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THE CONODONTA
G-
lesources,and
Morphology,Taxonomy,Paleoecology,
andEvolutionaryHistory
of a Long-Extinct
AnimalPhylum
3
azI
High-pressure
EE:
fIt;:t
WALTERC.SWEEL
TheOhioStateUniversity
F
by the
-i-:ztlog'
fr, :.1 a olutionb.-;:n-
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
w.c.s.
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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CONTENTS
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60
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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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The Conodonta
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1. INTRODUCTION
-gntrrus
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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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l?rl1
VJ-NOOONOJ AHl-
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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VJNOCONOJ
gHJ
tt
w
7
INTRODUCTION
G
:
:uccessive
:
:::1
a15. In
ItL :
lr-a:-t3J several
E R : -: adi scre te
, h ::-: \\ alliser
ms - rS !'ollec\\'alliser
ril--:
uE -: _.:i: deterk*- -:. :: ii:crete
4-'- -: --:le same
Er: :-. :-, idence
a r r-: :t useful
E:-:
FrE- '-::: -aaulTent
:One of
G:
: :Jiually
E
: '- . o i rhe
. .: them
I-: --:: . -::matic
&ts:
* .:r:s and
l{tl
d : . : : : -,q i nt o
{ I
-:i.tede.
lt ,",r.- ,': had
-:: , l a t an
fu TE1 =, _f s reI!Er-: - _i.- t: stnf lF
:--:: : ': O f m
F'-- t-:
Ber_s--:
Gr- \': :.::-toses
E-f,:::
:-: -:r5
In
fte : : " :
,jiag I :: ia: -: stnct
f i.:,- 1e-;31
:
_ lin plex
-i
E[T_ : : 5 a::plrve
F:L--r- ti-_drlent
!: ,ri-: !-rslical
G5- -- :'::ngthE : - --:-:l lnt erEE_--{ t: single
lTl --:':-::Lr ely.
ka-_': -_--:: olher
GE: r t.: -lsed ef3: : - * ::C evalD:
- :3dict ed
- - --:3lement
f
_:fi
: t!\m) rn
[:" : -rt erna:r .-., : t !
. n \ [ ar-
6c: :- -:: soul-
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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VJNOCONOJ AHJ
INTRODUCTION
f
:: a : -ninlscent
D-- . : r : : -5r of the
W!::: =.; t= 1mr =-.:hn The
c:ia-:--i
roles in
I L: - : : : : f at t he y
mg ;::-iates
for
R ::-:
:-r\\n the
C :er: -:--on rank
Eli.
__- ..:3s ledge
ill
-,r-- =ore inI s: : .-:tber of
I [ - : : ::,_SSSSer/ -
tr =;::::s:
hereI
!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
J. E -- ,.- l :i i li
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OI
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and
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The generic
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Group
Kansas.
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4 1 . l-
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Fer OrMinn.
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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VJNOOONOS AHJ,
ZT
SKILETAL ANATOMY
t ::
E1jJL
!b*r
:as ca\ ity,
--'1=rn-
r a--- :ored for
--:e trrst to
t&
I n --::: sections
rom::
dif-nr frlling
{me =wI
E". :-<ralized
!G:3:S
fu
rc
'Oet$-een
:: realness
:: rtreele-
fErj:=s
rhe fact
E -:e -ess resisI =E:es
studha;e
..f crown
I la::. ls shown
=: be visuft-. ..---; a bit of
r
:a rhis fact,
1fo :'::: about30
rb G:oss( 1957,
I it ::der for laI s:zn to basal
ES aave grown
of lam;ns:on
h--: of laminae
l.'---i e progres;r r:c- most sigEaiifded b-v tishs
a.ud others,
fuears
sew in
{ =:i reeth and
thus that the Conodonta were probably not however,only the first few lamellaeoverlapped
fish, as Panderand many othershave thought basally to form a very tiny basal cavity,
whereaslater lamellae became progressively
them to be.
as the crown
Elementsof the cephalicapparatusof most shorter than their predecessors
conodonts were dissociatedupon death by grew larger.Basalcavitiesof crownswith feadecayof the tissuesthat held them togetherin tures like this are termed basalpils, and the retife or by digestionofthose tissuesin the gut of mainderofthe attachmentsurfaceis described
basalmargin.
a predator.In only a few caseshave complete as a zoneof recessive
In most parts of the crowns of typical conbeenpreserved.After dissociation
apparatuses
most conodont elementswere moved about odont elements,lamellae are 0.2 to 1.2 pm
growth
with other sedimentaryparticleson the sea- thick. However,at placesof accelerated
platelike
floor and that agitation was apparentlysuffi- in elementsthat grewlaterallyto form
cient in most casesto separatecrowns from structures,or toward the extremitiesof those
basalfillings.In any eyent,most collectionsof that built elongatecomblike shapes,lamellae
conodontelementsconsistentirely (or almost may be as much as 5 pm thick. As noted preentirely)ofcrowns,and we havelittle or no in- viously, each lamella of a completeelement
We canseeedges
formation on the internal structureor extemal surroundsall its predecessors.
morphologyof the basalfillings for most spe- of lamellaealong the attachmentsurfacesof
cies.I suspectthat th€ basalfillings of a great crowns-however.and outlinesof someofthem
many species,perhapsthe majority, wereonly are cornmonly visible in transmitted light
slightly mineralized-or not mineralized al all. within the thinner partsof many crowns.Each
They aremostlyunknown asisolatedobjectsin lamellais built on a frameworkof organicmarocksthat contain lots of discretecrowns,and terial, and the tiny apatitecrystallitesthat mi
they do not occur in any of the completeap- neralizethis frameworkare orientedwith their
paratusesthat have been described,which prism surfacesparallel to the direction of
seemto have undergonelittle, if any, physical growm.
In reflectedlight crowns of conodont eledistortion subsequentto deathof the animals.
Whateverthe explanation,nearly all we know mentsare shiny and translucent.They are pale
aboutconodontsis derivedfrom a studyofiso- amberin color if they are from rocksthat have
not beenheatedabove 80"C for any length of
latedcrowns.
Crownsofconodont elements,like basalfill- time; otherwisethey may rangein color from
ings, are internally laminated.In morphologi- reddishyellow to brown to black if they have
cally simpleforms,like the onein Fig.2.1A,in- been held for any appreciabletime at higher
dividual laminaeare conicalin shape,and the temperatures.Elementsfrom some metamorentirecrown hasthe structureofa nestedstack phic rocksaregray,white, or evencrystalclear.
ofpaperdrinking cups.In the conicalspecimen Evidently elements such as this have been
for
of Fig. 2.1A, the basalfilling is still attached. deeplyburiedand kept at hightemperatures
However,ifthe two componentsofthe element such long periods of time that they have lost
were separated,the surface over which they eventhe fixed carbonresponsiblefor the dark
werejoined would be that of a conicalinden- colors of specimensfrom lower-temperature
tation in the abapicalpart of the crown. This rocks.
Thermally unaltered crowns rnay be uniindentationis termeda basalcavity,and it occupiesa part of the crown known as its b4se. formly translucentand ofthe samepaleyellowNote in Fig. 2.lA that basal edgesof succes- ish color throughout,or one may distinguish
sively younger crown lamellae extend some- within them more or lessirregular, fuzzy-edged
what farthertowardthe basalmaryinthan their massesthat are opaqueand of whitish appearpredecessors.
Thus, the surfacealong which ance.Crownsofthe first typearesaidtobe hyacrown and basalfilling wereoriginallyattached /irq those of the latter are tened albid. Many
is completelyenclosedwithin the baseof the crownshave both hyalineand albid areas,and
crown. In the element depictedin Fig. 2.t8, internal distribution of the white matter thal
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VINOOONOS AHI
nl
SKELETAL ANATOMY
ta:: io leaye the
r;:::-:ol ofthe tisEiil short, the
-Gli''-ti masses ln
F--i :nd lhe se6 :;: :-- ontogenete
...::essionsof
's-,:-?nt
tn taxlz;--of rhe life
E-::
ro the fact
qnl-: -_:o*ns of a
hs :::l
-\{iddle
zr :- -orm fract' - ,-r: margins.
Itc '-:;: internal
E:l-::-:s olfibers
rr--,: !-:h distincE|[-=- :: represent
:: :!\rodonts,
4
. Su:=: -ent studfrii: =: hraline
mc-_-i =e butlt of
t :i= ::de up of
I
: r-,-aled par-:
*r:
s-:---acesparrd :;-' re rodlike
p a c k ed
t::it y
l. €-': : Tl l u s, alr_.
erred in
-:::brm COnn;,: -.:
e;:.
:her \!ere
!-t : =-.rP distinE-,i
lt|ucture.
i i-- :s- or neur:= -:,'luded in
I
:s :,:, be much
: tc---: :n a wide
r$- 3:--ause those
rr
:: rhe funcbr -::-:ran basis
E. :-'.i ol atten=i recogniz- --: -ee descripri
IFt | :!.
It!:i--- :s el\'en rn
E' -_:: :tames ap-
t)
TABLE 2.1. Shapecategoriesofconodonl elemenls
Major shapecategory
Primary
divisions
Coniform (simple cones)
Geniculate
Nongeniculale
Ramiform (bars)
Alate
Teniopedate
Digyrate
Bipennate
Dolabrate
Quadriramate
Multiramate
Secondarydivisions
Breviform
Extensiform
Raslrate
Pectinform (blades,plates,
platforms)
Stellate
Pastinale
Carminate
Angulate
Segminate
plied to thosecategories
in Volume W (Supplement 2) ofthe Treatiseon InvertebratePaleontoloSy (clafk et al., l98l).
Pander,the first to describeconodont elements,divided his specimensinto two major
categories,einfacheZahne ax.dzusammenge'
setzteZahne(i. e., "simple or singleteeth" and
"compound or compositeteeth"). That division is still useful,althoughit hasbeencustomary for many yearsto divide the "compoundor
composite"categoryinto two or threedivisions
termed Ddrs,blades,and plates (ot platforms).
In the shapeclassification adopted by Treqlr'Jeauthors,Pander'ssimpleor singleteethare
coniform (cote-shaped)elemenls;barlike compound or compositeteelh are ramifurm (rayshaped)element$ and blade- and plate- (or
platform) like elementsare groupedtogether
as pectiniform (comb-shaped)elements.Elements describedin Table 2.1 as rastratehave
been regardedeither as peculiarly modified
coniform elementsby someauthorsor ascompound elementsby others.They werenot specifically identified in rhe Treatise classification
of shapecategoriesbut are here regardedas a
separatecategoryfor descriptivepurposes.
StelLplanate
Stelllscaphate
Pastiniplanate
Pastiniscaphate
Carminiplanate
Carminiscaphate
Anguliplanate
Anguliscaphate
Segminiplanate
Segminiscaphate
Bisegminiscaphate
Trisegminiscaphate
Coniform elementsare funher divided into
two categories(geniculateand nongeniculdte)
on the basis of the type of curvature.Seven
major types of ramiform elementsare recognized on the basisof the number of processes
and their relationshipto the cusp.Pectiniform
elementsare divided into five major groupson
the basis of the number and arrangementof
primary processes.
If pectiniformelementsdevelopplatformlikelateralextensions,they may
be further describedin terms of the resulting
shapeofthe attachmentsurface(planateor scaphate).Typical representatives
of eachof these
in
Figs.
2.2,
2.3,2.4,2.5,
categories
are shown
which
information
on
and 2.6,
also include
termsusedto describefeaturesofeach ofthese
elementtypes.
2.3.1 Coniformcrowns
Coniform elements,which are also known in
the literature as "simple cones" or just as
"cones,"are basicallyof conicalshape.Pander
divided elementslike this into two parts, a
more or lessexpandedbase,which enclosesa
subconicalbasalcavity (Ihe CavitasPulpaeof
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ol po^Jncsr ro tqEre4sraqlra sr leql sIX?EuoI
e seqpuE 'lrun eql Jo xod?eql ol sJed?l'pqos
sr qcqlA '(azrrds's,Jepued)dsn, e pu? :(Jepupd
'sEsuE) Jo ,{lrsJe run pu? €clJeuv
.^Aolotuoslsd elErqeuo^ul uo esupolf
Jo ilercos lecrSoloeC eql Jo /kolnoc
Iuo{ ur\rErporlqgu uo ruEJS?rC.su^\olcruloJruocJosed l loferu o^rf eqtyo A8olouluuefpue uoqq uaug .1i3 .B;g
al el ncruaOuou
p -itlsro^run
Er sruerS€rc
I lerolel = dl
! = dE :dsnc
'u
'c'z'3!d
wptnqqtH
mq ,.uuoJ
||- peurel
EJB 'sUod
D aprs qcee
buetsod e
SlJletutu^s
am lv'Kflo
FJJUr OSp
I uxroqs el?
