthrough havinga moredistinctlysinuouscarina:a more morphologically intermediate Pa elements conspicuouscentralnode;and a platform that with other membersof the lineagesto which is flexed upward (instead of downward) they are assignedinFiE 5.4'1. posteriorly. Species in the Palmatolepis branch repreSketchesI 7 and 20 to 27 in Fig. 5.47 are Pa sentedby sketches28 to 33 in Fig. 5.47 are elementsof a group of species(or subspecies) characterizedby small, narrow Pa elementsin that includes lhe type of Palmatolepis, P. per- which the smooth-surfaced platformwithdraws Iobata.Pa elementsofspeciesin this groupare phylogenetically from the anteriorend and belargeand conspicuouslysinuous;have narrow comeswidest and best developedposteriorof outer laterallobesthat generallybeara distinct the centralnode.Anterior of the centralnode, secondarycarina;and developa low, ridgelike the main axis ofthe elementis developedas a parapeton the inner sideanteriorto the central conspicuousblade.A more important characnode. ter of this stock, which was referred to the The Palmatolepislineagethat begins with subgewsDeflectoleprs by Miiller (1956),may species42 in Fig. 5.47 is distinguishedby Pa be the discovery by van den Boogaardand elementswith long, mostly smooth platforms Kuhry (1979)that Pb elementsare not "noththat lack an outer laterallobebut beara serrate ognathellan"but are archedpastinateforms of or ridgelikeparapeton the inner side,anterior a type that hasbeenreferredin form taxonomy of the centralnode.Speciesin this lineageare to severalspeciesof Tripodellus Sannemann.It inluded by many authors in the subgenus is difficult to seehow thesestructurescan be Panderolepis. homologizedwith the "nothognathellan"Pb van den Boogaardand Kuhry (1979) refer elements of speciesin other Palmatolepis linmost of the Palmatolepisspeciesin the stock eages,so it may tum out that this group will thar beginswith skerch 35 (Fig. 5.47) to the ultimately merit separate generic distinction. subgenusConditolepis.Pa elementsof most When (and i0 that day comes,the venerable speciesin this group lack an outer laterallobe, name TripodellusSannemann,1955 (not Dehaveovateto quadrateplatformswith a carina flectolepis M.ijller, 1956)will be available. that is weakly developed posterior of a large Conventional wisdom has it (e.8., Ziegler central node; and a sharp-edgedparapet that and Lane, 1987)that Polygnathuslatifossatus rims the inner margin to a point well posterior Wirth (Fig. 5.47.1),progenitorofthe Palmatoofthe centralnode. lepidae, developed from speciesof the PolygPa elementsdepicted.insketches26, 38, and nathus varcus stock (the far righthand branch 39 in Fig. 5.4'7have been taken to represent in Fig. 5.45).Speciesin that stockbuilt carmi three chronologicsubspeciesof Palmatolepis niplanate Pa elements with long, subquadrate

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Tgnathuslattfu* b Poh,gnathus, rnerable genus Itassary to reas15.17.2), P. uishr specieswith f are supposed ELJ. As a temribiiatus might rlrus and the irii. which has '$l- esymmetriI have not foly routeshere,or : rsed the rubric trrmal assignwell be de$t of the speDnsructed and -les

bsin and d ro this family td Polygnathus, r the attachment l- -\ moreor less dia-n groove, oc,.hn its position I b1 a broad flat fr may have a I of its lengh, or msiderably less dthe pit. 7 is complicated, . 5.48):the genus That is, Siphonbnognathus by :lhat namecould rue it had been cE.in 1944Branhtirute name.$ldle (1934) had boper (1939)the thus znd. Dinohblv parts of the della. Althou$t g,Falcodus,Elicfunodella to be d s-ork 1o prove

107

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Fig. 5.48. Elements typical of m\ltielement Siphonodella (Elictognathidae).

it. When (and if) that happens,Falcodus will and that they might not yet haye appearedin clearlybe the oldestvalid namefor the typical populationsof S. praesulcatd,the earliestand genus of the family Elictognathtdae, and EIic- least Siphonodellalike speciesof the genus,or tognathw, Dinodus, ard Siphonodella will be even in early populationsof S. sulcata.Those junior synonyms.The family name Elictog- earlyrepresentatives may haveevolvedPa elenathidaewill stand.however.becauseit is the ments with the essentialcharactersof Siphonoldestone proposedfor any ofthe genera(syn- odella,bnt they might have retainedthe genonymsor not) now includedin it. eralizedPb, M, and S-serieselementsof their Although a greatdeal of attention has been ancestors.In short, evolution of Siphonodella paid to the morphologyofelictognathidPa ele- may not have affectedall componentsof the ments,there is no publishedreconstructionof skeletalapparalus equallyor at the sametime, the complete skeletal apparatusfor any of the and there is no reasonto assumea priori that 20 speciesnow includedin the family. No one, non-Pa elements were morphologically the 10my knowledge,hasventuredevena guessas samein the apparatuses ofeveryspecies. to compositionofthe apparatusof any species With the exception of Siphonodellapraesulof Alternognathus(Fig. 5.49.1to 5.49.3),but cata Sandberg,all known Late Devonian repSandberget al. (1978)speculatedthat the Pb resentalivesof the Elictognathidaebelong in position was occupiedin the apparatusof at Alternognathus, which was created by Ziegler least some speciesof Siphonodella by anguli- and Sandberg(1984)for speciespreviouslyreplanateelementsofthe sortcommonlyreferred fened with question to Polygnathusor in1oEliclognathustthat the M position included cluded in Scaphignathus, whosespecieshave elementsthat havebeendescribedin form tax- Pa elementsthat are closely similar in moronomy as Falcodus angulus; and that various phologybut wereprobablyderived from Panform speciesof Dinodus occnpredpositions in dorinellina,not from Mehlina. the S series.As Sandbergand his colleagues The oldestknown speciesof Alternognathus, noted, however, no specimensof Dinodus, A. pseudostrigosus (Dreesenand Dusar) (Fig. Elictognathus, or Falcodus angulus have been 5.49.1), formed carrniniplanatePa elements reported from rocks with elements of ^trplron- with a narrow,asymmetricallydevelopedplatodellapraesulcata(Fig. 5.49.a)or from collec- form that bears only a few nodes marginal to tionswith earlyrepresentatives of ,Srphonodellathe sigmoidally curved carina, and with a narsulcata(Fig. 5.49.5).Theseobservationswere row, flat undersurfacethat has an elongate taken to mean either that S. praesulcata and, basalpit but no very distinctkeel.Youngerspethe earliestrepresentatives of ,S.sulcalawould ciesof Altemognathus(Fig. 5.49.2and 5.49.3) have to be removed from Siphonodellaon are characterized by Pa elements with more groundsof apparatusincompatibility, or that elaboratelydeyelopedplatforms,which havea the occurrenceof Dinodus, Eliclognathus, and. median carina that is separatedby a slight Falcodus angulus with the Pa elements typical depression from the anterior blade and rnarof alI other speciesof Siphonodellais a rcs,rlrof ginal rows ofnodes or short transverseridges. "similarity of niches." I suggestthat there is Although there is currently a substantial also the possibility that the really distinctive stratigraphicgap betweenthe youngestknown features of Siphonodella developed gradually specimensof Ahernognathusand the oldest

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the simplestand presumablymost generalized memberofthat genus,and not from oneofthe probablymore specialized, youngerspecies. As indicatedin Fig. 5.49,Siphonodellapraesulcata was succeededearly in the Carboniferous by ,t sulcata(Fi9.5.49.5),which was evidently the progenitor of highly variable ,S. duplicata(FiE.5.49.6,5.49.9, and5.49.13). Following an interval of substantialyariation,the duplicata zones,featurescharacteristicof Pa elementsin various segmentsof the S. duplicala populationsstabilized,and youngerpopulationswith thosecharactersarerecognizedas independentspecies,most of which rangedto the end of the KinderhookianEpoch.