! ssuoSelBJ
ue:q 0^?g
a te l nJrua6
iauroc leseqorelue
urO re| 'ul essq
ll^ec lgseq
,euroc lesgqoJelsod
essq lo ur6Js|rl ,addn
alsoS
le.l
eurJ9c
dsnc lo ' drl t o 'r ede
VJNOCONOJ AHI
9I
SKELETALANATOMY
f,
of cusp
Fg.n
of base
!b|
C,Oane
r
hr
have been recognized.Names for theseshape
categories
arelistedin Table2.1.Typicalforms
are showndiagrammaticallyin Fig. 2.3,which
also includes some information on terminology. Alate ramdorm elemenlsare bilaterally
symmetrical,lack an anteriorprocess,but have
a posterior processand a lateral processon
eachside of the cusp.In somedescriptivereports, alate ramiform elements haye been
termed "trichonodelliform" or "hibbardelliform" becausethe genera Trichonodella and,
Hibbardella (and several others) were origi-
l7
nally form-taxonomicconceptsbasedon alate
ramiform-typespecimens.
Tertiopedateramiform elementsalso have a
posteriorprocessand a lateralprocesson each
side of the cusp,but the lateral processes
are
not symmetricallydisposedwith respectto the
cusp and the posterior processis commonly
long and conspicuouslydenticulated.
Digyrate ramiform elements are like alate
elements,but they are individually asymmetrical; only rarely is the posteriorprocesswell
developedand denticulate;and lateral pro-
Fig. 2.3. Ramiform crowns and their orientation. Tetus used to describeth€m include the followns: ac = anticusp;ap: antetior process;bc = basalcavity;bp = basalpit;c = cusp;d = denticle;ilp = inner laieral process;
lp_= lateralprocess;olp = outer lateralproc€ss;pp = posteriorprocess;and zrm = zone ofrecessivebasalmargin.
Diagramsrcdrawn from Trcatise on InvertebratePaleontology,courtesyofth€ GeologicalSocietyofAmerica;nd
University of Kansas.
il
F
FE
ant
fiom
frh:sa*r
:r t ansas.
ts
I
kilEd
per
Fi
F[r
as 4/4te
elements,
co$ae that
might be
F
t
I
t
F rrr-h of the
!- -'tan" one
f, rre base are
p-c;'resses,
P 161 oo, nDIIIrlns
tFRTsses
+
post
q u a d r t r a m a te
on a
may
|d b ttre same
rhe cusp
] rfl
!d $ar ponion
D lorizonEl. A
Ce !::Dncave,or
processl
rd rhe alterior
-elor
and exlEEr.
rd a ra-miform
EAkm
eleDents
dolabrate
olp
ta.l|opodate
ahlo
f
F
oe
I
ItoloruoelEd
qF rqucsep
t 's'z '3!,{
[srJSJadns
htsq selcll
lsnJ eql Jo
E p [E]aA3S
Fod oqt t"
zrrals€J?qc
deap
^la^n
-eq [SoToruoq olqeqoJd oql ezrseqdue ol u?ql
uos?eJJeqlo ou JoJJr .el"lntlue8 s" poqursap
suuoJ eqt JoJ tuau)p uttoltuot autqopp
^{ou
uorsseJdxoeql esn ol et?udordd? eq lq8lt'u fl
'tc?J uI 'sseroJdJouelsod eleFcuuep Euheq
e
uI
sluourele uloJruoc
elElnorueE
uro+ JoJ
^Iuo 'tcedsoJsrql q .pedsp
-Jrp,(eql
l?Jet?lur ped?qs
{crd ,{IuorullIoc eJ? puE ssecoJd touelsod
e
e^?g stuauap unlrruDr aptqDpe
^lrro
'eteuueolq
ural
eq1 ;o ,{1qrqz
Iurlneu
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lI? e qlr^r aseec
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-4uopol,{proc,, Jo ..urJoJrlapoepulq,, peulrel
ueeq e^?q sluaurole urJoJrr[eJ oleuuodrg
'o1?lnorluep"ss$oJd Jouotsod eqt puE ol"In rl
-uep sr ssecoJd Jouelu?
egl
(snpoutsttg
pue
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.
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=
sorceds
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Jo
equJsep
pJsn
Jo
ot
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uors
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sglcquep $lcel sseJoJdJouolu? eql ,,(Iuoruluoc
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'se^rnJ
"
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^luouj
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sI ssecoJdJouolu" eql ,sluer[gJaujJoJrrrr€Jelpu
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-uodrq lsou.I uI .dsnc eql 01 lcodseJqlr,t\ uoqrs
c4eurals^s ? ur lnq 'sluoluela t4artsroJ asrpatJ
-od ur Jouelsod puE Jouelu? eJ? qorqA{.sassec
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eif-r^trp
ulto!
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r{srnEuqsrp
ol
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ur
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t?rll
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suo$snJsrp uI'sasnleJEdd? luo{rrelerllnut
-Je Jo slueuoduoc el?J,tErpeqt loJ .,urloJrql?u
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se
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sldecuoJ crruouox?l-urloJ ,{l[?ul8uo eJa/ snql
-ouSouuoug pue,snqpu8o&tz,snpoqndsol
-JaH 'snpolno ereue8 sq1 ssn"Jag .suoqceJrp
olrsoddo ur e,rrnc fluoruuroc
serlrue4
lpr{l
-xe Telsrprl1l1'\'pedole^op
ow sessac
'sluaLuala
uttolraot apwotlqnru parujel eq tq8ru (pore
-^ocsrp eq re^e,{us pFoqs) sosseJordlnoJ ueqt
,stuauala
eJoru qll.t\
esoql
s€eJeg,tl
unltraD.t
apuotupDnb sE pequosep
er?
(l?Jel?l o,{l pue
^Iet?udorddp
'Joualsod
'Jouelue)
sessaJoJd
.oJ?J
JnoJ qlrn
asoql
eJ"
sessecoJd
fu"ruud
eerql ueql^{eJ
eJou
qll^l
uJoJnu"u
EuerrJele
.,{uouoxq
lueurelorlJnrll pu?
Kurouox?l
rrrJoJ
uea,{qeq
uorlrsu€Jl
Jo J? Jelur
eql ur penedd" luql $Fo,,l\e^rldursep ur
..tutoJ
-uuoporuoudoeu,, puz
..uuoJrluopouou^c,,
peurJal ueeq
osle
e^?q
s
uos
trl JO Sluau
-efg 'sorceds luopouoc snoue^
Jo sesnlpJ?d
-oe lelela{s eql ur ed^l srql Jo slueueTe ueeal
^lpnbeun
VINOCONOJ EHT
8I
SKELETALANATOMY
ide skeletalapr
Fecies. Eleb been rcrmed
bprioniodontil+tpeared in the
]frm E\onomy
rban three
ic
-e fu sirh four
tuo larcral)
Et-;dirumate
F
-.
fce srrh more
fc== be discovF"lri.amiform
tively deep basalcayity and a posterior maryin
characterizedby a prominent flangelike "heel"
at the posterior end and, typically, a seriesof
seyeral denticles between the heel and the tip
of the cusp. A few rastrate elementslack denticlesbetweenheeland cuspapexand are thus
superficiallylike geniculateconiform elements.
l9
A few others(e.g.,the upper pat in Fig. 2.4)
lack a "heel."
2.3.4 PectiniformUowns
In 1879Hinde usedthe term "pectinateteeth"
for comb-shaped
conodontelementsofthe sort
Fi9.2,5, Pastinateand stellatepectiniform elements,their scaphateand planatederivatives,and terms used to
describethem. (SeeFig. 2.3 for explanation of abbreviations,)Diagams redrawn from Treatise on Invertebrate
Paleontology, courtesy of the Geological Society of America and University of Kansas.
F
i
7
F.
g*ra
Belo-
Parebel! cotriform
to the
&ren$
+
ant. -_
post
past|nale
but
post ._
Sendcles on
6e cusp. No
cI
in the
r slsematrc
K)r one was
sas coined.
compressed,
rirh a rela-
stelliplanate
ant <_
-
pastiniscaphate
oost
PoSl -_
-
ant
anl-
-
stelliscaphate
-+
post.
3a0co,ro
lEr.tal
rrJ l9s9q
rur6as
lU^E. lssBq
-eq eceJns lueurqcene eql pus dsno eql qt?eu
-eq lld ps?q ? ol paJnpar aq f?ur ,(lr ?J aqt ro
:dsn3 aql qt?euaq
spu?dxe q3rq^l
^DuE3gruErs
suorsuelxo olrTqEnoJl Jo
',$l^?J psEq aql Jo
-eAooJA ur pasolcuo eq ,(eu ecslns lueluqcel
-1? eql Jo suorsuelxe sessecoJdesoql Jo sued
rJ^lol Jql uo 'slueu:Ja uloJrtlleJ Jo sesseJoJd
uEqt Jeq8rqpuE possoJdruoc,tll?Jelel eJ?sossec
-oJd'sluourele uJoJtuqced ssalurloJleld lsour
uI 'dsno eql luo{ ,terre puelxo l?ql sossecord
JnoJse ,(ueu sEol auo pue lrd Jo ,qr^ec l?s?q ?
Jo xod? eq1e^oq? pelenlrs sr l?ql dsnJ e Eur^Brl
ur Jelruns I[? eJe suorsuedxe pralel e{llrrrJoJ
-leld ro -elqd ]cpl l?qt su,rorc pod?qs-quroJ
'sed^l aseqlJo slueluele eqrJcsopot
pesn surJel aqlJo eruos epnlcur osle qcrqa'r'z
p'de '9'Z '9'e 's8rJ ur u.{rogsele sluouela urroJ
-runcedFcrd,(I 'I U olqel ur pelsll ore sluoul
-ele uuoJrurlrod roJ asuDaJJ eql ur pesodoJd
srrrreJ 'suoq?urquroctueraJrp ur 'elerJdord
e lsnf
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uo pes?q
e^nducsepE ^reJ
asr^epol
^Solouruual
elqrssodsE^\
lr'fuoEelm od?r{sJofeu olEurs?Jo
sluelJ? se su \oJcJo sedlt eseqlTI?AurpJESoJ
,(q 'uorlrppzu1 snonqo ,(Solouoql"ql sa{Errr
E ur pequtsop eq pFoqs snqt pu?
l?qt
^?,r
sasnlEr"dde
lelale{s snoue^ ur suoprsodJ?II
-Iursperdncco ? su.uoJleldpu? 'sel"ld 'sapelq
s",r lr esnEJeqsroqtn?
13ql reelc elmb ueql
^q Jo..selqd,, poqqnp
snol\eJd
..suuoJ1eld,,
stueuela ^q
opnlJur ol (.,uJoJrurped,,luJoJ eql
uI) ujr01eqtJo uorlrugappeuapsorqI 'ra^e,ror{
'sluoPouoJuo orljnlo^ aslpalJ 186l etll uI
'sJoqlnEluonbosqns
,(q ..sepelq,,peuJel
^uetu
(
F
pue
Jo
eqlJo
l"Jlaoroec
,,t'ororuosled
orerqoue^ul
uo
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ol pesn
suuol puE'so^qe^uep
^rorcos oleuEldpu? oleqd"rs
".,r".rr'iilrufl*?,fi''l,qliilS
^sounoc
Jreql ,sluouele uloJruncod eteurturec
pue elqnauy_ .9:i .a1i
:
$Y
oleqoecsrurturec
ol Su Itt r Sc
FFy
ge^Iurl pue
o nas reddn
bs 't'z '3r.{
'mpolstou
o eurosJo
uroJ eleurl
rr$od 'Jou
*uql o^Eq
jpuus0d
Frp qcueJq
lo Jouel
I iesseroJd
D stuauap
luelue8uEr
SuIpuodop
I ur PePr^
EroJl?ld
EEq po!e^
gqr a,toorE
d aql r{teeu
e
elelnbue
a l e u el dr l nE ue
-
sern6ll lte -.-
d
ur6Jeut
leseq a^rssacol lo euoz,ull2
s s aaor d l er el et l l epuo a e s.d l s
sseJoJd roualsod -dd
ssacord leJalel AJeul|ld-dld
ulJolleld_ld
taal_r
al .r l u a p - p
d u l u c- l t
e u u e a ,e c
d sn c- a
d leseq-dq
a p e tq - tq
a^ oor6 l e se q _ 6 q
A l r^ e. l e se q ,Jq
oteltns luetrlqcelle-se
sseco,o lot,ralue.de
a^ oor6 l euUe cp e - 6 e
oz
VJNOOONOJ AHJ
2l
SKELETAL ANATOMY
may havea narrow medial lodina) the anterior process is commonly
neaththe processes
groovethat is flankedlaterallyby zonesof in- adenticulateand is represented
only by a conspicuousflangelikerim on the anlerior margin
vertedbasalmargin of varying width.