'Sionodella from ft | -il -1. rcgularis; t tgt S- duphcata, qudruplicata; )S 'rc:cha-

rd the only one i,d€velopednartat are carminItal stages but r6es. which exIsrcrior to the n of S. praesulI carina that is ndose marginal Ss- Zieglerand '. S. praesulcata lEeudoslrigosus,

109

that appearedearly in the Mississippian(Fig. 5.50.5),anterior endsof cup segmentsare directly opposedon oppositesidesof the blade. In Pa elementsof two somewhatyoungerMississippianspecies(Fig. 5.50.6and 5.50.7),howjoin sides ever, anterior endsof cup segments of the blade at slightly different points. This conditionheraldsonethat is commonto the Pa elementsof all speciesof Gnathodus,but it is combined in Pa elementsof Protognathodus praedelicatus(Fig. 5.50.6)and P. cordiformis (Fig. 5.50.7)with featuresof surfaceornamentation that are more like the Pa patterns of older speciesof Protognathodus than those of somewhatyoungerspeciesof Gnathodus. Shortly after Protognathodus praedelicatus (Fig. 5.50.6)appearedin Early Mississippian new 5.9.7 Famib Gnathodontidae, seas,it was joined by geographicallywideConodontsassembledhere (Fig. 5.36) devel- spreadpopulationsof conodontswith similar oped late in the Devonian frorn Bispalhodus but substantially more elaborate Pa elements. rrdbil,1 (Spathognathodontidae).They are Theseconodonts,which representthe first specharacterizedby a basically seximembrateskel- ciesof Gnathodus,differ from P. praedelicatus, etal apparatusin which Pa elementsare car- their presumedancestor,in a numberof ways. miniscaphate,and Scelementsare alate,with a First, Pa elementsare conspicuously asymmetdenticulated posterior process.The sides of ric. The anteriorend ofthe cup on the concave gnathodontidPa elementsflare laterallyposte- (or "inner") side of the element invariably rior to midlength,and their uppersurfacesrnay joins the btadeat a point well anterior of the be smooth or ornamentedby a few scattered point at which the anterior end of the cup on nodes or by longitudinal or radial rows of the conyex (or "outer") side meetsthe blade. nodes. Little attention has been paid to the Furthermore,nodeson the uppersurfaceofthe complete skeletalapparatus,so taxonomy of inner cup segmentcommonly join to form a gnathodontidconodontsis basedalmost en- distinctive ridge- or comblike parapet, vthich tirely on featuresof Pa elements. may be short and weakly definedor long and The oldestgnathodontids,from rocksof lat- prominent. In Pa elementsof most speciesof est Famennian(Devonian)age,are referredto Gnalhodus,the outer cup is more broadly exProtognathodus(Fig. 5.50.2to 5.50.6),whose pandedthan the inner one; it may be essenspeciesformed carmlniscaphatePa elements tially smooth,may bear inegularly distributed with attachmentsurfacesin broadly expanded nodes,or may have longitudinally,radially,or basal cavities (or "cups") that occupied the concentricallyarrangedrows of nodesor low posteriorhalfofthe under surface.Pa elements ridges. In the highly variablepopulationsof Gnalrof the three late Devonian speciesof Prolognathodus difer primarily in the manner in odus that spread with the initial explosive which the upper surfaceofthe posteriorcup is burst, Lane, Sandberg,and Ziegler(1980)recornamented;one species(Fig. 5.50.2)formed ognized three rnajor groups. The group that Pa elementswith smooth-surfaced cups;Pa ele- centerson G. delicatus(Fig. 5.50.8),and also ments of another(Fig. 5.50.4)bearshort rows includesG. cuneiformis(Fig. 5.50.1l),is charof nodeson either side of a subcentralcarina; acterized by Pa elements with a long, well-deand homologouselementsof a third species fined parapet.The groups4pified by G. lypicus (Fig. 5.50.3)havea node or two on either side (Fig. 5.50.9) and G. punctatus(Fig. 5.50.10) of the carina. In Pa elementsof theseLate De- have Pa elenents with short parapets.The latvonian soecies-and in those of a fourth one ter groups are distinguished by the fact that

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(1) I Zegler(1980). frti.16) P. pruedeli@tPifotm$i (12) G. pzdosemtglaber

ht is also clearly Ets fiom the two nples of G. typ5-50.12. r-d(1980),who are llilogenetic inter. 5-50.recognized andre;. 5-50.14) is formedPa eler of rhe two early , a long, well-deinendy widened

denticlesin the posteriorsegmentofthe carina. on reLaneand his colleagues did not speculate lations between G. bilineatus and earlier species of Gnathodus.However, since simplification in the morphologyofPa elementsappears to have beenthe hallmark of evolutionarydevelopment in the three major groupsof early Mississippianspecies,it seemsunlikely that G. bilineatus, with its highly complex Pa elements, is very closelyrelatedto either the lypicusot punctatus grcups, eventhough posteriorly widened denticles are characteristic of the latter. Thus, if thereis a relationshipbelvteenG. bilineatus and any group of early Mississippian gnathodontids,one might be postulatedwith the delicatus Eroup,which specializedin building Pa elementswith long parapetsand might also have taken up denticlewidening in midMississippiantimes. Belka (1985) deives Gnathodusbilineatus from his species,G. praebilineatus, which is said to be "... a perfect horneomorph of Gnathodus delicatus." Although this would seem to vindicate my "guess" that the G. bilineatus slock had its origrns in the delicatus group,I shouldalsonote that BelkachoosesG. semiglaber(Fig. 5.50.13),a member of the punctatusgroup,asthe likely ancestor,and not a speciesofthe /elicatus grcup.This would require reversalof morphologictrendsin the G. punclatus group rccognizedby Lane, Sandberg, and Zie{er (1980)by requiring that posterior denticlesofthe carinarevertto simplicity in G. praebilineatus(only to becomecomplexagain in G. bilineatusitself),and that the parapetbecome longer,despitea tendencyin the puncta,rJ group for the parapetto begin short and become shorter! Gnathodusgirtyi (Fig. 5.50.15),which appearedat aboutthe sametime as G. bilineatus, might be a somewhatsimplifiedmemberof the same group. However, Belka (1985) has recently describeda new species,G. awtini, which is said to have Pa elementstransitional in morphology betweet lhose of G. texanus,an end memberof thepunctatusErorrp,and G.gir/yi. Derivation of G. girtyi from G. austini, however,would require that a short parapet, with a high anterior node,becomelongerand more uniform in heiglt; and that the posteriorly wideneddenticlesof G. texanus,G. aus-

lll

,infs presumedprogenitor,be abandonedin favor of a sirnpler arrangement.I opt at this stagefor a scenario that requires fewer reversals in trend-that is, for a relationship to tJIe delicatus grotp (and, perhaps, G. bilineatus), which has a long parapetand posteriorlysimple denticles. 5.9.8 Lochrieqand Vogelgnathus referred Two additionalMississippianlineages, tentativelyto the Spathognathodontidae in Fig. 5.36,merit discussionhere,separatefrom the Gnathodontidae.One lineageis formedby the several speciesof Lochriea (also known as Paragnathodw)(Fig. 5.51), the other by the two known speciesof VogelgnathusNotby and Rexroad(Fig. 5.51).Speciesof both Lochriea and Vogelgnathus are represented by bedding-planeassemblages, hence there are few mysteriesabout skeletalanatomy. There are substantialquestions,however,about their relationshipsto other conodonts. Lochriea commutatrs (Bransonand Mehl) hasa seximembrateskelelalapparatusin which the Pa posilionwas occupiedby carminiscaphateelementsthat are closelysimilar to those of Protognathodusmeischneri(Fig. 5.50.2)in that the cup is smoothon its upper surface.Indeed,it would be easyto confusePa elements ofthe two specieswhich, however,differ in relative height of carinaand basalcavity and in the fact that denticlesof the carina tend to widen laterallyand developa distinctive cancellatepatternrn L. commulatus.L. cracoviensisBelka,the oldestknown speciesof Lochriea, is distinguishedby Pa elementswith cups of more elliptical outline and laterallyexpanded, surficiallycancellatedenticleson both anterior blade and posterior carina.Other speciesof Lochriea,like other speciesof Protognathodus, developedPa elementswhose cups are variously ornamentedby nodesand ridges. Lochriea uacovienJri appears stratigraphicallyin the biozonejustabovethe onein which Protognathodus cordifurmis is last recorded andjust belowthe onein which Z. commutatus and Gnathodusbilineatus(Fig. 5.50.14)make their debut.It is easyto postulatea relationship betweenZ. cracoviensisall:dL. commutatus, but difficult to relate either sDecieslo Para-

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THE MAJORCONODONTCROUPS

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rix processand dose laterally qr are smooth s of Vogelgnar of late Missistlrcir phylogeil Norby and tottr name and t that the Pa futhodus pass tages that are *ments of ZoIs might be an tved through tklus stabilis, r other gnatho: been considrceding discus5t an attactrYe a rhat Lochriea Leage separate a and Gnathoriable to create r- I recommend : until the relaI relaled genera E[ considered, ls in the same Fesent the hisEd by so many : made out yery

Fibodonridae).

A Fig. 5.52. Upper views ofPa elements typical ofvarious genera ofthe ldiognathodontidae. (A) Declnognathodus; (B) Idiognathoides; (C) Neognathodusi (D) Idiognathodusi and (D Strcptognathodus.