Platformlesspectiniform elements are di- of the cusp. Pastinateelementsof this type
vided in Table 2.1 into five major categories, havebeentermed "dichognathiform"in some
dependingprimarily on the number and ar- of the descriptiveliteraturebecausethe genus
rangement of processes.Stellate pectindorm Dichognathuswas basedon such elementsin
elements(Fig. 2.5) have at leastfour primary the heydayof form taxonomy.
processes,of which two are anterior and posCarminate and angulate pectinifurm eleterior. One or more of these processesmay ments (Fig. 2.6) have two primary processes,
which are regardedasanteriorand posteriorin
branch distally to form secondaryprocesses.
(Fig.
2.5) position.The categoriesdiffer only in that the
Pastinate pecliniform elements
which are ante- longitudinal axis of carminate elements is
have three primary processes,
rior, posterior,and lateral in position. In pas- straight, or essentiallyso, in lateral view,
tinate componentsof the skeletalapparatuses whereasthat of angulateelementsis archedbeof some Ordovician conodonts(e.9.,Neomul- neaththe cusp.Carminateelementswerecomtioistodus, Multioistodus, Phragmodus, Plec- monly termed "spathognathodontiform"and
trig. 2.7. Segminarepectiniform elements, their scaphateand planate derivative$, and terms us€d to describe them.
Upper sets ofdiagrams from Treatise on lnvertebrate Paleontology, courtesy of the Geological Society of America
and University of Kansas.
fr ' 5. 1.6.and
Jte rcrmsused
F.
l dae- or platp d similar in
pe rne apexof
L oanr asfour
L tbe .u.p. rn
I demen$, prohigher than
I- On the lower
bas of the athd in grooveh rprel cavity,
the cusp;
ra
-rlh bsal pit berrt srface be-
proccaloa
brsogminiscophato
trisogminigcsphots
iIJql suoD?l
E1. JrSoloqd
Fr3ui lqtrru
IXSUOTSnO
$ 'la^eA\oH
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u aql plseeJ
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F"a[ 1? Uoeq
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r s?^1 qcrq^\
tFoloqtuotu
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Fr q3Eeleql
meerqgc
l4-,fuleurru,(s
ale tEql snl?J
6q?Jl-arnl€
rIP SneeJq?d
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p Jo suorsJe^
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m 6 z '8rdJo
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roJeJ se 's?l?J
lals aql uI
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alqeu? eJ€
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xt
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VINOCONOJ
AHI
SKELETAL ANATOMY
It Th.se. in turn,
.1 en dolabrate
rliologic
inter5 '-:< 6rst to recfu -..-ies. and he
d. 1pes. Subserr€rt-d out, and
th .iereat [email protected] ihe skeletal
ris :iat are repE3
I illustrate
rries n Fig. 2.8,
IIr .]s€q'here in
series int
n or seven
--:on
'E ::e complete
L *apes of elet..a
are
-ries
E-1
difficult
to
fcncgrneric sper &enls
alone.
tr3 of elements
I c-E
in the appah
aFaents on the
!r=_jopedarc eleiipr_:
or dolabrale
j
U
in the symmetry-transitionseriesmay be of
considerablehelp in distinguishingbetween
conodontsthat belongin differenlsuprageneric
stocks.Thus it is important to pay carefulattention to the correctassignmentoftransitionseriescomponentsduring the systematicstudy
of any collectionof discreteelements.
In a few conodontspecieswith apparatuses
composedsolely of coniform elements,it has
beennoted that positionsin a symmetry-transition seriesmay be occupiedby elementsthat
are variable with respectto longitudinal curyature. Fahraeusand Hunter (1986) have recently termed these variable gtorrpscurvaluretransitionseriesa\d haveillustratedthosethey
regard as characteristic of the apparatusesof
Panderodus gracilis, Protopanderodusvqricoslatus, ajf,d Drepanoistodus sp. aff. D. suberectus.
In the skeletalapparatusof Panderodusgraby Fihraeusand Hunter,
ciri, asreconstructed
four ofthe five componentsform a symmetrytransitionseries(Fig. 2.9).Specimensin row A
ofFig. 2.9 are all bilaterallysymmetrical,alate
coniformelements,but theydiffer from oneanother by slight variations in curvature;specimensin row B. whichare slightlyasymmetric
versionsof the alatespecimensin row A, also
exhibit some variation in curvature, as do
thosein row C, which are otherwiseevenmore
asymmetric versions of elementsin row B.
Fihraeus and Hunter tius recognizefive curvature-transitionserieswithin a skeletalapparatusthat alsoexhibitsa three-or four-member
symmetry-transitionseries.
Fihraeus and Hunter evidently conclude
that eachindividual of Panderodusgracilis had,
a skeletalapparatusin which there were five
morphologicallydistinct components,eachof
which was representedby a six-membercurvature-transitionseries.Thus there must have
beenat least 30 elementsin the skeletonof a
complete individual. Fihraeus and Hunter
regard the nature and number of such curvature-transition seriesas featuresof potential
supragenericvalue in conodont taxonomy.
However.they do nol seemto have given serious considerationto the possibilitythat they
rnight merely be charting the breadth of morphologic variation within the particular populationsthev samDled.
A
Fig.2.9. The curvatur e-transilion seies of Panderodus
gracilrs(Bransonand Mehl). Each of the five or six elements oflhe apparatusis shown to increasein curvature
away fiom the planeofbilateral symmetry.Inner lateral
views on lefl; outerlaleral views on dght. Redrawnfrom
F6hraeusand Hunter ( 1985).
2.5
SkeletalApparatuses
We have thus far consideredthe composition,
classification,and descriptiveterminologyfor
discretecomponentsof the skeletalapparatuses
ofconodonts.However,it hasbeenknown for
at least the last 50 years that in many conodonts,perhapsin all of them, the skeletalapparatusincluded elementsof severaldifferent
shapecategories.Thus, in order to compare
one conodontwith anotherand to build a biologically reasonabletaxonomy for the group,
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VINOCONOJ
IIHJ
SKELETALANATOMY
for elements
B nol intended
of€lement$;it
rnajor poslon IDverteSociety of
Eaming ho-
4rFratus. In
€f rhe systems
rhis and conscheme illus210. That
frct that the
qEcres reprecan be
catego-
S in Fig.2.10.
are occu['specialized
digyrate
two types of
ma;- be desigpmitions they
aDd Pb posi
it-shaped
(or
may be fiched
drgyrate
ramiform elements or geniculate coniform
types.Elementsin the threemajor positionsin
the S seriesform a symmetry-transition series
of the sort first recognized by Lindstrdm
(1964).The Sapositionis occupiedby an alate
coniform or ramiform element;the Sb position
by a digyrate or tertiopedate element; and the
Sc position by a bipennate or dolabrate rami
form element. However, the nomenclaturejust
mentioned (and illustrated in Fig. 2.10) provides formal identification for just the midpoint and the two extremesin the S series.In
the apparatusesof many conodonts, however,
there may be more than three morphologically
distinct componentsof the S seriesand, to describe and locate them, it may be necessaryto
invent intermediate categoriessuch as Sa-b or
Sb-c.In Fig. 2.10 thereis alsoa positionin the
S serieslabeled Sd, which is occupied in some
reconstructed septimembrate apparatuses by
quadriramate ramiform elements.It is not appropriate to use Sd as the designationfor intermediate positions in the symmetry-transition
series,as some authors have recenfly done, In
most skeletalapparatusesthe Sd position is not
filled at all.
2,6 SymneFyof Elements,Elem€ntPsirs,
and Apparatuses
It takes rrery little experiencewith collections
ofdiscreteconodontelementsto discoverthat,
like componentsof the human skeleton,most
of them corne in mirror-image pairs. The only
ones that depart very obviously from this pattem are alate coniform and ramiform elements, which are themselvesbilaterally symmetrical, and a few straight pectiniform
elements that are developed in the same way
on both sides.Indeed, it was not until almost
1960that it becameobvious that the right and
left (or dextral and sinistral) mernbersof some
element pairs are not exact mirror-image replicas of one another; and the incieasingly widespreadpractice of multielement taxonomy has
led to the discovery that bilateral asymmetry
wasrather common in the skeletonsofat least
certain groups of conodonts.
Iane (1968)suggested
that four major types,
or classes,of symmetry could be recognized
among various element pairs. These are illus-
trated diagrammaticallyin Fig. 2.1L In Lane's
scheme,C/ass I symmetry is exhibited by elementsthat are themselvesbilaterallysymmetrical. For suchelements(e.g.,alateconiform or
ramiform elements) no pairs can be established,and it is possiblethat they occurredsingly in skeletal apparatusesor were paied with
other individually symmetrical Class I elements. C/aJs11 slrmmetry is a feature of individually asymmetricelementsthat are represented by approximately equal numbers of
"rights" and "lefts" in largecollectionsof discrete specimens.As lane (1968) noted, the
bulk of known typesof conodontelementsapparentlybelongin this class.
ClassIII symmetry is exhibited by conodont
elementpairs in which the "right" and "1eft"
Fig.2.ll.
Various symmetry classesexhibited by elements typica.l of (I) R hipidognathusi (Jl\ Xaniognathus;
(llla) Eognat hoduii (lllb) A mory hoghalhus.
4J rnz 18961)
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VINOOONOJ EHJ-
SKELETALANATOMY
i
-'-csr cannot be
r::Tor-lmage (or
h:-<lement pairs
g:nerally held
t
;rfoe=selves bilatr ccrbers of ele: p::s may have
r*-=-eron or body,
F-E sides of the
lE :on'acknowlls :irh Class III
E:r
a monkey
E::---{r
to sugd :errs rn their
I ti-. e been bilatr -'r-lleral asymE i- --Ie skeleton
Ef,lnents
were
k
=e animal as
n. r-teral symleature to
-:r.r oirhe mode
b:r
G -'llnodonts in
L
k-s:---om. S. M.,
k:c=:.
F. H. T.,
I-=istrom. M.,
\ G ,9El ). Conrc.::a
PaleontolL I \fi5cellanea),
flme:,-a and Univ.
rl:s-ion
of conllfD- R 1986).The
h:-=:al part of
l4ttri:-.ruses (PanE- C-.nodonlata).
---189.
Dar
IE =r Conodonb
ft
;e: Gatrungen
C:xodontida).
r a.:,:-bed and its
I ,i;:. Soc.Lonnir:s
from the
|' :, -le Cambroih-. ard Genesee
fu = Canadaand
the United Stltes. Quafi. J. Geol.Soc. London
3 5 ,3 5 1 -3 6 8 .
Lane, H. R. (1868). Symmetry in conodont element-pairs."L Paleont.42, 1258-1263.
Lindstriim, M. (1964). Conodonts.Elsevier,Amsterdam,London, New York, 196pp.
and Zregler, W. ( 1971) F€instrukturelle
-,
Untersuchungenan Conodonten. l. Die Uberfamilie Panderodontacea.Geol PaloeonL5,9-
Mikromorphologie der Cor.odor]]!e[ Paleontographica, 128, ll 5-l 52.
Roundy, P. V. (1926).The micro-fauna.In, P. V.
Roundy, G. H. Girty, and M. I. Goldman, Mississippian formations of San Saba County,
Texas. 2. U. S. Geol. Sum. Prof. Paper 146,5-
Sr".t, W. C. (1979). Late Ordovician conodonts
and biostratigraphy of thae western Midcontinent Province. Brigham Young Univ. Geol.
Pan-d'er,
C. H. ( 1856).Monographie der fossilen
SLudies26,45-85.
Fische des silurische Systemsder russisch-bal- (1981). Macromorphologyof elements
tischen Gouvernemefis. Akad. lliss., St. Peand appa.atuses.Pp. W5-W20. ID Treqtiseon
tersburg,9lpp.
InverlebretePaleontology(ed. R. A. Robison),
Pietzner,H., Vahl, J., Werner, H., and Ziegler,W.
Pt. W, Suppl. 2, Conodonta. Geol. Soc. Amer(1968).Zur chemischenZusammensetzung
und
ica and Univ. Kansas,202 pp.