5.9.9 Family ldiognathodontidae Harris qnd Hollingsworth,1933 Idiognalhodontidconodontsare distinguished by carminiscaphatePa elementswhoseupper surfacestlT,icatly bear three longitudinal rows of nodesor denticles.One of theserows, the carina, is a posteriorcontinuation of thefree blade, which is assignedan anterior position and may account for more than half the total length of the element.Conceptually,at least, the other denticle rows are maryinal to the carina; in fact, the carinacurveslaterallyat varying distancesposteriorto the end of the blade to join one ofthe marginalrows or is replaced across much of the platform by a median groove or trough, so that many idiognathodontid Pa elementsappearto have only two denticle rows. The complete skeletal apparatus is known for only a few of the speciesnow includedin the Idiognathodontidae. In thosespecies, the apparatus is seximembrate, includes

an alateSa elementwith a denticulatedposterior process,and is in other respectsclosely similar in compositionto that of the presumably ancestralGnathodontidae. Declinognathodusnoduliferus(Fig. 5.52.\), the oldest knowr idiognathodontid,is drstinguished by carminiscaphatePa elementsin which the carina curves laterally to join the outer marginal denticle row a short distance posterior of the end of the blade. In Pa elementsofspeciesof Idiognathoides(Fig. 5.528), which appea6 shortly afler Declinognathodus, the posterior end of the blade curves abruplly to one sideandjoins the outer maryinaldenticle rorv wihout forming even a short carina. Furthermore,Pa elem€ntsof some speciesof Idiognathoides exhibit ClassIII symmetry; that is, sinistral and dextral specimensdiffer in morphologic detail, but not in overall pattern. Early speciesof Neognathodw(Fig. 5.52C) built subsymmetrical Pa elements in which a well-developed carina extends nearly to the

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THE MAJOR CONODONT GROUPS

d photographic E tbat represent a elongatecup is ro the poste:en subparallel an of course, JStreptognathord to that genus Rl) larger spechs'erse ridges d lobaleareas r rt€ platform's I h.gesl Pa elets illustratedby l!85). the carina rt of the platEard by transI atrtrolaterally ndose marginal hents are the lnathodus spemll Straka,and lar cryptically, tErs to have r to ldiognathtmsylvanian." t dominated by th those domifr the Pennsylplicated underr€s€ntsa maJor Eronomy, m*-art) Slreptondzs. as stratirdicate, and if z.SlreptognathE)- as ontogeatically in Fig. n and Permian rldiognathodus caicshiftsofjuders into adult iritial ancestor, r€ and more diI earlyPermian f,!' through the Fosis affected drs. If such an ned in the detrtant group of

ll5

are of Early Mississippian (Kinderhookian) ageithe youngeslare ofearliest Triassicage. The rootstockofthe Anchignathodontidae is formed by a successionof Carboniferous, Permian, and earliestTriassic speciesherein assigned collectively (and probably quite loosely) to Hindeodas.Only a few ofthese specieshavebeendiagnosedand describedin fully modern, multielement terms, so future work may well establishthat severaldifferent lineagesare involved.In line with the practicein many other families, each of those lineages would merit recognitionas a genus,and there are namesalreadyavailablefor most of them! The oldestanchignathodontidknown to me, Hindeoduscrassidmtalus(Bransonand Mehl) (Fig. 5.5a),representsa group of Early MissisFig. 5,53. Ontogenetic sequencein Pa elements of sippian (Kinderhookian)speciesin which carIdrognathodus- Smallest specimens have morphologic miniscaphatePa elementshave a cup-shaped fea]]'Jfesof Slrcptoghalhodus. D.'aw,n froni photographs basalcavity beneaththeir posteriorhalf and a in van den Boogaardand Bless(19E5). finlike anterior blade that consistsof three or four denticlesthat decline in length and deconodonts,I suspectit involved severallin- creasein width posteriorly. These elements eages,u/hich may have been affectedto differ- wereoriginallydiagnosedin form taxonomyas ent degreesand at diferent times. Sorting this various speciesof Spathodusor Spathognathoout will requirecarefulbiometricstudiesofcol- dus, and it is not possible at present to deterlections from rock sequencesthat represent mine if they representone or dozensof species long-continuedstability of depositionalcondi- in a multielementsense.In any event, Pa eletions. I doubt that it rvill be worked out in the ments of this type are regularlyassociatedin cyclicdepositsby which the Carboniferousand collections from Kinderhookian strata with Lower Permianare represented in much ofthe bowedangulatepectiniformelementsthat have world, despitethe fact that collectionsfrom relativelyshortsubequalanteriorand posterior and have been identified previously such rocks are commonlyrich in specimens processes that represent the Strepognathodus-Idiognath- asspeciesofSzbDryanrolrffBransonand Mehl, 1934. Such elementsqualifu morphologically odzJplexus. and fraternallyas Pb componentsof speciesin the H. crassidentatusgJrou'p. Maybe, after this 5.9.l0 FamilyAnchignathodontidae Clark, group has been revised and its taxonomy 1972 brought up to date, Subbryantodus will t.urn Ozarkodinideconodontsincludedin this fam- out to be the oldestavailablegenericnamefor ily have a basicallyseximembrateskeletalap- rt. paratusthat includescarminiscaphatePa eleCudotaxis piceslingl Chauff, based on maments, angulate Pb elements with relatively terial from somewhatyounger,middle Mississhortprocesses, and alateSaelementsthat lack sippianrocks,alsoseemsto be a memberofthe any trace of a posterior process.At presentone crassidentatusgroup, and I seeno way to disanchignathodontidspeciesis assignedto letitinguishit genericallyfrom otherspeciesin this ota-,cis,which is peculiar in that its apparatus plexus. appearsto havelackedelementsin the P posiHindeodus scitulus (Hinde) (Fig. 5.54) reptions, and all the othersare included in Ilin- resentsa secondcomponentof the Mississipdeodus, wbich is probably interpreted far too pian species-grouphere referred loosely to broadly. The oldest anchignathodontidsknown Hindeodus. H. scitulus has a distinctive car-

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audus. Pa ele,idcztatusgrotp tnds in cuplike le unit and like rmarion of the . tte most diagz of Hindeodus [s ertensiform te Sb position b" One lateral rbeb€ntsharply t distance from mmonly devel'oflong, needleq./as group ape \tississippian

1r7

(Meramecian)age,and the lineageclearly ex- includesancestorsof Homoiranognathus, the tends through the remainder of the Carbonif- lesswell-known Permian Rabeignathusand,Irerous, the Permian, and into the lowermost anognalhus,and the Early Tiassjc IsarcicellaTriassic.Initially, in 1970,I interpretedPa ele- In the Treatise, Clark and I included all these ments of the youngestspecieskrown, H. typi- genera (except Homoiranognathus) with Hincdlrj (Sweet),as componentsof the unimem- deodusand,Aethotaxts in the Anchignathodonbrate apparatusof Anchignathodustypicalis, 1idae.However, information published since whereasoccupantsof the Pb, M, and S posi- the Treatise manuscript was prepared makes tronswereregardedascomponentsofthe mul- me uncomfortablewith lhat assignment.Thus, timembrate apparatus of Ellisonia teicherti I now treat Diplognathodusand its kin as an Sweet.Later (Sweet,1973)it becameobvious independentgroup with familial status. The that Anchignathodus typicalis and, Ellisonia family Sweetognathidae was established(as leicherti werc names for different parts of the Sweetognathinae)for this group by Ritter apparatusofthe samespecies, which ultimately (1986) who, however, included it within the (Sweet,1976;Sweet,in Ziegler,ed,.,1977)was family Anchignathodontidae. shown to be closely similar in skeletalarchitecIn terms of the morphologyof their pa eleture to other species of Hindeodus as inter- ments, speciesof Diplognathodus(Fig. 5.55) preted in a multielementsense.Before all of are in many respectshomeomorphic replaysof this had beensortedout, however,Cl ark(19'12\ the simpler speciesof Late Devonian-Early had baseda family-groupnameonlnchignath- Mississippian Prolognathodusand.mid- to late odas(superfamilyAnchignathodontacea), and MississippianLochriea. The pa elementsin this must prevail for the family under consid- question(Fig. 5.55)are carminiscaphate struceration despite the fact that Anchignathodus tures with a cup that expandswidely to the has been considereda junior subjectivesyn- sidesand extendsto the posteriortip. Upper onyrnof Hindeodussinceat least 1977! surfaces of the cup are characteristically The skeletal apparatus of Hindeodus cristu- smooth, but rare specimensin a large sample /ff (Youngquistand Miller), the Mississippian may havea nodeor denticleor two. Esoeciallv type-species, is now well known,asarethoseof distinctive of lhe Pa elementsof DiplignathoLate Permian H. julfensis (Sweet) and Late dzs, however,is division of the med.iandentiPermian-EarliestTriassic1L lypicalis (SweeI). cle row into a high anteior free blade and, posFor the most part, howeyer, Pennsylvanian terior of it and abovethe cup, a lower carina and Permianspecieshave beenneglected,and that may consistofa few denticlesbut is more their skeletalapparatuses are largelyunknown. typically fused to a smooth idge ot spatula, In fact,a majority ofauthors.whodealwith late which has a subquadrate lateral profile and Paleozoicand early Triassicconodont faunas drops off steeplyto the posterior end of the recognizeonly one speciesin this long interval, cup.Diplognalhoduswas said to have a multiH. minutus (Ellison) which, despitesubstantial membrateapparatusat the time it was estabdifferencesin morphology of its various skele- lishedby Kozur and Merrill (in Kozur, 1975), tal elements,is evenregardedasthe seniorsyn- but no illustrationswereprovided.It is signifionym of FL typicalis, whosetypesare from the cant to note, howeyer, that a hibbardelliform Lower Triassic.The timited materialavailable (i.e., alate)elementwith a long posteriorproto me suggests, however,that critical analysis cesswaslisted as a component. of larger collectionsthat span greater stratiIn the same year that Kozur and Merrill graphic intervals will result in recognitionof (1975)establishedDiplognathodus,perlmutter many species.But that is a job for the future. rnterpreted an aggregation of elements that form a recurrent group in severalsamplesfrom the Lower Permian of Kansasas the skeletal 5.9.11 Family Sweetognathidqe Ritter, 1986 apparatusof D. expansus.Perlmutter'sreconThe group of Carboniferous and Permian spe- struction,which includesan alate Sa element cies refened to Diplognathodus, Sweetogna- with a long posteriorprocess,follows the plan thus, and.Neostreplognathodusnakes up a dis- mentronedvaguely by Kozur and Merrill in tinctive and probably related stock that also their genericdiagnosis.However,perlmutter's