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AI^IOJVNV'IVruINV-A'IOH,&\'g
WHOLE-ANIMALANATOMY
29
2mm
G_-i:3nt aspects.
:f:::s-,1!cted from
E- iirrally comE
ts.:.ld
specimen
ir
i in m (1 .6in. )
,.0 7to 0. 08
i ==
lE:r-sron at one
!:c,-:t apparatus
r.=:::1ic lobe.On
r*: _'rdged to be
t =:: :reted to be
f::i
r. the bestlr i:!i diagnostic
ak ::,r of indisr:i .r'rtenor end.
hc ::-.:r- the axis
L: : ilrn ct l i near
ft::: :s a seriesof
!bi: srop€ toward
rs a-. more dise-, :d specimens
EE :: :.1.( 1986),in
Fe : \--like shape,
E:: rhe anterior.
rtt='r specimens,
I
a]rost to the
I{: -:r'er. in that
E=:sr \-'s appear
di---'< as a result of
k-: --eburial and
ft=: and still the
c$::s:r Specrmens
-l - --:e V-shaped
E:r 3h laken to
| ::re compared
(or muscle
b=5
L B--: lhese inter'equivocal."
d
aL- i iS6) describic= rhe Scottish
Fig. 3.1. Clydagnathlls? sp. cf. C ca,rutbrmis. (A\ Cafiera-lucidadmwing of part; (B) and (C) enlargedvrews of
head showing conodont assemblagein part (B) and counterpan(C).Specimenfrom "shnmp-band" in Granron
Sandstones(Dinantian), Edinburgh,Scolland.Redrawn from Bflggs,Clarkson,and Aldridge (1983).
Dinantian locality, these V-shapedstructures
are identified, with no further discussion,as
"somites." This interpretationwas obviously
quite influential in the much more subjectiye
discussion
ofanatomyin the 1986paper.even
thoughit is in no way requiredby the evidence.
Outlinesofthe anteriorend are preservedin
only one of the ScottishCarboniferousspecimens(Fig.3.1).Althoughthis end is termedthe
"head" by Briggset al. (1983)and Aldridge et
al. (1986),thereis no evidencein it ofa brain.
The term cephaliclobe may thus be more appropriate. As noted previously, the cephalic
lobe of the best-preserved
Scottish specimen
enclosesa completeand apparentlylittle distorted conodontapparatus,which permits the
specimento be identifiedas Clydagnathus'l
sp.
cf. C. cavusformis.
In the only Scottishspecimento preservethe
cephaliclobe,right and left componentsofthe
insteadof
skeletalapparatusare superimposed
arrayedon oppositesidesof a line of bilateral
symmetry.It is thus reasonable
to supposethat
the cephaliclobe of this specimenis preserved
in lateral aspect,that is, that the plane of the
bedding approximatesthe plane of bilateral
symmetry.The deepcleft at the anteriorend of
the cephaliclobe may thus mark the position
ofthe mouth, and the positionofthe apparatus
within the cephaliclobe may indicatethe general position of a buccal cayity. If the entire
specimenshownin Fig. 3.1 has beenflattened
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\_\.aA
.'. ,
4/
_r'
>-. \
hP;r.
I
k::.
ol .e.t.tu.
of Scotland.
d =e apparatus
t i ';nfonunately
I o-rhrr anatomic
J:!e elements in
tr
a basal plate,
f E::sculature as! Er:r more puzig rhar might be
fr :-.rrrcture comd :-ae hagfishes,
r Fbst|ate for the
r d- [email protected] of
I (onplex apparamens provide no
! r-olodont appaqrrielium
or exHer
elements of
I eld might thus
I b.'5 or whether
I tt':s may have
rsi.
supports for
Lc
in a buccal
ll
The trunk of all four of the Scottishspeci- requirethat all segmentsofthe axial structure
mensis 1.2to 1.8mm wideand preserves
evi- in everyspecimenrepresentthe sameorganor
denceoftwo distinctiveanatomicfeatures.The organsystem.For most ofits lengthin all ofthe
most striking consistsof a seriesof V-shaped Scottish specimensthat show it clearly, the
structures,which were apparentlysituatedon axial structurejoins the apicesofthe V-shaped
the sidesof the animaland whoseapicespoint structuresofthe trunk. No suchrelationshipis
anteriorly. In the most completelypresewed shownin the anteriormostpart of the trunk of
specimen(Brigeset al., 1983;Fig. 3.1) these the best-preservedspecimen (Briggs et al.,
structures are rather indifferently preserved 1983;Fig. 3.1),however,and the axial structure
and are confined to the posterior half of the in that specimenmay not be continuouswith,
trunk. In the three more fragmentaryspeci- or represent,the sameanatomicfeature,asthe
mens (Fig. 3.2) describedby Aldridge et al. discontinuouspair of axial lines in the poste(1986),the V's arebetterpreservedand may be rior half of the trunk. Aldridge er al. (1986),
seento extendnearlyto the anteriorend ofthe who allowedtheir interpretationofthe Scottish
trunk. The shape,serialrepetition,and lateral specimensas primitiye chordatesto restrict
situationofthese V-shapedstructuresare rem- their survey of anatomic possibilities,suggest
iniscentofthe myotomes(or muscleblocks)of that the double lines in the anterior parts of
Branchiostoma(or "amphioxus") (Fig. 8.1) severalspecimensmight representsome parts
and of other chordatessuch as the hagfish or all of a nerve chord, notochord,or dorsal
Myxine (FtE.8.2).
aorta.I suggest
that one might alsoconsiderinBut chordalesare not the only organisms terpretingthis pair of lines as the incomplete
thatmightyieldsuchV-shapedimpressions
on trace of a nemertinerhynchocoel,the tubular
compression.For example,nemertines(phy- cavity within which living representativesof
lum Rhynchocoela)such as Nectonemertesthe curious group of invertebratesstore their
(Fig. 8.1) are characterizedby a long, straight eversibleproboscis.Suchan interpretation,of
intestine,which bearsnumerouspairedlateral course,would not rule out interpretingaxial
diverticula that might, on flattening and lines in more posterior parts of the trunk as
producepatternssimilar to those traces of a gut with numerous lateral
compression,
exhibitedby various parts of the severalScot- diverticula.
tish Dinantian conodonts. Nemertines, of
If the axial structurein the posteriorpart of
course, are pseudometamericbut acoelo- the lrunk of the Scottishsflecimensis intermate invertebrates,probably derived from pretedas a gut, it appearsto have extendedto
flatworms and not known to have a fossil the posterior extremity of the tail in the two
record.
specimensthat retain it (Figs. 3.1 and 3.2).
The secondfeaturerecordedin variousways That would mean that theseanimalslackeda
and with varyingfidelity in the trunk of all four postanalsegment,or true tail, featuresthat are
Scottishspecimens
consislsof a line. or a pair characteristicnot only of chordatesbut of inof subparallellines, more or less coincident dividuals in virtually all the deuterostome
with the longitudinalaxis ofthe body and ex- phyla. On the other hand, if one assumesa
tending from a point just behind the cephalic nemertinemodel, even with tonguein cheek,
lobe on the anteriorto the posteriorend ofthe the problemvanishes.for in that g.roup-as in
body. Aldridge er al. (1986) note rhat these most invertebrateswith a completedigestive
lines might representa number of different tract-the anal end of the digestivetube is sitstructures.For example,they might be tracesof uatedat the posteriorextremity of the body. I
a notochord,a nerve chord, the gut, a major agreewith Aldridgeet al. (1986)tbat anatomic
longitudinal blood vessel,or a mesenteryor interpretation of the axial structuresof the
septumdividing the body cavity into compart- Scottishspecimensis inconclusive;however,I
ments.I agreethat a gut is the anatomicfeature suggestthat, in combination with other feamosl likely to be representedby most of the tures,thosestructuresmight well tum out to be
lengthof the axial lines in most specimens.as significantin reconstructingdetailsof conoHowever,evidenceavailablethus far doesnot dont soft anatomyas the V-shapedstructures
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VINOCIONOJ AHJ
WHOLE.ANIMALANATOMY
3.3 HistologJof Demineralized
Tissue
i F^ire
ft=:,:'.
Formation
published
19f,-,, The WauG
-One-rs prelrile bmken edge
atrd repretrG
E E:anor part of
;fu,.v,a
eriginal
frte .-tphalic apilceg anes transll r::.k and with
Smith,
lE--rorlllat this
t-esla5trafillus was
qE\ls.d
halyes,
ds longitudinal
t
iobe and the
&55a specimen
rai ro rhe plane
lEr1le. considerin the preI--'.5.:hr My was
Had it
trcssed
to have
r +ears
I Carboniferous
t ba..e been de|l oo compaction
lFiles sery little
aaatomy of
lo$
E Lbar vague,
lrss original seglcr aI { 1987),the
pible
segment
lecrlr ransverse
There is no indiI (ia.racreris tic of
soft-bodiedanimalswith a slender,vermiforrn
body
a couple of inches long that may have
Fdhraeusand Fahraeus-vanRee (1987) have
been
either
laterally compressedor dorsovenidentifiedcells,fibrousmaterial (thou8htto be
trally
depressed.
The anterior end of the slenat leastpartly collagen),and luminaeofvarious
der,
elongate
body
is developedasa distinctceshapesin scrapsof demineralizedtissue dephalic lobe in known Silurian and Dinantian
rived from the pectiniformelementsofSilurian
specimens of Ozarkodina confluens. Cells of specimens and includes the apparatus of
vanous types have been identified; some are mineralizedelementsby which conodontsare
relatively large,with nuclei 3 to 5 pm across bestknown. The slendertrunk is divided into
and long dimensionsas much as 20 pm: others serially repeated transverse structures that
are smaller, contain numerous nucleoli, and occur on either side of or surround an axial
structurethat may representthe gut, a notohavea stronglyeosinophiliccytoplasm.
chord,
a doral aorta,a longitudinalseptumor
Luminae, which have the form of irregular
mesentery,
or somecombinationof thesestrucspherical spaces,and single and branching catures.
The
transversestructures,which are V
nals,are relatedby Fihraeusand Fihraeus-van
shaped
in
Carboniferous
specimens(but not in
Ree to the irregular frothy spaceswithin conothe
Silurian
one)
have
been
comparedwith the
dont elements that have been termed white
myotomes
(or
muscle
blocks)
of amphioxus
matter. Flbrous matter is sparselydistributed
and
fish
and
thus
taken
as
evidence
of body
among cells, occurs in bundles and, for the
segmentation.
Those
interpretations
are
not remost part, lacks nuclei. Some is dense,basoquired,
however,
and
the
transverse
structures
philic, and apparently associatedwith elonjust aswell representthe paireddigestive
gatedsinuouscells.The fibrousmatterwithout might
nuclei is consideredto be collagen.Fihraeus diverticulathat characterizemany acoelomate
and Fihraeus-vanReeemphasizethe fact that nemertrneworms and, as such,indicate only
pseudometarnerism.
The posteriorend of two
onty 20 to 25 percentof the tissuefragments
of
the
flve
known
whole-bodied
fossil conthey studied is collagen,whereascollagenacodonts
is
bluntly
aciculate,
and
its
margin
countsfor the entirematrix ofmost othertypes
bears
traces
of
ray-supported
finlike
structures
of mineralizedtissue.They also point out that
the largenumberofcells in demineralizedcon- on oneor both sides.Linearaxial structuresexposteriormarginin bolh specimens.
odont tissueindicatesthat it ". . . is not bone tend to the
Ifthese
represenl
tracesofthe gut, the Carbonor dental tissueas we know it," becausebone
iferous
conodonts
may not havehad a postanal
". . . is eithertotally non-cellularwith an extrasegment,
or
true
tail.
cellularmatrix largelyconsistingofcollagen. . .
The five whole-bodiedfossilconodontsnow
or, in cellularbone,it is dominatedby bundles
known
raisealmostas many questionsas they
ofcollagenwith the occasional'trapped'osteoprovide
answers.It is a comfort to know that
cyte." They go on to statethat the demineralthe
skeletal
apparatusis apparentlyassociated
ized conodonttissueof their study is also unwith
the
cephalic
lobe,for most ofus had conlike that found in living inarticulate
cluded
from
seat-of-the-pants
functional analbrachiopodsor mollusks.They do not exclude
ysis
that
this
would
turn
out
to be the case
the possibilitythat primitive vertebratesmight
years
many
before
the
famous
Scottishspecihaveformed suchtissues,nor do they fule out
mens
were
discovered.
It
is
also
no surprise
possibility
the
that the tissuesthey studied
that
the
specimens
indicate
an
elongate,
verrepresent a ".. . unique approach to biomiform
shape,
since
that
is
a
shape
common
to
mineralization."
mobileaquaticanimals.However,eventhough
the skeletalassemblages
includedin the wholebodied
fossils
seem
to
have
beenonly slightly
3.4 Summary
distorted by compression,they all apparently
Existingevidenceon whole-animalanatomyis lack basalbodies,and it is not possibleto desparsebut may be takento confirm the conven- termine how they may have been related to
tional wisdom that conodontswere prirnarilv muscularsystemsor whetherthey were com-
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'f d'e8pupN pu? ''D g q 's8:lug 'd 'I^t 'tIlItuS
'dd 16'[email protected] 1S
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VJNOCONOC EHJ-
lGment rn con*-1 t-rpe and in
-l:1.
l . r 197 3 ).ConI tt Bear Gulch
aC...'-dont PaleoI G;,-'i. Soc.Am.
4. TAXONOMY
G- a-ld Kluessenf"j-rcdred biota.
4.1 FormTa-xonomy
aib,- A new exr -3e Lower Silfu- Tens. Roy.
$,
h-uon of the Pa
---90 in Pad
L R- J, .{dridge).
lre-
der fossilen
;le
r&r russisch-bal+t .t\rd. Wiss.,
d t"lridge, R. J.
h
te lower Sid --:e apparatus
I csqodonts. 9lE-{-r (ed. R. J.
bssrer- 180pp.
Pander'sclassificationofconodontswasproposedlargelyas a meansof giving order to the
In Chapterl, I pointedout that ChristianPansystematicdescriptionof specimensin his colder, who discoveredand named conodonts.
lections.He made no attempt to give it prowrestled in his 1856 monograph with the
found biologicmeaning,nor did he useit asthe
choiceoftaxonomic base.He wrote (my rough
basisfor discussingeyolutionarydevelopment
translation):
or addressingother biologic questions.Evidently
the classificationwasto serveonly until
We don't know what sort of teeth we have before us;
wheth€r they belongto the jaws, the palale, the lips or
discoveryof more completespecimensmadeit
the tongue; whether each particular shape conslitutes possibleto determineif all elementsweremorsufficientbasisfor establishmenlofits own genusor spe- phologically
alike in the body of a singleindi.
cies; or whether different forms could have come from
vidual,
or
if
the body of the sameindividual
one and the samebody.
might havehadan arrayofmorphologicallydifIn the absenceof a living analogue,and un- ferentelementswithin it.
able to answerthe questionshe raised about
Although Harley, Smith, Hinde, and Hadfunction or anatomic position or about mor- ding contributed to conodont taxonomy bephologic monotony or yariability of elements tween 1856and 1926by describingdiscreteelein the samebody, Panderchosean admittedly ments of distinctive shapes as genera, no
utilitarian classificationbasedon shapeof in- further consideration of supragenericcategodividual elements.This approachto the classi- ries seemsto havebeenmadeuntil 1926,when
fication of biotic groupsis sometimestermed, Ulrich and Basslerpublishedtheir influential
rather pejoratively,form taxonomy.
taxonomicscheme.As you will recallfrom earTo beginwith, Panderassignedconiformele- lier discussions
in this book, Ulrich and Bassler
mentsin his collectionsto an informal category stated their firm conclusion that conodonts
termed "simple teeth" (Einfache Zahne), and represented
not only the teeth,but alsothe derramiform andpectiniformelementsto another, mal platesof extinct groupsof fishes.In additermed "compositeteeth" (Zusammengesel.ztetion, and probablyrnoresignificantly,they also
Zahne). The s.impleteeth (Fig. a.1) were then concludedfrom their studiesof recent fishes
further divided on the basis of crosssection, that, exceptfor the fact that they may occuras
into 7 categories,
which Panderdescribedand right and left mernbersofa pair, all teethin the
named as genera.Finally, each of these was mouth of the supposedliving relativesof the
subdividedon the basisof cuspcurvatureinto conodonts are essentially alike. Thus they
groupsregardedas species.Compositeteethin wrote ". . . eachkind is characteristicof some
Pander'scollectionswererepresented
by 7 ob- particular genusand species."In short, they
viouslydifferentmophologiccategories.
among professedcompletecoDfidencethat a classifiwhich he recognized18 specieson the basisof cation basedon the shapeofdiscreteelements,
featuressuch as mode of denticulation,pres- a form taxonomy,wasa true expressionofbioenceor absenceof carinae,anglebetweenpro- logic relationships.
cesses,characterof platform extensions,and
In their 1926 paper Ulrich and Basslerigthe like. Altogether,then, Pander recognized nored the one genus(Periodon)proposedby
and named 7 generain each major category Haddingin 1913,but distributedall the others
(simpleand compositeteeth)and, collectively, known at that time, and 15 new ones,among4
56 species,all basedon an analysisofshape,or supragenericcategoriesdescribed and named
form.
as families. Pander's"simple teeth" category
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VJ,NOOONOJ lHl-
9t
TAXONOMY
lridia)
and de:sembled into 8
: tgo suborders
hodontiformes)
irhose elements
h srborder NeuE \-rcian genera
rlought to have)
bas
the suborEilshed for the
lresented by elef irEmal strucG Eere thought
Erendl' shaped
r|rd ioro catego)bsrs of lhe supr tle variously
i on x'hich they
bn<. The three
fu:dae.
for ex! hsil record by
rirh
relatively
rllich
were
I
roirhe fish that
i
oi conodonts
J irdiridual,
dislfu a,,-mein Volat*ate
PaleonP- h rlat highly
rF
entirely by
Ddouts were asC-onodontophorI r Iamrtres. lne
h Branson and
iorders
Neurohmes were reEsrent by Hass,
![\ze those subrc ramilies (Co,-d ldiognathotEmilies so that
EL-sones recog;nred *'ith 8 in
r4 in that of Ut-
I ahernative to
pied
as a pospractice
rnto
IEI
37
until 1879,when Hinde employedit for a spe- and an essentiallycomplete "natural assemcies from the Devonian of New York. In the blage" is preservedin the headof the Scottish
"multielement" way of looking at things, the "conodont animal" discussedin defail in
skeletalapparatusofa conodontis madeup of Chapter 3. So, since at least 1934,there has
numerous parts, not necessarilyall alike in beenno good reasonto defenda form taxonmorphology.The basefor taxonornyis thus a omy for conodontsasa natural one.The pracgroup of elementsthat can be shown, or can tice persistedthrough 1962,I suspect,because
reasonablybe inferred, to have occurredto- most studentsof conodontswer€ more congetherin the body of an individual conodont. cernedwith biostratigraphicapplicationsthan
An apt analogy,ofcourse,is with the vertebrate with biologictaxonomy.
skeleton,which consistsofnumerous morphologicallydifferentparts.
Hinde (1879),the first to practicemultiele- 4.2.I Multielementmethodology
ment taxonomy, based his concept of Polyg- The major difficulty with putting a general
nathus dubius on an assortmentof elements schemeof multielementtaxonomy into pracpartly exposedon the surfaceof a slabof black tice for conodontshasbeenthe fact that "natshale.He took physicalproximity on the shale ural assemblages"
and fused clustersare relaslab to indicate original association,although tiyely rare and are not uniformly distributed
the namehe chosefor the first speciesto be di- stratigraphically,whereasthe collectionsavailagnosedin a multielementsensesuggests
that ableto most systematicists
arejumbled assorthe was not completelyconvincedthat all ele- ments of elementsthat may have been commentscamefrom the sameindividual.
ponents of several different multielement
Other, more convincing "natural assem- apparaluses.Thus, except for speciesrepreblages" of elementswere describedindepen- sentedby natural assemblages,
a meansmust
dently in 1934by Hermann Schmidt in Ger- be devisedfor determiningwhich elementsin
many and by Harold Scottin the United States. collectionsof discretespecimenswere origiThe assemblages
availableto thosepaleontol- nally associated
in the sameskeletons.
ogistsareall from Carboniferousrocksofabout
Perhapsthe most instructive way of reconthe same age and are made up of about the structingthe skeletalapparatuses
of conodonts
sametypesof conodontelements.Becausethe from largecollectionsof discreteelementsis a
compositionof Schmidt'sand Scott'sassem- basically empirical method. This procedure
blagesis closelysimilar, and becauseeachhad wasappliedasearlyas 1958by Huckriede,who
severalspecimensof sirnilar or identicalcom- noted that in his collectionsof Triassic conposition, chanceassociationwas easily ruled odontscertainforms occurredmoreor lessregout, and the assemblages
were describedand ularly with certain others.From his observainterpretedas the skeletalremainsof individ- tion that certain types of ramiform elements
ual conodonts.E. B. Bransonand Mehl (1936) wereregularlyassociated
with one or the other
and C. C. Branson(1957)attemptedto dismiss of the two most common platformedpectiniScott's specimensas coprolites,but Scott re- form elements,Huckriede proposedthat the
plied (to Bransonand Mehl) that ". . . it would " Conodonten-Satz"
(literally, conodont-set)of
be strangeindeed to find a group of animals Gondolella navicula (the platformed pecriniwith sucha balanceddiet that the excretalma- form element)includesa pair of angulatepecterial would consisttime after time ofone pair tiniform elements(refened,thenLoOzarkodina
of prioniodids, one pair of spathognaths,
one tortilis), a\d ramiform elementsthat wereinpair of prioniodells,and approximatelyfour cludedin 1958in 9 form speciesassignedto 6
pairs ofhindeodells."
form genera.Huckriedecomparedthe associaThere are now more than 500 "natural as- tion he recognizedempiricallywith naturalassernblages"of conodontsknown from Cam- semblagesdescribedby Schmidt (1934, 1950)
brian, Devonian,and Carboniferousrocks,al- and Rhodes(1952)and evidentlynoted agreethough not all have been described and ment in at leastmajor features.He appliedsimillustrated.In addition, fused clustersof ele- ilar reasoningto a reconstructionofthe ",Salz"
ments now and then appearin acid residues, of Polygnathus tethydis, with results that are
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'(6961) tnqo)
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eql Jo pu? eseq crurouox?l eql Jo uoIsIAeJ Jo Jsn JeJUrduJ Jpsrll 'uotls ut 'apJu{tnH
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VJNOOONOCAHI
IAXONOMY
indonts did so
r--ald
a major
bE ard of the
lEries in the reH s-ith enthusi
I ad voD Bitter
fusring
meth- iterily
6o"5hcrete elements
groups that
E
I apparaluses of
lql
sp6511u1a4
rf pincal methbers. atd BergI rhai it became
E to subdivide
tinsial or facies
tRlement
spetl s.lme conclub hern een elellh apparatuses
*s of the clusLFt- survey, alrihl
drat these
rs€ator is dealof samples
C
ftrms
and are
-er
iicate rhat rergrrence of 15
ELs in l0 sam$'ould be
nist
h
lr rhat table
Dq-currence of
b Ibar might be
14
:_i
I
I
x
I
x
I
\
x
\
\
39
pairsfrom Table4.1
TABLE 4.2. Jaccardcofficientsfor element-type
Type 1
l0
11 12
56
22
-_
I
2
3
5
6
7
8
9
l0
61 10 18 33 56 J0 ?1
11 40 44 29
22 11 80
88 ?0 60 7l t00 6't 33
-807063887844893850
-9050?08050905040
-4060?056805644
1t 63 29
78
44
18
1t
80
33
13
57
60
38
14
56
22
00
88
70
60
7t
00
67
33
78
33
38
- , : t r 3 1 31 3 3 3
-
ll
t2
l4
s6
56
33
29
100 50
56 44
-50
t)
7l
80
33
44
50
56
29
33
30
100
56
100
50
33
lf
i{ot? Decimal pornts omitted:"100" = 1.00;"50" = 0 50, e1c.
consideredto representthe skeletalapparatuses
of various conodont species.Severaldifferent
coefncients
havebeendevelopedfor usein stating the degreeofassociationbetweentwo entities,and eachstudenthas a preferencefor one
or another.Kohut (1969),for example,usedan
index of affinity developed by Fager and
McGowan (1963),whereasVon Bitter (1972)
employedthe Jaccardcoemcient.Clustersderived from the data ofTable 4.1 are essentially
the same whichever similarity coefiicient is
used; so, becausethey are easierto calculate
than Fagervalues,I havederivedJaccardcoefficientsfor all possiblepairs of the 15element
types in that table. The Jaccardcoefncient,as
usedby Von Bitter, may be given as
_a
J=-
a+ b + c
in which a is the number of samplesin which
both the elementtypes consideredoccur, D is
the numberjn which one of them is present
alone,and c is the numberof samplesin which
only the secondelement type occurs.Obviously,ifthe two typesofelementsdo no1occur
togetherin any ofthe samplesavailableJ : 0;
but if eachof the two is representedin every
samplethat containseitherone of them,J = l.