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l&menrs ig',trrs

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ln diferencesin ItEir skeletalapI Diplognathodus fueb-' related as Hr Pa elements Mus expansus, r(1975),is taken Es of otherspers profitableto !d io 1977-that E closely related t to Hindeodus. lis suggestionis b oldest named tphanus, is sepe of L. commu!trt to the entire d interpreied to nillion years of xt of this inforr. b-vnoting that l9E2) have illuselements from l eastern Canada

I l9

that appear to represent Diplogndthodus and are from a stratigraphicinterval that is well within the rangeof Lochriea and.Vogelgnathus. Close sirnilarity in morphology between Pa elementsof DrplognathodusexpansusPerlmutter (1975) Sweetognathus merrilli Kozur ^nd. that the Sweetognathus (1975) suggests slock developedfrom diplognathodontanancestors in the Early Permian. Speciesof Sweetognathus (Fig. 5.55),like thoseof Diplognathodus,had, a quinqui- to seximembrateskeletalapparatusin Babeignethus which alateelementsin the Sa position havea long, denticulatedposteriorprocessand structures in the Pa position are carminiscaphate elementswith a relativelyshort anterior blade Fig. 5.56. Pa elementstypical of speciesassignedto that continuesposteriorly acrossthe broadly Rabeighat hus and I sarcicella (Sweetognathidae). expandedbasalcup as a carinathat is basically an adenticulateridge, but may be replacedby a singlerow oflow broad nodesor supplemented yet completelyconvincedthat this pattern(or laterallyby a pair of subparallelrows of such that of Pa elernentsof Rabeignathus) merrts nodes. A characteristic feature of the Pa ele- recognitionat the genericlevel. ments formed by speciesof Sweetognathusis Sweetognathuswlritei (Rhodes, 1963) was development of a pustulosepattern on the also the apparentancestor,in the Early Permupper surfaceof posterior nodes.Such a pat- tan, of NeostreptoqnathodusClark (1972) (Figtern was also developedon Pa elementsby 5.55), a stock of stratigraphicallyimportant speciesof Lochriea, which may suggesta rela- sweetognathidspecieswith Pa elementschartionship. Evolutionary development of the aclerizedby subparallelrows of nodes sepaSweetognathuslineagehas recently beentraced rated by a well-marked median groove or chanby Ritter(1986). nel. It is also of interestto note that nodesin Ritter (1986)has demonstratedthat species the posterior rows of Pa elementsof Neostrepol RabeignathusKozur (1978)(Fig. 5.56)have tognathodusspeciesseemnot to have the disa quinqui- to seximembrateskeletalapparatus, tinctively pustulosesurfacethat is characterissimilar to that of Sweetognathus.The Iwo tic of comparablenodes in the Pa elements known speciesofRabeignathusare recognized, formed by speciesof Sweetognathus,Homoirhowever, by their distinctive Pa elements, anognathus,Rabeignathus, and Lochriea- T||re which are like the double-rowed carminisca- significance of these features is diftcult to asphate elementsof someSweetognathzsspecies sesswith information presentlyavaitable.It is but differ in having one or two additional sub- possible,of course,that Neostreptognathodus parallel rows of somewhat rnore irregularly representsa stockdevelopedde novofron Dipustulosenodes latenlly. Rabeignathus,which plognathodus, whose stratigaphically disconalmost certainly developed frcm Sweetogna- tinuousrecordextendsalmostto the end ofthe thus, has a r/ery short range in the Early Permian. Permian. The Late Permian Pa elementsfor which Homoiranognathus wasestablishedby Ritter Kozur, Mostler, and Rahimi-Yazd (1976)es(1986) for a singleEarly Permian species(1L tablished Iranognathus arc, in many respects, huecoensis)known only from carminiscaphate homeomorphic replays of those on which Pa elementson whoseupper surfacean aden- Ritter (1986) basedHomoiranognathus.Evoticulate,pustulosemedian ridge is flankedlat- lutionary patternsin Permian stocksare curerally by two to four subparallel rows of pus- rentty diftcult to reconstructbecausecollectulose nodes. Elementsof H. huecoensisare tions from criticat intervals and facieshave yet surelydistinctive in appearance, but I am not to be madeor describedin detail. However,it

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THE MAJORCONODONTCROUPS

liss III symmetry; E of a pair may be r e\ en thoughthe i end ofthe same H membersof a rlle of denticula4ftarures. ; Cloghergnathus, frognathus (all ild Pa elementsin lerior end of the ldiare between its . Patrognathus apI a-ndrangesinto iia-Upper Crenr of this group ap, some two zones ld are either conch- into the next lis disjunct strati lbe very difficult to f tbe four genera. ghergnathw differ o\' in the profile lce blade,whereas ra'nticlesin Pa elel ma)' be replaced Ess of nodesthat I featuresare cerI the Pa elements b samegenus,but l:nce on the geb its debut in the tha, Patrognathus rtbe Lower Missis:nulata Zone, inr lhat built Pa eler end of the blade fix end of the row h of the doublehis is also the case Eears in the late mft4s and continMississippian,and *Yed in the latest mugh the Pennsylf the Permian. oany singlefeature mbination of feaIte Pa elernentsof

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ClogheEnathus

Adetogntthus

Clydagnathus Fig. 5,57. Elementsand apparatusestypical ofspeciesofthe Cal.usgnathidae.

Clydagnathus, Cavusgtathus, and AdaognalruJ. Apparatusesof Cavusgnathus speciesapparently exhibit Class IIIa symmetry in that right and left Pa elements are distinguished solely by being bowed in different directions. Right and left Pa elemelrtsof Adetognathus,on the other hand,are morphologicallysomewhat differentand thus exhibit ClassIIIb symmetry. In addition, Pa elementsformed by speciesof Adetognathushal/e essentiallyno fixed blade, whereasin comparablePa elementsof Cavusgnathus srycies as rnuch as half the length of the blade may be incorporatedinto the marginal platform row with which the blade is confluent. I would ascribemy consistentinability to separatespecimensrepresentingspeciesof CavusgnathusandAdetognathusto rny lack of ex-

periencewith thesegenerawere it not for the fact that specialistson Carboniferousconodonts appear to have problems lhat arc aI leastasgreatasmine! In any event,elementsof cavusgnathidconodontsare common in rocks that recordmarginalmarineenvironmentsthat werecharacterized by shallowwaterofvariable salinity, and they are useful as guides to such environments. Ancestorsof caYusgnathidconodontshave beenidentifiedby Sandbergand Ziegler(1979) in Late Devonian populationsof spathognathodontid ozarkodinidesreferredto Pandorinellina insita (Stauffer).In thosepopulations,Pa elements (Fig. 5.38) are dominantly singlerowed,carminatepectiniformstructureswith a prominentfinlike bladethat is deflectedto the rightat its posteriorend. However.minor seg-

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VJNOCONOJ

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THE MAJORCONODONTGROUPS

[t986). That study [en1. and distriYdtognathus speb collections were Drxsfucting comE onl) the Pa eleh qere mineralimed them. drzs species are *rucures with a merior V-shaped dtransversely ned by adcarinal , srbmedial carina fu of the platform. iro fiee and fixed Fal length,is conts margin of the flte oppositemarirdvely denticuBitter. Sandberg, Rmjer taxonomic irui Pa elements in keel,which enrior of midlength '6e pit by narrow rgln (termed an lr. Sandbergand ficb may include s!-e. is developed b carina and parl of f{estognathus roryhology of the ladorm maryin; a Ent ofthe anterior of the secondary lt ofbroader and sal margrn mard posteriorto the I rEpresent early Aognathusspecies b Pa elementsof E scaphateunder =sive basalmarhct that the latter lronouncedin its y. suggesBto von I (1986)thatMes&gnathus (family