Ofcourse, ifJ = I for a pair of elementtypes,
we must considerthe possibilitythat thoseelement types were originally parts of the same
skeletalapparatus,eventhoughthey may differ
greatlyin morphology.
In Table4.2I givethe Jaccardcoemcientsfor
all possiblepairsofthe 15elementtypeswhose
distribution is given in Table 4.1. Note that
valuesof thosecoemcientsvary from a low of
0.22to a high of 1.00.What we needto do now
is rearrangethe dataofTable 4.2 to revealclusters of elementtypes more closelyrelated to
eachother than to other clusters.This may be
done in severalways. Kohut (1969)chose,in
effect, to rearrangethe similarity-coefficient
matrix itself, as I have done in Table 4.3, by
rnovingthe highestvaluesin eachrow and column to the diagonalmedianof the matrix. Von
Bitter (1972),on the other hand, usedthe unweightedpair-groupmethodto developa dendrogramthat graphicallydisplaysthe relationship between pairs and clusters of pairs of
elementtypes.I havederived the dendrogram
of Fig. 4.2 from the similarity-coemcientdata
of Table4.2.
In both the rearrangedmatrix of Table 4.3
and the dendrogramof Frg.4.2,note rhat there
are threewell-definedclustersofelementtypes.
Two of theseclustersinclude six rnorphologically different typesof elements,whereasone
has only three. Furtbermore,both the rearranged matrix and the dendrogram reveal the
samethree clusters,which may appropriately
be identified as recutent groups.
In a multielement view of the conodont
world recurrentgroupsare the basisfor taxonomy if their componentsalso exhibit several
other important features.For example,if a recunent groupis to be interpretedasthe skeletal
apparatusofa conodontspecies,
onemight rea-
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TAXONOMY
4.3 MultielementClassifications
t5
l
i:
rr
4r
5r
_=
i:
l3
J-l
t9
{0
=rJ
t_
rl
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ES0
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r RiauoDship to
tcdsr ro be con;rarus of a con!- demonstrate
rca-nc1 in rank
Ried
clusters
he studhJr
-ials to support
criterion
;wal
F as rhe appap r: has become
against
lFaled
;; oi some con-rarl
envrronl!
contributed
+ ihrs \ ardstick
Ef,n
rtr many
41
ment or redefinition ofcertain categorieseslab-
lished in that volume. PersonalprejudicedicLjndstrdm (1970)compiledthe first classificatatesothers.Thus the classificationadoptedin
tion of conodonts that recognizedrecurrent
in somedetailin Chapgroups of morphologicallydifferent element this book and discussed
ter 5 wilt appear,at least superficially,to be
typesas the basictaxonornicunits. Conodonts
quite different from that in the Treatise. Some
were groupedinto 2 orders (Westergaardodibrief commentis thus in order.
nida Lindstrtjm and ConodontophoridaEiFirst, I havebecomeincreasinglyskepticalas
chenberg),with the latter (the "conodonts
to
the amnitiesand relationshipsof the tiny,
proper") further divided into 8 superfarnilies
phosphatizedfossilstermed protoconweakly
(Distacodontacea,Panderodontacea,Chirogodonts and paraconodontsand lumped tonathacea, Prioniodinacea, Prioniodontacea,
getherin the Treatiseas the order ParaconoBryantodontacea,Gondolellacea,and Polygnathacea).These major groups were defined, dontida. These superficially conodontlike
forms are quite different in internal structure
insofar as possible,in mullielementterms, as
from the onesdescribedin Chaprer2, for which
werethe familiesinto which they weredivided.
Bengtson(1976)hasprovidedthe generalterm
Altogether,Lindstraimwasableto assigna total
Although Bengtsonhas shown
"euconodont."
of73 generamore or lessconfidentlyto one or
how paraconodontsmight have evolyed from
anotherofthe 2l familieshe recognized.
protoconodonts,neitherhe nor anyoneelsehas
The taxonomic schemeoutlined by Lindyet documentedthe stepsby which euconostrbm (1970)obviouslyinfluencedthe one dedonts might have developedfrom paraconovelopedby authorsofrevisedVolume W ofthe
donts.
Szaniawski(1982),on the other hand,
Treatise on InvertebratePaleontology(Clatk et
hasdemonstratedthat conicalelementsof one
al., 1981),which waswritten betweenl97l and
of the protoconodonts(Phakelodustenuis)arc
1976but was not publisheduntil late 1981.
essentiallyidenticalin jntemal structureto the
Treatise atJlhots included alt conodonts in a
graspingspinesof modern anow worms such
singleclassof a phylum Conodontaand folas Sagitta. Thns, Phakelodus tenuis, a protolowed Lindstrdm in assigningthe animalsI regard as "true conodonts" to one order (the conodont,may well have beenan early representativeof the phylum Chaetognathaor of
Conodontophorida),which included 44 famisome now-extinctgroup closelyrelatedto the
liesdistributedamong9 superfamilies.
The 179
and not a conodontat all. Szageneraof "true conodonts"recognizedweredi- chaelognaths,
(1987),whosecarefulhistologicstudies
niawski
agnosed,insofar as possible,in multielement
are certainlyimportant in this regard,continterms and described in the terminological
uesto hold that the proto-, para-,and euconoschemedescribedin Chapter 2 of this book,
donts(i.e..the Paraconodontida
and Conodonwhich was developed speciallyfot lhe Treatise.
tophorida of the Treatiseclassification)form
parts of a singleevolutionarylineage.He does
not regardthe Conodonta(in which he would
4.4 A RevisedMultielementClassification
includethe Paraconodontida)
aschaetognaths,
At the time ofthis writing, skeletalapparatuses despiteconsiderablehistologicsimilarities,but
of speciesthat representmore than 100genera as representatives
of a group that developed
havebeenrecognizedfrom naturalassernblagesfrom a common ancestor.It is my conclusion,
or fused clusters or have been reconstructed which is further developedin Chapter8, that
from recurrentgroupsidentifiedin largecollec- the Conodonta,Paraconodonta,
and Chaetogtions of discretespecimens.For the most part, nathaare similar but divergentgroups,all posthe classification adopted in the Treatise pro- sibly of phylum rank, that developed in the
videsa satisfactoryframeworkwithin which to generalradiationof major structuralplansthat
assessrelationshipsof the speciesso recon- characterized the Cambrian history of the
structed or recognized.However, discoveries biosphere.
since 1976,when the f,"eat,semanuscriptwas
It would be an unjustifiedextrapolationfrom
essentiallycomplete,dictate some rearrange- Phakelodusto assignall 15 generaincluded by
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YJ,NOCONOJ :IHJ
TAXONOMY
h:e are the Disbi-ncation, augrber of genera
lcher superfamaoval
of a few
l'rte Conodonta
E of all other orx-a included in
hctive
bi- to
of
lparatuses
raslrate eled
fr-rous longituhderodontides,
I erceslral stock
, ppear first in
I range upward
prokihcation
lIirh skeletalapr- or septimemhents
or their
hh of the P polffs major con: and content to
@ of the Trea*o include in it
iodontidae) astte DistacodonE gmups (Pygo*lae ). which are
3a ard Ozarkoz I have found a
r &m ies of the
r rhe superfamifrlellacea of the
iludes some of
i aurhors in the
r. Prioniodinide
i.mbrate apparailbrm digyrate
:P positions. EleF tr'ith peglike
tsi of the order
x develop platl I recognize 34
ie rn r}tis order.
followed
drme
r!
Pol;gnatha: large order ineued
by sexi-
43
F6hraeus,L. E. (1983). Phylum Conodonta Pander, 1856 and nomenclatural priority. ,Sl,sl.
zool. 32(4), 455-459.
Hadding, A. (1913). Undre dicellograptusskiffern
i Skanejdmte nagra liirmed ekvivalenta bildningar.Lunds Univ. Arsskr.,Avd. 2 9(15), l-90.
Harley, J. (1861).On the Ludlowbone-bed and its
crustaceanremains.Q. J. Geol.Soc.London 17,
542-552.
Hass,W. H. (1962).Conodonts.Pp. W3-W69. In
Treatiseon InvertebrqtePqleonlology(ed. R. C.
Moore). Pt. W. Geol. Soc. America and Univ.
Kansas.
Hinde, G. J. (1879). On conodonts from the
Chazy and Cincinnati group ofthe Cambro-Silurian, and from the Hamilton and Geneseeshaledivisions ofthe Devonian in Canadaand
the United States.0. ./. Geol. Soc. London 35,
351-369.
Huckriede, R. (1958). Die Conodonten der mediterranen Trias uod ihr stratigaphischer Wert.
Paldont. 2.32, l4l-17 5.
Kohut, J. J. (1969). Determination, statistical
analysis,and interpretation of recurrent conodont groups in Middle and Upper Ordovician
strata of the Cincinnati Region (Ohio, Kentucky, and Indiana). J. PaleonL 43(2),392-412.
Lindstrom, M. (1970).A supragenerictaxonomy
References
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Am.
Paleont.
in
and
Indiana.
alents Ohio
Rhodes, F.H.T. (1951). A classificationof Penns0(229),21 t-441.
sylvanian conodont assemblages.J. Paleont.
Branson, C. C. (1957). Comment on the Moore/
26,886-90r.
Sylvester-Bradley "Parataxa PlaL" Bull. Zoo l.
Schmidt. H. (1934). Conodonten-Funde in urNomencl. 15, 169.
sprunglichemZusammenhang.Pqliiont. Z. 16,
Branson, E. B. (1938). Stratigaphy and paleon76-85.
tology ofthe Lower Mississippianof Missouri,
(1950). Nachtragezur Deutung der ConoPanl. Univ. Missouri Studiesl3(3), l-208.
Decheniana104, I l-19.
donteL
(1933).
Conodont
studand
Mehl,
M.
G.
-,
ie^snumber l. Univ. Missouri Studies8(l), 5- Scott, H. W. (1934).The zoologicalrelationships
of the conodonts.J. Paleont.8, 448-455.
(1936). Geological afrnities and taxon- Smith, J. (1907).On the occurrenceofconodonts
-,
in the Arenig-Llandeilo formations of the
omy ofconodonts, Geol, Soc.Am. Proc.Abstr.
SouthernUplands ofScotland. Truns. Glasgow
1935, 436.
Nqt. Hist. Soc.,n. s.7(3),235-252.
pls.
(1944).
pp.
235-246,
Conodonts,
93,
-,
94. ln Index Fossilsof North America (ed..H. Stoll, N. R., Chm., Dollfus, R. Ph., Forest, J.,
Riley, N. D., Sabrosky,C. W., Wright, C. W.,
W. Shimer and R. R. Shrock). Wiley, New
and Melville, R. V., Secretary(1964). IntemaYork.
tional code of zoologicalnomenclatureadopted
Clark, D. L., Sweet, w. C., Bergstriim, S. M.,
by the XV International Congressof Zoology.
Klapper, G., Austin, R. L., Rhodes, F. H. T.,
Int. Trust Zool. Nomencl.,Londoq 176 pp.
Miiller, K. J., Ziegler, W., Lindstraim, M.,
Miller, J. F., and Harris, A. G. (1981). Cono- Sweet, W. C. (1970). Uppermost Permian and
Lower Triassicconodontsofthe Salt Rangeand
d,onta. h Treatise on Invertebrcte Pqleontology
Trans-Indus ranges,west Pakistan, pp. 207(ed. R. A. Robison), Pt. w, Suppl. 2. Geol. Soc.
275.h Stratigraphicboundaryproblems,PermAmerica and Univ. Kansas,202 pp.
ian and Triassicof lVestPakistan (ed. B. KumEichenberg, W. (1930). Conodonten aus dem
mel and C. Teichert). Univ. Kansas,Dept. GeCulm des Harzes.Paldonl. Z.12, l'77-182.
ology, Spec.Publ. 4.