t23

Cavusgnathidae), whosescaphatePa elements havea double-rowedplatforrnand a bladethat joins the right maryinalrow. Pa elementsof Mestognathusare homeomorphic with those of Scaphignathus(Fig. 5.41),which is thought to have evolvedin the Late Devonianfrom Pandorinellinainsita (Fig. 5.38;- as did Clydagnathus,the proximale ar\cestorof Mestognallrus.Short of a thorough restudy and revision of Pandorinellina, the Cavusgnathidae (including Clydagnathus), and, Fig. 5,59, Elements typical of sp€ciesincluded in the Meslognathus,however, I seeno way ofaccom- Coleodontidae. modating these remarkable homeornorphs (and their common ancestors)in the samesupragenenccategory.Thus, somewhatanomalously, I suppose,| rctain Scaphignathusit the is difficult, if not impossible,to diagnosepropSpathognathodontidae, whereas its younger erly becausethe type-species of Coleodus,its homeomorph, Mestognathus,is accommo- typegenus.is basedon fragmentary specimens datedin a separate family, the Mestognathidae. that make comprehensiyemorphologicinterPossiblythe bestway to insuretaxonomicsym- pretationimpossible.In general,howeyer,spemetry would be to establish a separatefamily cies in this taxonomically isolated group also for Scaphignathus,which would raise the formedrelativelylargeelementsthat havehyapossibilility,of course,of a separatefamily for line, fibrouscrowns,which (in the caseof,S/ereachconodontgenus! eoconusandMixoconuJ)consistofrobust conMestognathusis interpretedas a denizenof iform elements or chains of such elements harsh, nearshoremarine environmentschar- weakly (in the caseof Archeognathus)or stoutly acterizedby high saliniry and probably land- connected(in Coleodusand.Neocoleodus)with ward of the shallow,inshoreenvironmentin- their neighbors at the base. Undersidesof habited by its progenitor,Clydagnathus-von crowns are flat, broadly convex,or longitudiBitter, Sandberg,and Orchard(1986)speculate nally grooved,and in numerousspecimensof that restriction to such an extreme envron- Archeognathusand,Coleodzscrowns surmount ment may help explain the rarity of Mesto- a prominent basalstructurethat consistsofan gnathus and.the fact that it may have mineral- elongatebar that exhibitsa conspicuousdownized only the Pa element$ of its skeletal ward projectionand apparentlylacksa cavity apparatus. or excavation.No basal structureshave been observedwith elementsof .S/ereoconus, Mixoconusor Neocoleodus. Barskov,Moskalenko,and Starostina(1982) 5,10 OrderUnknown illustrate numerous bonelike featuresin the Bransonand 5.10.1 Family Coleodontidae basalstructureof Siberianspecimensreferred Mehl, 1944 to Coleodus,bntK)apper and Bergstrdm(1984) This family is retained for ColeodusBranson were unable to identifi, comparablefeaturesin and Mehl, 1933,Archeognathus Cullison,1938, the basal structure of Archeognathus.Altho]j;gh andNeocoleodus Bransonand Mehl. 1933.and the crownsof elementsassignable to speciesof possibly related forms such as Stereoconus the Coleodontidaeclearly exhibit the intemal Branson and Mehl, 1933, a.nd Mixoconus structure of conodonts, the basal structures Sweet, 1955.Elementsof speciesassignedto have no counterparts elsewherein the Conothesegeneraare shownin Fig. 5.59. donla. No relationshipto olher groups is In connection with their thorough recent apparent. The Coleodontidae,if a natural study of Archeognathus,Klapper and Berg- unit, might representa separateclass of the stritm (1984)point out that the Coleodontidae Conodonta.

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THE MAJORCONODONTGROUPS furdse Acad. We54, rt. B- (1979).Sta: Palmatolepis apdoDlophorids) at d specific level. 1r5 from the Fort Horician), Mararc- {3(5), ll374h1.- and paleonFatr of Missouri. r IJ{3). r-208. - C- | 1947).Lower lEk\'. J. Paleont. ; G. ( 1933).Condodts from the rado: Bainbridge Do City (Lower jr-llissouri StudG no. 3: ConShaleof Missouri. l? l -159. rlJs lcriodus ard I J. Pateont. 12, 233-246 i\ Index E \I . Shimer and 'aL ft. R. (1983).Paea frmily Icriodonlifiw. Fossilsand *t'oniar. Icriodus nl. et Palaeo . 6, fuatigraphie und es der dstlichen ial ){. Jb. Geol. rr. D. G. (1977). conodonts from 51,'772-796'ar. [tte multielement trs. Doliognathus .- 59\2), 299-309. ria.n crisis and its coDodont taxont7-158. Clark, D. l 2). Suwey of r North America. gudies 9(2), 102teqstrom, S. M., Rhodes,F. H. T., , Lindstriim, M., G- (1981).Cono-

dolta- In Treatise on inyertebrate paleontology Pt. W, Suppl. 2 (ed. R. A. Robison). ceol. Soc. America and Univ. Kansas,202 pp. Cooper, B. J. (1975). Multietement conodonts from the Brassfield Limestone (Silurian) of southern Ohio. "/. Paleont. 49,984-1008. (19'77\.Toward a familial classif,cationof Silurian conodonts.l. Paleont.5f , 105?-1071. Cooper, C. L. (1939). Conodonts from a Bushberg-Hannibal horizon in Oklahoma. "l PaIeont. 13, 379-422. Croft, J. S. (1978).Upper Permian conodontsand other microfossilsfrom the Pinery and Lamar Limestone members of the Bel[ Canyon Formation and from the Rustler Formation, west Texas. Unpubl. M. Sc. Thesis,The Ohio State University, Columbus, 176 pp. Drygant, D. M. (1974).ProstyeKonodonty Silura i nizoy Devona Volyno-Podolya (Simple conodonts ofthe Silurian and lowermost Devonian of the Volyn-Podolian arca). Paleont. Sb. Lvov Univ. GlO, 64-70. Dzrk, J- (1976\. Remarks on the evolution of Ordovician conodonts. Acta Palaeont. Polonica

2r,395-455.

(1983).RelationshipsbetweenOrdoyician Baltic and North American Midcontinent conodont faunas.f'offils and Strata 15, 59-85. Dzik, J., and Drygant, D. M. (1986). The apparatus ofpanderodontid conodonts.Ielrara 19, -

r33-141.

Ethington, R. L., and Brand, U. (1981).Oneotodus simptex (Fumish) and the genus Oneotodus (Conodonta)."/. Pqleont. 55, 239-24'1. Ethington, R. L., and Clark, D. L. (1982). Lower and Middle Ordovician conodonts from the Ibex area, western Millard County, Utah. Brigha.m Young Univ. Geol. Studies 28(2\, t160. Fahraeus,L. E. (1984).A critical look at the Treatise family-group classificationof Conodonta: An exercisein eclecticism. Lethaia 17. 293305. Fahraeus,L. E., and Nowlan, G. S. (1978). Franconian (Late Cambrian) to early Champlainian (Middle Ordovician) conodonts ftom the Cow Head Group, westen Newfoundland.. J. Paleont.72, 444-47L Fortey, R. A., I-anding, E., and Skevington, D. (1982). Cambrian-Ordovician boundary sections in the Cow Head Group, western Newfoundland. Pp. 95-129 in The CambrianOrdovician boundary: Sections, fossil distributions,and correlatioflJ(ed. M. G. Bassettand W. T. Dean). Nat. Mus. Wales,Geol. Ser. 3, 27 pp. Furnish, W. M. (1938).Conodontsftom the Prairie du Chien (I-ower Ordovician) beds of the Upper Mississippi yalley. J. Paleont. 12,318340.

Gagiev, M. H. (1979). [Conodonts from the Devonian/Carboniferousboundary depositsof the Omolon Massifl. Guidebook, Tour 9, Biostratigraphy and fauna of Devonian-Carboniferous boundary deposits. l4th Pacifrc ScienceCongress,Khabarovsk,USSR, August 1979,Suppl. 2, 104 pp. (In Russian,\ryithEnglish diagnoses ofnew generaand species.) Harris, R. W. (1964). Subgeneraof the conodont ge\!s Multioistodus in Simpson-Burgen (Ordovician Arbuckle) conodontsf.om Oklahoma. Okla. Geol.Notes 24. 108-lI8Harris, R. W., and Harris, B. (1965). Some Wesr Spring Creek (Ordovician Arbuckle) conodonts from Oklahoma. Okla. Geol. Notes 25, 34-4'1Harris, R. W., and Hollingsworth, R. V. (1933). New Pennsylvanian conodonts from Oklahoma. Am. "L Scl., ser. 5,25(t47), t93-2O4. Hass,W. H. (1959).Conodontsfrom the Chappel Limestone of Texas. U. S. Geol. Sury. Proll Paper 2941,365-40Q. Helrns, J. (1961).Die " nodocostata-Gruppe"der Gattung Polygnathus.Geologiet0, 6'14-'71t. Huckriede, R. (1958).Die Conodonten der Mediterranen Trias und ihr stratigraphischerWert. PalAofi. 2.32. L4l-175. Huddle, J. W. (1934). Conodonts from the New Albany shale of Indiana. Bull. Am. Paleont. 2r(72), 136 pp. Jeppsson,L. (1974) lL9'151.Aspectsof Lare Silurian conodonts,Frssils and Strata 6,19 pp. (1983). Silurian conodont faunas from Gotland,.Fossilsand Stratq 15, l2I-144. Kennedy,D. J. (1980).A restudyofconodonts described by BRANSON & MEHL, 1933, from the JemersonCily Formation, t-ower Ordovician, Missouri. Geol. et Palaeont.14,45-76. Klapper, G., and Barrick, J. E. (1983).Middle Devonian (Eifelian) conodontsfrom the Spillville Formation in northern Iowa and southernMi[rrcsota.J. Pq.leont.57(6), 1212-1243. Klapper, G., and Bergstrom,S. M. (1984). The enigmatic Middle Ordovician fossil Archaeognathus arrdits relationsto conodontsand vertebrates.I Paleont. 58,949-9'16. Klapper, G., and Johnson,D. B. (1975).Sequence in conodont ge rs Polygnathusin Lower Devonian at Lone Mountain. Nevada. Geol.et Palaeont.9,65-83. Klapper, G., and I-ane, H. R. (1985). Upper Devonian (Frasnian) conodonts of the Polygna/ltis biofacies, N.W.T., Canada. J. Paleont. s9(4),904-951. Klapper, G., and Murphy, M. A. (1975).SilurianI-ower Devonian conodont sequencein the Roberts Mountains Formation of central Nevad,a.Univ. Calif. Publ. ceol. Sci. ttl, t-62. Klapper, G., and Philip, c. M. (197D.Devonian conodont apparatusesand their vicarious skeletal elements.Ietra ia 4,429-452.

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THE MAJORCONODONTGROUPS r and Llanvirnian rcrthern Sweden. L r C, R. (1982).Rer fuovician-Early d Group, Clarksrt Sci. 19, 14'74I

:S- ald Bames,C. j a new multielele latest Ordovit- Geol. Surv. Cant!s!). phylogeny, r ot the conodont -{rbuckle Moun-ff- l410-1433. aaafumarion (Lower -donbgat. EltL Geol. et Pa. U- ( 1974).Pennlnts- IIa: The dinidu' Geol. et fua of the Notch trrician), House

ar.i-139.

lrrsions of some 'frovician conoievolution. U/riy. 4.1pp. lb der Conodonr Der ons. l. Die Stnckenb. Naturho Einteilung der

z x. 109-n7.

lf, ( 1957). Early bce) conodonts 3r- 1069-I 108. I ( | 9E2l,ower Dec-kindlei Zones), t to Pvblications, ED€nt genus,{pdria-o of The Canh {lcheringa 4, Eposition of the &anson & Mehl of the Canning ,-]IR J. Austrul.

B_(1e68) F9691.

rkontology of the Creek Member of brian) in southLKentucky. 1/dirts of Nora Fm.,

Toko Range,Glenormiston, pp. l2-15. In Hill, phylogeny of post-Early Permian crisis br'sseftD., Playford, G. and Woods, I.'1., eds.,Ordowhitei Zone corodonts with comments on Late vician and Silurian Fossils of Queensland. Paleozoicdiversity. Geol. et Palaeont.20, 139QueenslandPalaeont.Soc.(Brisbane). 165. Norby, R. D., and Rexroad,C. B. (1985). Vogelg- Sandberg,C. A., and Dreesen,R. (1984).I-ate Denalhus, a r'ew Mississippian conodont genus. vonian icriodontid biofaciesmodels and alterIndiana Geol. Sur,. OccasionalPaper 50,l-14. nate shallow-water conodont zonation. Geol Norris, A. W., Uyeno, T. T., and Mccabe, H. R. Soc.Am. Spec.Paper 196,143-178(1982). Devonian rocks of the Lake WinniDe- Sandberg,C. A., and Gutschick,R. C. (1984).Disgosis-LakeManiroba oulcrop beh, Maniroba. tribution, microfauna, and source-rock potenGeol. Surv.CanadaMem.392, 280 pp. (Also istial of MississippianDelle PhosphaticMember sued as Manitoba Mineral Resources Div., of Woodman Formation and equivalents,Utah Dept. Energy and Mines, Publ.77l.l and adjacentstates.Pp. 135-178.In HydrocarOrchard.M. J. (1980).Upper Ordovicianconobon source rocks ofthe greater Rocky Mountain donts from England and Wales. Geol. et Paregion(ed,.J. Woodward, F. F. Meissner,andJ. Iaeont. 14.9-44. L. Clalton), Rocky Mountain Assoc. GeoloPander, C. H. (1856). Monographie der fossilen gists,Denver. Fischedes silurischenSystemsder russisch-bal- Sandberg,C. A., and Ziegler, W. (1973). RefrnetischenGouvernemetts.Akad. Wiss. St.Petersment of standard Upper Devonian conodont burg,9L pp. zonation based on sections in Nevada and West Paull, R. K. (1983). Dennirion and strarigraphic Germany. Geol. et Palaeont.7,9'7-122significanceof the Lower Triassic (Smithian) (1979).Taxonomy and biofaciesofimporconodonr Gladigondolella meeki n. sp. in the tant conodonts of Late Devonian styri4ctrrvesternUnited States.J. Paleont.59(l), 188Zone. Unired Shtes and Germany. Geo!.et Pat9 2 . laeont. 13,173-212. Perlmutter, B. (19?5).Conodontsfrom the upper- Sandberg,C. A., Zle9l,e\ W., IJuteritz, K., and most WabaunseeGroup (Pennsylvanian)and Brill, S. M. (1978). Phylogeny,speciarionand the Admire and Council Grove groups (PermzonaIi,on of Siphonodella (Conodontz, I)pper ian) in Kansas.Geol.et Palaeont.9,95-l15. Devonian and I-ower Carboniferous). Newsl. Puchkov, V. N., Klapper, G., and Mashkova, T. Stratigr. T, 102-120. V. (1981).Natural assemblages of Palmatolepis Sannemann, D. (1955). OberdevonischeConofrom the Upper Devonian of the northern donten (toII"). Senckenbergiana Lethaea 36, Urals. Actq PalaeonL Polonica 26(3-4\, 281123-156. 298. Satterfield,I. R. (1971).Conodontsand stratigraRepetski, J. E. (1982). Conodonts from El Paso phy of the Girardeau Limestone (Ordovician) Group (Lower Ordovician) of westemmost ofsoutheast Missouri and southwestIllinois. "/. Texas arrd southern New Mexico. New Mexico Paleont. 45,265-273. Bur. Mines Min. Res Mem. 40, l2l pp. Savage,N. M., and Bassett,M. G. (1985). CaraRepetski,J. E., and Ethington, R. L. (1983) Rosdoc-Ashgill conodont faunas from Wales and sodus manitouensis(Conodonta), a new Early the Welsh Borderland. Palaeont. 28Gl- 679Ordovician index fossll.J. Paleont. 57(2),2897 t3. 301. Serpagli,E. (1974).Lower Ordovician conodonts Rexroad, C. B. ( l98l). Conodonts from the Vifrom Precordilleran Argentina (Prowince of San enna Limestone Member of the Branchyille han)- Boll. Soc. Paleont. Ital. 6, t7-98. Formation (Chesterian) in southern Indiana. (1983). The conodont apparatus of Indiana Geol. Surv. Occasional Paper 34, l-16. Icriodus woschmidti z,ie9ler Fossilsq,nd Strata Rexroad, C. B., and Nicoll, R. S. (1972). Cono15, 155-161. donts from the Estill Shale(Silurian, Kentucky Sparling,D. R. (1981).Middle Devonian conoand Ohio) and their bearing on multielement dont apparatuseswith seventypes ofelements. taxonomy. GeoL et PqlaeonLSBl, 57-74. J- Paleont.55, 295-306. Rhodes,F. H. T. (1953).Some British Lorver Pa- Stouge,S. S. (1984).Conodontsofthe Middle Orlaeozoic conodont faunas, Philos. Trans. Roy. dovician Table Head Formation, western NewSoc. London Ser. B. 237,261-334 foundland. Fossils and Strata 16, 145 pp. (1963).Conodontsfrom the topmost Ten- Sweet, W. C. (1970). Uppermost Permian and sleepSandstoneofthe easternBig Hom MounI-ower Triassic conodonts ofthe Salt Range and tains, Wyoming. "/. Paleont. 37, 401-408. Trans-Indus ranges,West Pakistan, pp. 207Rieber, H. (1980). Ein Conodonten-clusteraus 275- In Stratigraphic Boundary problems: der Grenzbitumenzone (Mittlere Trias) des Permian and Triassic of West Pakistan (ed,-BMonte San Gioryio (Kt. Tessin/Schweiz).lzr. Kummel and C. Teichert), [Jniv. Kansas Dept. Naturhist. Mus. Wien 83, 265-214. Geol. Spec. Publ. 4. Ritter, S. M. (1986). Taxonomic revision and (1976). Skeletalanatomy of the Late pa-