Fager, E. W., and Mccowan, J. A,. (1963). Zooplankton speciesgroups in the North Pacific. Ulrich, E. O., and Bassler,R. S. (1926). A classification ofthe toothlike fossils,conodonts,with
Science 140(3566), 453- 460.
or septimembrate apparatusesu,ith carminate
and angulate pectiniform elements (or their
platformed analogues)in P positions. The ozarkodinides, which appear to have been the most
diverse of all the conodonts, did not appear
until late in the Ordovician and persisted into
only the earliest part ofthe Triassic. At present,
this is the largest conodont order, with 55 genera distributed among 12 families.
Chapter 5 is devoted to a description of the
principal components of each of these conodont orders and to a discussion of the distnbution and geologic history of these groups. A
synopsis of the classification followed here is
also included as Appendix A. Apparatuses of
typical speciesof many conodont generaare illustrated at appropriate places in Chapter 5,
and illustrated discussionsof additional species
are to be found in the Catalogue of Conodonts
(Zieglel 1973, 1975, 197'1,198 1), of which four
volumes have been issued to date,
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dnorS-sselas roJ
posn euEu lsrg eql sI tluop
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^urJo
vt NoooNoJ aHJ
! Fa-nson'sCono] rbr a class-group
E fu tacl lhal priorllrsirp b). the ftleld4-*-e {Sroll, 1964).
L ie name ConolryjJm Conodonta
lJ- :9tl) is aDimI eEsl:r (ex order
trt Fiiraeus's protr
ro Pander,is
irt
of
-assificadon
t i..1]liJe
classifia name proL-ria
bhd-r (1930). AI'd-€rcup
names
E.earl!
op€rated
I b*re beennoted
I E:nenberg by 7
fu--._ place) of an
E\lacfarlane's
E :oq-ever, so it
EE o[ any of the
kiradon
of the
I:E
:O the suborproposed
\l.bl
ri
in 1944.
--=6
E-r s\nonyms of
t
:alegories for
:rcognized as
!F
a*
Conodonta.
5. THEMAJORCONODONTGROUPS
shallow basalcavities,which were assembled
by
evolvedmembersof most ordersinto elabIn Chapter6 I look into generalfeaturesin the
with
orate sexi-to septimembrateapparatuses
eyolutionaryhistory of the Conodonta.Before
ramiform elementsin S
complexly
denticulate
attention can profitably be paid to such airy
and M positions and a varied assortmentof
matters, however, some attempt must be made
pectiniform elementsin the two P positions.
to characterizethe major conodontgoups, ilThe stratigraphicrange of the Conodonti is
lustratetheir typical members,and point out
from Upper Cambrianto uppermostTriassic.
tbe featuresthat provide internalunity for each
Although the Cavidonti seem to have apgroupand thosethat indicatetheir relationsto
peareda bit beforethe Conodonti in the Iate
other major groups.
Cambrian,the latter diversifiedalmost exploAs notedat the endofchapter 4, my concept
sively and were clearly the dominant conof the Conodontais limited to the group disodonts throughoutthe history of the phylum.
cussedcollectively as the order ConodontoThere is currently no evidencethat the Conophorida in the Treatise(clark et al., l98l).
donti hs Teridontzs) are descendedfrom cavHere,bowever,that groupis elevatedgeatly in
idont (e.g., Proconodontus)ancestors,which
Cavtaxonomicstatusand treatedas2 classes,
might suggestthat the Conodontaare polyphyidonti and Conodonti. These are further diletic. The possibility remains,of course,that
vided into 7 orders:the Cavidontiinto Proconthe Cavidonti and Conodontiwerederivedinodontida and Belodellida;and the Conodonti
dependentlyfrom ancestorsin the group asinto Protopanderodontida,Panderodontida,
signed to the order Paraconodontidaby ZreaPrioniodontidd,Prioniodinida,and Ozarkodi/ise authors. At present,however, there are
nida.
reasonablestructuralgroundsfor beingskeptical aboutaffinitiesbetweenparaconodonts
and
5.2 CavidontiandConodonti
conodontsand, on thosegrounds,I haveomitted the Paraconodontidafrom this discussion
The conodontsI assignto the classCavidonti
(and from the Conodonta).Answersto all of
are distinguished by thin-walled, smooththesebasicquestionsaresurelylurking in asyet
surfaced elements,primarily coniform, that
undiscoveredor undescribedcollectionsfrom
formed unimembrateto quinquimembrateapparatuses,most of which apparently lacked Cambrian rocks.
structuresin either of the P positions.Cavidonts made their debut in the Late Cambrian
(Cavidonti)and
(as Proconodontus)and persisted(as Belodella 5.3 The Proconodontida
Its Fanilies(Fig.5.1)
and.Dvorakia) into the Devonian.
The Conodonti, establishedby Branson Speciesof Procozodontus(Fig. 5.2), founding
(1938) as a subclassof the vertebrateclass stock of the Proconodontidaeand the oldest
Piscesbut here regardedas a classof another conodonts known, built an apparently uniphylum, includesthe bulk of the Conodonta. membrate skeletal apparatus composed of
This largeand important stock apparently orig- relatively large, de€ply excavated,smooth-surinated in Late Cambrian forms assignedby faced, hyaline coniform elementsthat harre
Mifler (1980) ro Teridontus.Typically, these subsymmetricallyoval transversesectionsand
conodonts formed longitudinally striated ele- keelson the anteriorand/or posteriormargins.
ments,mostly with relatively short basesand The marginal keelsof some elementsare mi5.1 Inhoduction
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ue qlIA dsnJ oqt surofreqlo eqlJo lsgt s?ereq^\ u?uqueJ lselq eql ur per?edd" '(6961 'rellln)
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VJNOCIONOJ AHI
THE MAJORCONODONTGROUPS
Lc tanding, and
ba.-r Pander, are
: Enrll Cordylodr:crts (Fig. 5.2)
ih qhich oneeled a.ndgeniculate
k to the asymIr Lhepresumed
t! :re alsodistinEuses- but the
lt ercavated eleb s5on processes
E dolabrateelet posteriorpror srooth curYe,
tle cuspwith an
le denticulated
h slieletalcomE orher is a ram-
Polonodus
*E:dae.
Fryxello&&iodonridae.
F ryxellodontus
Proconodontus
Cambrooistodus
lapetognathus
Eoconodontus
Fig. 5,2. Elementstypical of speciesassignedto various generuof the Proconodontida.
:tE
!:
iform version of the geniculatecomponentof en with Proconodonlusin a singlefamily, alCambrooistodusapparatuses.Miller (1980) thoughthis would surelycomplicatediagnosis
suggeststhat Cambrooistodusand Cordylodus of the Proconodontidae!
developedindependentlyfrom Eoconodontus. The two known speciesof Iapetognathus
Ifso, it might be betterto includeall threegen- (Fig. 5.2) are representedin Early Ordovician
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uorlspurl
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pue eqlJoJ*prtuopoS d eql pot?erJ
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sod,qtueurolooqt Jo eajql 'eurle/(q,{llBcrs?q
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sluopouoJluopr^"c paJu"^p€aJorxaqlJo ouo
pu? (I'S '3rd) e"prluopoflexACeqtJo requeur
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snluopouo)otd
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'Je^e,{roq'uorlzu?Alprlu?lsqnsltqlqxa s?srr8
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snr0 pu?
snqpuSopdDlJo ed,4lalSuls
tuourele
e,{Iuo sezruSooeJ
(9961 'uofuraelg pue 'tur
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sI sluouele snqlDuSoladol
Jo ssecoJdpel?lnrq
-uap eql 'sr l?ql 'uoqelueuo ur reJrp
^llue
-J?dd?lnq snlereddz snpo{pto) oql Jo sluau
-odruor et?rq€lopalqncueA eqt lseEAnslBrll
Euoruele rrrJoJrrrreJ,{qeleJls (uBrcopeueJl)
VJNOCONOC IIHJ,
THE MAJORCONODONTGROUPS
I b Lindstritm
gecimen, which
forographs sugrilbrm element
y inegularly dislr rbrm equally
ibsrrarions of a
E Aom the Early
na.rm
Mitler's
E
least 3 speof two
-pnsed coniform
tl.ared
ry and bear surk of concentric
kributed ridges
c develops four
' ad two posteEnmc
nodose
: other more ir! bsically trirary lobes may dii
elements of
cridBnce as yet
&r eloped more
,lrn none of the
. bsed on very
f, rccrntly been
tu
in11983),
brictan species
paruses of sped !-oniform elerbe homologous
theelobed eler of 'r'hich are
lmenr\ with ininel)
denticurrgins. One of
E described as
Fmerricat; the
r BeEstrdm (in
lL One of the
lenrs in the apcicall! trilobate
b rhe anterior
ble element of
r sp{-a$'ling elediate and posI simpler than)
;rrauses ofPoryrarn idal quad-
riramate and tertiopedateelementsof the Pygodus apparutus form a partial symmetrytransition series,they are reasonablyassigned
to S positions.The sprawlingcharacterof the
other two elementsis like that associatedin
other conodont specieswith P-position elements,hencethey are assignedto thosepositions.That alsosuggests,
ofcourse,that the two
known elementsin the apparatuses
of Polonodas speciesmight atsohave beenoccupantsof
P positions,although obvious elaborationof
the apparatusin the evolutionofPygodasfrom
Polonodusmay merelyhayemaderoom for additional positions.
coniform elements,but the entire apparatus
has not yet been reconstructed.Walliserodus
species(Fig. 5.3) had quinquimembrateapparatusescomposedof keeledand costateconiform elements,one a symmetricatform with
anteriorand posteriorkeelsand interpretedas
an occupantof the Sd position.Dvorakia (Fig.
5.3) differs from LValliseroduspintarily in that
apparaluses
of its two known speciesapparently lackedelementsin the Sd position.Belodella, name giver of the family (and the order)
(Fig. 5.3), includesspecieswhoseskeletalapparatusesare similar to thoseof Walliserodus
but are distinguishedby S elementswith finely
denticulatedposteriormaryins.
Speciesof Ansella(Fig. 5.3),principal genus
5.4 The BelodellidaandIts Families
of the Ansellidae,built quinquimembrateskel(Fig.s.l)
etal apparatuseswith a geniculateconiform
In Fig. 5.I (and in the classificationoutlinedin elementin the M position and an S seriesof
Appendix A), I include three closely related elongate,deeplyexcavated,surficiallysmooth
groupsof conodonts(Ansellidae,Dapsilodon- rarniform elementswith adenticulateor irregtidae, and Belodellidae)in a newly established ularly serrateposterior margins.Elementsin
order of the Cavidonti for which I proposethe the Sa position have three keellikecostaeand
nameBelodellida.All the conodontsassernbled are triangular in cross section; Sb elements
in this new order formed quadri- to quinqui- have sharpanteriorand serrateposteriormarmembrate skeletal apparatusescomposedof gins and a planoconvextransversesection;Sc
typically thin-walled, smooth-surfacedconi- elementsare like thosein the Sb positionin
form elementswith \'ery deep basal cavities. the apparatusesof most known speciesbut
Thoseelementscommonly havedistinct ante- areofbiconvex crosssectionand,in a majority
rior, posterior,and lateral keelsor costaethat of species,have no posterior denticles or
developfine, needlelikeserrationor denticula- serTatlons.
tion in the apparatuses
of severalspecies.
The apparatusof Hamarod.us(Fig. 5.3) apThe central stock of the Belodellidais the parentlyhad a geniculateconiform M element
Belodellidae.Treatisearolhorsassigned,S/a1o-and a symmetry-transitionseriesthat included
dus, Belodella, Coelocerodontus,and Walliser- a quadriramateelement with a few anterior
odus lo this family. I add Dvorakia (Klapper and lateraldenticlesin the Sd position,and latand Barick, 1983),which was describedand erally adenticulatealate,tertiopedate,and donamed after the Trcatisewas published.The labrateelementsin the Sa,Sb,and Scpositions,
better-known belodellids formed quadri- to respectively. Typical of the Hamarodus appaquinquimembrate apparatuses of slender, ratus,however,are a pair of deeplyexcavated,
deeply excavated,smooth-surfaced,basically laterally compressedelements with serrated
coniform elements,ornamentedby longitudi- postero-and antero-basalmargins.Evidently
nal keelsand costae.The latterbearminute ser- theseelementsoccupiedP positions.Affinities
rations in severalspeciesbut are undenticu- of Hamarodusarc anythingbut clear.Elements
latedin others.
assignedto the S positionsare surely not exThe oldestbelodellidsare includedin Slolo- actly like thosein comparablepositionsin Andus (Fi$.5.3), which Lindstr6m (1955)erected sella, which also lacks elementsin the P posifor conodontswith deeplyexcavatedconiform tions. But Ansells alsolacksfeaturesthat would
elementswith threeor four asymmetricallydis- ally it very obviously with any other major
posed longitudinal keels or coslae. Coelocero- group of conodonts,including the Periodontidontus (Fig. 5.3), which may not be distinct dae(Prioniodontida),whereit wasassignedin
from Stolodus,includessymmetricallycostate the Treatise(Clark et al., l98l). Thus I follow
polecrplrl iFeels $ snpoltsdDe q]lt
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VJNOOONOS AHJ
THE MAJORCONODONTGROUPS
tinctive pattem of chevron-shapedridges adjacent to the anterior margin. At present,BeJselodusis known only from Ordovician rocks.