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6. EVOLUTIONARY PATTERNS

Fnten der Eifelknchbarten Fat Lahaea 5t,

6.1 Introduction

1979)-Evolutionb mnodont genCteol-et Palaeont. rg des hiiheren ! im Gebiet des ) mit Hilfe von hnt. Abh. 127, Devonian conofr.le (?) of Iowa. ! und Phylgenie I und ihre stati2x- I-andesamtes Eh-\' of the EuLlt Symposium Fd w. C. Sweet, z -lmerica Mem. f conodonts. E lhandlung 3, lJohnson, J. G. ivision ofthe yarts?Upper DevoJn€Iicz. Geol. et ll9E7). Cycles in Fonian to midb, Palaeobiology idge). .Ellis Hort (19E4). Palma.r IEfl of standard lion. Geol. Soc. L d -\ustin, R. L. shodus Etoup Devonian and c, Pelaeont. 8,

data basesthat are the enq' of their peerswho studylargerfossils.This is duepartly,I believe, As noted in ChaptersI and 2, conodontsoccur to the caution automaticallyprovided by very abundantlyin most typesof marine sedimen- largepopulationsamplesfrom stratigraphically tary rock; they are widely drstributed geograph- well-controlledsequenc$.In somepartsofthe ically; and, as a group,they have a long strati- geologiccolumn, densespacingof large samgraphic range. Consequently, conodonts are ples provides an almost continuousrecord of exceptionallyuseful as providers of biostrati the developmentof populationsthat can be graphic information to geologistsinterested in seento have had and to have rnaintained very a broad spectrumofproblems, and thoseofus great internal complexity for appreciableinterwho study conodontshave expendedmost of vals of time. Not uncornmonly,it is difficult or our efort in supplying biostratigraphic service irnpossibleto expresswhat we know about to the geologiccommunity. We have had little such populations within a generally accepted time left over in which to addressthe more es- systematic framework that yr'as designed to otericaspectsofwhat might be consideredtruly handle a rigidly typological taxonomy. And paleobiologictopics. this, of course,has resultedin hiding a good On the plus side, however,our serviceori- deal of very useful information in categories entationhasresultedin the assemblyofan ob- namedand treatedas speciesor generain acjective, if widely dispersed,data base,which is cord with principlesestablishedand rigorously several orders of magnitude larger than that maintainedby Ihe InternationalRulesofZooavailablefor any other group of Paleozoicor logical Nomenclature. Triassic "macrofossils" and is also one that But studentsof conodontshave had addimay be viewed within a biostratigraphic frame- tional excusesfor caution.That is, the subjects work currentlycapableof resolvingthe 300-my oftheir studyrepresenta groupof animalsthat history ofthe Conodontainto 152divisions.In evidently does not occur in the living bioaddition, of course,we now have a fairly clear sphere.This means,of course,that thereis litidea that recurrentgroupsof morphologically tle unequivocalanatomic information availadifferent element types are more suitable than ble for conodonts and that biologicalty individual elementsas the basis for species- meaningfulsystematicand ecologicconcepts level taxonomy;and, as I havetried to showin must be generatedquite indirectly Chapter5, speciesmay now be assembled into Now, however, there are in place at least the supraspecific categoriesthat provide consider- broadoutlinesofa defensibletaxonomy,and it ableinsight into the patternsofconodont el/o- is probably time to take up the biologically lution. At present,our organizedcollections (and philosophically)challengingtask of evoprovide little information as to the modesor lutionary interpretation.To this end, I assemprocesses by which evolutionarydevelopment ble in this chapter some observationson the was effected,but pattemsmay ultimately sug- evolutionary patternsI recognizein the 300gestprocessesthat may also be testedagainst my historyofthe conodonts,and commentson the objectiYeevidence. a few examplesthat seemto suggestprocessas Micropaleontologists,as Lipps (1981) has well. Perhapsrny observationsand comments emphasized,are not noted for generating,test- will proye sufficiently outrageousto stimulate ing, or evaluatingfundamentalpaleobiologic more penetratingstudiesofevolutionary mode hypotheses, despitetheir accessto finely tuned and process.I hope so.

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cyclesin conodont diversity and also delineate about 20 second-ordercyclesthat supportthe conceptadvancedby Zieglerand Lane (1987). The logs,ofcourse,extendthat conceptto both earlier and later intervals of time than Ziegler and Lane considered. In the tabulationsdepictedin Figs. 6.1, 6.2, and 6.3 I have includedonly the 1446species

(of 246 genera)that I regard as reasonablydistinct in a multielementcontext,even though completeapparatuses havebeenestablished for only a few of them. Clearly, as additional data becomeavailable, certain peaksin the diversity logs may be emphasized and certain depressions filled in. But I suspectthat the general patternwill remain much the same.

Fig. 6.3. Speciesdiversitylog (uppcr solid line) and log oforigination-extinction ratios (Iower dashedline) for 48 Permian and Triassic zones. Stippled segments of origination-€xtinction logs indicate times when more species evolved than becameextinct; asterisksindicate times ofsignificant extinction separatilgdiversily cyclesidentifred with Roman numemls XIV to XX.

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tst diYersityin h- in the interrt€ Ibexian and E whiterockian i diwersity to a rtre early Mot- corresponds in Dost u.idespread rdepositedin the America (Sloss, rlme Middle Orftersity of conryrdly to a Late bllowed (with a in at the Ordor generaldecline rian hto the earildle Ordovician : correspondsin r during which Tippecanoe SeBlionh America rth America,an iau rhrough the od by formation c KaskaskiaSe|Fig. 6.2 that inalnost symmetly Devonian and rsiqv maximum ! nme. tia Sequencein fNorth America r and lesswideEara which colrgressive-regresmed Absaroka. I outlines of the roka cycle are rr-Mississippian qtich show a l5rlranian and a , decline in the i to interpret the Ite cratonic cy{- record major

eustaticepisodes,and first-ordersegmentsof the species-diversity logs in Figs.6.1, 6.2, and 6.3.The simplest,I believe,is to postulatea direct relationship betweendiversity of conodont speciesand the expansion(and subsequentreduction) of suitable habitats recorded by rocks of the transgressive-regressive sequences.

eage,which foundedthe Paraconodontida and, in the very early Ordovician, launched the Prioniodontida. Early diversification of the Cavidonti wasalso accomplishedearly in Cycle I. The Proconodontidasurelyreachedthe acme of their evolutionary developmentthen, and only Polonodus and,Pygodus are known from rocksdepositedduring later Ordoviciancycles. The Belodellida,which I interpret as descen6.3 Second-OrderCycles dants ofthe Proconodontusstock, appearedat Superimposed on the first-order cycles re- the climax ofdiversity in CycleI but werenot cordedin the logs of Figs.6.1, 6.2, and 6.3 is conspicuouscomponentsof Ordovician conthe record of 20 second-ordercycles,eachwith odont faunasuntil the interval of CycleII. the generalcharactersofthe onesrecognizedby The 40 conodontgeneraand 227speciesthat Ziegler all.d,Lane (1987)in the Devonian and are largelyconfinedin their known rangeto the Early Carboniferous.By and large, each of interval ofcycle I representtwo groups.Oneis thesecyclesbeginswith an interval oflow spe- made up of mostly shortJived experimental ciesdiversityin which the ratio of speciesorig- stockssuch as the Clavohamulidaeand Corinationsto speciesextinctions(the "evolution- dylodontidae, which appeared briefly during ary index") is below1.0,followedby an interval the initial radiation of cavidont and conodont of higher diversity-that is immediately pre- lineagesbut were apparently not ancestral to cededby an innoyative episodesignaledby a anything else; whereasthe other, typified, for spurt in the evolutionary index. Cyclesare ter- exarnple, by Rossodus, Tripodus, and, Prionminaledby moreor lessabruptdropsin species lodzs, includesgenerathat foundedor are indiversity which, for the most part, seemto be termediate parts of evolutionary lineagesthat, the expectableconsequences ofa declinein the in modified form, assumegreaterimportance evolutionary index in immediately preceding in later cycles.Theselalter lineageswere evizones. dently subjectedto more or less continual As is clearfrom the logsin Figs.6.1,6.2, and, pruning during the prolonged regressivephase 6.3, whose horizontal scaleis approximately that characterizedthe waning stagesin North proportionalto the length of time represented America of the Saukcratoniccycle,so thereis by the intervals shou,n, second-order cycles no evidenceof their catastrophicor massexwere of greatly different durations and hence tinction at the end of CycleI. seemunlikely to have beenthe resultsof any Note that by early in the Ordovrcranconregularlycyclic mechanism.As the following odonts werecommon not only in the tropical remarkswill show,thesecyclesare complexin- to subtropical waters of North America, Austemally,and it may be difficult to find a general tralia, Siberia,and probablypartsofChina, but explanationfor them. also in seasat much higher latitudes,such as those that overspreadEurope, western Australia, and western South America. Although 6.3.1 CyclesI andII there were remarkable parallels in the evoluIn the latestCambrian,Ordovician,and Silu- tionary developmentof low- and highlatitude rian history ofthe Conodonta,I recognizefive Ordovician faunas, already evident in the insecond-orderevolutionary cycles. These are terval of Cycle I, there was limited exchange identified by Roman nurnerals I to V in Fig. betweentheir chamcteristicfaunas.Very pos6.1. sibly the existencethrough the Ordovician of Cycle I, which began in the Late Cambrian well-developedconodonl faunas from the with the appearanceof the first conodontsin equatoralmost to the pole contributedto the the shallow low-latitudeseasof North Amer- exceptionallyhigh diversity ofthose faunas. ica, Australia,and China,is largelya recordof Cycle II is related, at least in North America the rapid diversification of the Teridontus lin- whereit is best known, to a short-livedtrans-

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.As indicatedin Fcies to be the hida which dir ard dominates Llcle III is genfnction and is mbined effects hg of the world rr glaciation.Of Esentedby speEs only 7 suregocks were all Qunas in low he of the lin(or colder* ainued into the

y of the Conod-order cycles, Flonged interbrery, followed :Easintroduced btionary-index Fenlock Epoch. drh diversificarilies IcriodelliI rle ozarkodiltidae, PlectoIdae. The patmtrnue in later rhich are charhtionary bursts Foduced iteraIte icriodellid, [' from succesB.

e are sparseand tler Cycle V is b introducedby nionary vigor, rde of innovarDse from the i/es from the Co4ssognathus m-ery in diverdectinedalmost ime.

135

lygnathusestablishedin earlier phasesof the cycle,and diversity droppedto 2l in the midAs noted in Section 6.2, Ziegler and I-ane Givetian low-diversity interval that initiated (1987) have recently recognizedsix second- Cycle VIII. A bit later in the Givetian (late order cycles in the evolutionary development Middle Devonian), iterative developmentof of Devonian and Early Carboniferouscono- "Polygnathus" latifossatus from the spathodonts.Theseare identifiedin Fig. 6.2 as Cycles gnathodontid stock led rapidly to establishVI to XI. ment of Mesotaxis, Schmidtognathus,KlapperCycle VI began with a short interval in the ina alnd,ultimately, Palmatolepis,which I have late Silurian and earliest Devonian during set asidein Chapter 5 as the new family Patwhich the diversity of conodont species matolepidae.Speciesof these genera,which droppedto a low of 6. This critical interval in were apparently adapted to the increasingly the history ofthe Conodontawasterminatedin widespreadopen marine enyironmentsof the mid-Lochkoviantime by an innovative burst, late Middle Devonian, together with new effectedby vigorous speciation within Ozarko- forms developedfrom surviving membersof dina and iterctive deyelopmentfrom the spath- Ihe Polygnathus stock and short-liyed species ognathodontid, icriodellid, and/or distom- of Ancyrodella that also evolved iteratively odontid stocksofa flood ofspeciesassignedto from spathognathodontid ancestors, accounted Amydrotaxis, Ancyrodella, Pedavis, Pelela h m

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dense,oxygen-poorwater of the thermocline, to which Gondolella was best adapted and in which ldioprioniodus also thrived. As Heckel (1977) noted, that water may also haye been rich in settling organic matter, phosphate,and heavy metals,and thesemay also haveplayed a part in defining Gondolella's depth-zone habitat. Adetognathusand.Ellisonia were certairLlyat the otherend ofthe ecologicspectrumin Pennsylvanian seas.The common occurrenceof specimensof Adetognathus,for example, in rocks that representobviously nearshoreor marginal marine environments(like the outside shaleof FiE 7.'l), in dolomitic stratathat intertonguewith evaporitesequences, and also (in greatlydiminished abundance)as componentsof higher-diversityconodontfaunasthat include forms associatedmore commonly with normal marine conditions suggeststhat ldetognathus$tasearyhalineand adaptedto life in well-oxygenated nearshore waters that were susceptibleto wide variationsin salinity.Mer-

163

rill recognized ihe Adetognathus biofacies in 1962and, in subsequent reports(Merill, 1968; von Bitter, 1972),it hasbeensuggested that ratios between specimensof Adetognathus and, Idiognathodus or Streptognathodas, which dominate environmentsmore offshore,might provide a quantitative senseof distancefrom shoreand a measureof relativesalinity. Environments intermediate between the shallow, well-oxygenatednearshore waters with highly variable salinity and the much deeper,colder,more dysaerobicwatersof the thermoclineseemto have supportedthe most diverseconodontbiotasin the Pennsylvanian. Throughoulthe Pennsylvanian, dominantconodonts in these intermediate environments were members of the ldiognathodus-Streptognathodusplexus,and rocks that accumulated in them have been assignedto a ubiquitoi]s I di ognathodus-Strept ognathodus biofacies (Merrill and von Bitter, 1984).But theseintermediate environmentswere also the ones in which the closely related anchignathodontids

Fig. 7.8. Hypothetical c.oss sectionsofPennsylvanian seaat maximum extent in the U.S. Midcontinent. Upper diagram relates sedimentary facies to inferred quasi-estr.rarinecirculation pattem; lower diagram shows inferred living distribution of conodontsas reconstructedfrom tieir occurrencesand frequenciesin various sedimentarv facies.Redrawn and slightly modified from Swade(1985). prevartrng w rnd

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APPENDIX B. STRATIGRAPHICRANGE CHARTS in and

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In charts on the next 14 pagesI indicate the slraligraphicrangesof 562conodontspeciesin a biostratigaphicframeworkthat includes156 conodont-definedbiozonesand, in the Lower and Upper Carboniferous, severalintervals for which there are currently no widely accepted In choosingwhich conodont-biozonal schemes. of the nearly 5000namedconodontspeciesto include in the following charts,I followed no particular guidelines. By and large I have included specieswhose first and/or last occurrencesdefineboundariesin the biozonal framework, and I haye omitted species with exceptionallylong rangesand/or poorly known or morphologically nondescript apparatuses. Representativespecimensof many of the species included are illustrated in Chapter 5.

I and 938 t 1933 f,ehl, Xehl,

Mark Kleffner, a doctoral student at The Ohio State University, generously abstracted information on the rangesof important Silurianspecies from the much more inclusivecomposite slandard section he has assembledby graphic means.In compilingotherchartsin this appendix I have supplementedmy own recordswith information from sourceslisted below. Thus authors of those rcports must be credited for most of the hard work that went into building the charts in which their data are used. Of course,they must alsosharesomeofthe blame for any errors!

References Higgins, A. C., and Austin, R. L., editors (19E5). A Strutigraphkal Index of Conodonts. Ellis Horwood, Chicheste\ 263 W. Klapper, G., and Johnson, J. G. (1980). Endemism and dispenal of Devonian conodonts."[ PaL 54(2), 400-455. Klapper, G., arld Ziede\ W. (1979). Devonian conodont biostratig.raphy. Spec. Papers in PoIqeont. 23, 199-224. Kovacs, S., and Kozur, H. (1980). Stmtigaphische Reichweite der wichtigsten Conodonten (ohne Zahnreihenconodonten)der Mittelund Obertrias. Geol. Paliiont. Milt. Innsbruck

r0(2),4't-'18.

I-ane, H. R., Sandberg,C. A., and Ziegler, W. (1980). Taxonomy and phylogeny of some Lower Carboniferous conodonts and preliminary standard post-Siphonodella zonation. Geol. et Palaeont.14, ll'l-168. I-ane,H. R., and Struka,J. J., (1974).I-ate Mississippian and early Pennsylvanian conodonts, Arkansas and Oklahoma. Geol. Soc.America Spec.Paper 152, 144 ppRitter, S. M. (1986). Taxonomic revision and phylogeny of post-Early Permian crisis bisseliwhitei Zone conodontswith commentson Late Paleozoicdiversity. Geol et Palaeont. 20, 139165. Wardlaw, B. R., and Collinson, J. W. (1986). Paleontology and deposition of the Phosphoria Formation. Contr. Geol. Univ. Wyoming 24(2), to7 -142.

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Index

Absarokasequ€nce,132, Acanthodontidae,54, lE5 Acanthodus,54, l85, FiE. 5.7 Acodus,6l "Acodtu," 60 Acontiodus,53 Adcarinal trough, 100 Adenticulate process, I 6 Adetognathus,120, l2l, 122, 136, 137, 160, 16l, 163, 188,Fig. 5.s7 Aethotaxis,ll5,ll7,137,161, 164,I87, Fig. 5.54 Agnatha, 177 Alate g€niculateconiform el€ment, 16 Alat€ nongeniculateconiform element, 16 Alale ramiform element, 17 Albid (crowns),13, 14,52 Alternognathus,89,96, IO1, loE, 188,Fig. 5.49 pseudoslrigosus, 107, 108,Fig. 5.49 Ambalodusgalerus,99 Ammocoetelawa, 177 Amoryhognathus,63, 64,7l, 134, 156, 165, 186,Fig. 2. 11, 5. 15 ordovicicus, 26 Afi p hioxus. See B ruhchtostoma Amydrctaxis, 89, 93, 95, 99, 135, 187,Figs. 5.39,5.40 Anchiglathodontacea, I I 7 Anchignarhodontidae,90, 92, 1t 5, I17, 188 Ahchignathodus,l17 typicalis, ll7 Ancoradella,89,93, 98, 187 ploeckehsis, 98 Ancyrodella,89, 96, 106, 135, l4l, 187,Figs. 5.39,5.41 Ancyrodelloides,89, 93, 95, 96, 98, 99, 106, 135, l4l, 187,Fiss. 5.39, 5.40 Ancyrognalhus,lO2, 187,FiE, 5.46 Ahcyrclepis,187 Angulatepectiniform element,2l Annelids,l7l, t72 Ansella,49,50,167,185,Fig 5.3 Ansellidae,42, 49, 185 Anterior process,16 Antler flysch trough, 157 Antognathus,69, 10, 186 Apatognathus,81, I16, 186,Fig. 5.30 Aphelognathus,73,91,93, 134, 154, 155, 156, 187,Fig. 5.37 "Aphelognathus,"9l, |86 gigas,9l kimmsv/ickensis, 9l Aplacophora, 172, 173 Appalachignathus,15, 16, 186,Fie. 5.26

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