Dapsilodusis also representedin the Ordovician, but it is probablybest known and most
in Silurian strata.
typically represented
5.5 OrderProtopanderodontida,New
d
This division of the Conodontaincludesmost
of the conodontsthat built uni- to multimemcomposedof longibrate skeletalapparatuses
tudinally striated, laterally unfunowed coniform elements. Expresslyexcluded are the
panderodontideconodonts,a goup that built
apparatusesof coniform and rastrate elements
with deeplateralfurrows;and the oistodontids,
JI
whoseapparatuses
include pastinateconiform
(or "acodontiform") elementsin an array of
coniform typesthat is otherwisetypical of the
Prioniodontida.
With the exceptionsnoted,the Protopanderodontida is the major conodontgroup identified in the Treatiseas the superfamilyDistacodontacea,which was based on the family
Distacodontidaeof Bassler(1925). Unfortunately, no one knows much about Distacodus
Pander,1856,whosetypesarepresumablylost,
or about the Distacodontidae,basedon it by
Bassler(1925). Consequently,I suggestthat
Distacodusberegarded.asa nomendubium ar.d.
that it not be usedas the basisfor any supragenerictaxonomicunits. The name Protopanderodontida, although a mouthful, is taken
Fi9.5,4, The Protopanderodontidaand Panderodontida.The families Protopanderodontidae,Clavohamulidae,
Acanthodontidae,and Drepanoistodontidaemake up the Protopanderodontida.The Panderodontidaconsistsof
the singlefamily Panderodontidae.
z
9
o
G
thca
&Components
Iosed
by Sb, Sc,
r elementmay be
P R OT OPAN
i
\*
r oblique orna;ior margrn.
B 5.3) formed
rirh nongeniculmenrs in appaFg 5.3) are gens areknown
I a relationship
raed bv the dis-
s
\d
z
@
D R E P A N OIS TODO NTI DAE
C L A VOH AM
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VINOCONO]:IHJ
THE MAJORCONODONTGROUPS
n*-ldontidae.
h<nrs like those
I a sharp, narrow
Srr of. the distal
rlrf,
6 tas redefinedby
@- j.5) includes
tlh<€S were COmI elements of cirs
These
-ction. on the
dsntadon
h oi maximum
fum a few low,
&E. -\ currently
rmediate be'f]L.a;
iodusi I am
-t
r rerir separation
, l9-J has a bir ot- deeply excat Erth relatively
I rl}e is short and
tho apically; the
rd more slender
rilar morphologlseudooneotodus
Eilie apical denrdiagnostic ofthe
lower Middle
rred
by Stouge
zrs ir Upper OrErd inlo the Delus separatedby
a considerable stratigraphic interval from the components of multielement Tropodus
Oneotodus,its closest morphologic relative, comptus,tbe type species,as form speciesof
and this suggests
that ancestryand possiblyfa- Paltodus a\d. Scolopodus,which suggeststhat
may be there is still uncertainty a'bout Tropodw in
milial assignmentof Pseudooneotodus
elsewhere.
Lower Ordoviciancircles.
(Fig. 5.5)built biSpeciesof Semiacontiodus
representedin earliestOrSemiacontiodus,
membrateapparatuses.
One elementtype is bi- dovician rocks by a number of speciesprevilaterally symmetrical,anteroposteriorlycom- ously assignedto I contiodus,wasprobablyalso
pressed,and has a posterior indentation or ancestralto the long-rangingEarly Ordovician
costaflankedby posterolateral
costae;the other stock named Glyptoconusby Kennedy(1980).
elementis asymmetricand hasa costaon only The only speciesof Glyptoconus(Fig. 5.5) that
one side.Sometimeearly in the Middle Ordo- has been adequatelydescribed,G. quadrapli:"ician, Semiacontiadzrgaverise to ,S/auferella callas(Bransonand Mehl, 1933),has a multi(Fig. 5.5), whose speciesformed a bi- or tri- membrateapparatusof hyaline coniform elemembrateapparatusthat differed from those ments that have only slightly expandedbases
formed by Semiacontiodusspeciesin that the and slightlyrecurvedcuspsthat arequadratein
basal part of the finlike lateral costaeis dis- crosssectionand bear prominent longitudinal
tinctly notched. Dzik (1983) suggeststhat grooveson the lateraland posteriorfaces.
Semiacontioduswas also ancestralin the MidA particularlyimportant eventin the history
dle Ordovician to Scabbardella(Fig. 5.5), of the Protopanderodontidae
wasdevelopment
whose only known speciesformed a bimen- in the earliest Ordovician of the short-lived
brate apparatus of strongly compressed ele- stocknamed Utahconusby Miller (1980).Spements, one type of which is essentiallysyrn- ctesof Utahconus(Fig. 5.5) formed a bimemmelrical and hasanlerior and posteriorcostael brate skeletalapparatusof albid elernents,one
the other is asymmetricand more broadlycos- ofwhich is a primitive pastinate(or acodontitate on one sidethan the other.
form) coniform element. Utahconus probably
Shortly after Sezlacontiodus made its debtJt wasancestralto RossodrJ,the oldestdescribed
in the very early Ordovician, it apparently memberofthe Prioniodontida.Utahconusmay
spawneda stock of protopanderodontide
con- alsohavebeenthe ancestorof the Acanthodonodonts characterizedby a multimembrateap- tidae,anotherof the protopanderodontide
famparatusof albid, costate,and noncostateconi- ilies. ParutahconusLanding (in Fortey et al.,
form elements.Landing, Bames,and Stevens 1982) is similar, but its elementsmay bear
(1986)haverecentlycoinedthe genericname small nodeson the baseand along the lateral
Variabiloconus
for the bestknown of thesecon- costa.
odonls,V. basslui (Furnish,1938)(Fig. 5.5).In
Fig. 5.4I suggest
that Variabiloconus
may have
Lindstom, 1970
(Fig. 5.5),a 5.5.2 Family Clavohamulidae
beenancestralto Protopanderodus
long-rangingstock of Ordovician conodonts This family was establishedfor a small group
with a bi- to trimembrate skeletalapparatus of bizarreprotopanderodontide
conodontsthat
composedof albid elementswith prominently formed unimembrate skeletalapparatusesof
sharp-edgedposterior and posterolateralcos- albid elementsthat are surficially granulose,
tae.In the very latestphasein the historyofthe nodose,or spinose.Speciesof Hirsutodontus
Protopanderodusstock,the upper margin ofthe (Fig. 5.6) built apparatusesof Teridontus-like
basewasextendedposteriorlyand cameto sup- coniform elementsthat developedspines or
port a singlelargedenticle.
nodesat the base.The basalpart ofcomparable
Speciesof TropodusKennedy (1980) (Fig. elementsof Cldvohamulus(Fig. 5.6),however,
5.5), like those of Protopanderodus, formed spread laterally to form a hemispherical
multimembrate apparalusesof deeply exca- mound with a nodoseupper surface.Elements
(Fig. 5.6),on the otherhand,
vated symmetricaland asymmetricalconiform of Serratognathus
elements,all of which bear three to firre prom- are anteroposteriorlycompressed,bilaterally
inent keellikecostae.Repetski(1982)describes symmetricalstructureswith a knifelike lower
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Jouelsod pua louetuz dj?qs s€q,{luoruuloc
pu? 'lmulelxu s fll?lelqrq ,pesseJduoc sr
lusuele auo 'eule^q ro prql"
elE leql
sluorrJeleurJoJruoceleFcrueSuou
^plEa,n
Jo s5snl?led
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uoqsJo s,rol
^lasolr
snoJsrunuqlr^\ ar?J Jouelue
u? pu? ulSJeur
'oEprlnureqo^elJeql Jo
Eroue8ol peu8rssE
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nilbrm elements
he- One element
4rmmetncal, and
k and posterior
E bur the cusp is
D rie base,which
Ii5 Lindstrtim es'ntdodus for elei i !-onvenientto
The type
k!el.
may have
-i..m.
nrus of a species
d
henceis not
rd this famity.
F.d here to the
t one another in
m:lodu species,
hta row of nodes
beas an anterior
d r}rebilaterally
ilalin a speciesis
notch
Tle family is
-srincdve
-.
*t
Fdhraeus
L<- Paroistodus,
Ee of an impornide conodonts
! Lsad the name
L-{s notedearlier
Edrs is best rei thus is not suitxic categories.
E apparently the
trnridae, has a
rirh a complete
.sinctive genic'X position,and
qmmencally bi
B in the P posi'tr basalmargin
b ilaftened, or
Eacter suggests
mmber of the
dEstor.
frg- 5.8) had a
ronly described
rld probably be
THE MAJOR CONODONT GROUPS
55
of recessivebasal margin at the anterobasal
corner.Nongeniculateelementsthat form the
symmetry-transitionseriesin the apparatusof
Paroistodus speciestend to develop bases
whose upper margins are drawn out posteriorly. Dzik (1983) and Stouge(1984) have
bolh suggested
that this processbecamedenticulatedin the youngestspeciesof the genus,
P. horridus(Barnesand Poplawski,1973).
5.6 The Panderodontida
Patotsrodus
Fig. 5,8, Elementsand apparatusestypical of species
assignedto generaof the Drepanoistodontidae.
consideredquinqui- or evenseximembrate.
An
essentiallyerect, nongeniculateelement with
equally concavesidesand sharp anterior and
posteriormarginsoccupiesthe Sa position.Sb
and Sc positionsin the transitionseriesare
filled by regularlyrecurvednongeniculateconiform elements, which are flatter on one side
than the other and commonlyhavebasalmargins that are diferent in conformation.M elements are geniculateconiform elements,commonly with long cuspsand much shorterbases.
P positions were evidently occupiedby subsymmetricalnongeniculateconiform elements
with an acutelyangularanterobasalcornerand
sidesthat are almost equally convex. Certain
speciesof Drepanoistodusattained an essentially cosmopolitandistribution in the Ordovician, but the genusbecameextinct at the end
ofthat period.
Paroistodus(Fig. 5.8) specieshad a bi- to
quadrimembrateapparatus that included a
symmetry-transitionseriesof laterally costate
nongeniculateconiform elementsand a geniculateconiform elementin ihe M position that
characterislicallydevelopsa conspicuouszone
Speciesassignedto this distinctive division of
the Conodontaare distinguishedby tri- to septimembrate skeletalapparatusescomposedof
longitudinallystriated,taterallyfurrowed coniform or rastrateelements.The latter are basically coniform typesbut have a few denticles
alongthe posteriormargin ofthe cusp.In only
a few apparatuses(e.g.,that of Plegagnathus)
are denticulatedparts of any elementsdrawn
out into distinct processes.
In my opinion the major diagnosticfeature
ofthe Panderodontidais the distinctive longitudinal fissureor furrow that is developedon
one or both sidesof everyelementofthe skeletal apparatus,and I have excludedfrom the
order all ofthe conodontswhoseelementslack
indication of this feature. This means, of
course,that my conceptofthe Panderodontida
is considerablynarrowerthan that adoptedin
the Treatise.
In the narrow senseadoptedhere,the PanParapanderderodontidaincludesPanderodus,
odus,PseudobeIodina, BeIodina,Culumbodina,
Plegagnathus, Parabelodina, and Neopanderodus. Pseudopanderodus(l-anding, 1979) is excludedbecausethe ostensiblylatestCambrian
coniform elementson which it wasbasedhave
subsequentlybeen determinedto be representatives of Panderodus that were somehow
mixed into collectionsfrom much older rocks.
The oldest panderodontidespeciesknown
had apparatusescomposedof furrowed coniform elements,and it appearslikely that they
were ancestral to species of genera such as
Pseudobelodina,Belodina, Culumbodina, and
Plegagnathus,whoseskeletal apparatuseswere
formed entirely of rastrate elements.Consequently,at first blush,it would seemfeasibleto
assignpanderodontideconodontswith appa-
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arppanttusesof Para!lsed clusterofParofPodolia. From
--
,4, ro Semiacond- most recently,
f Parapanderodus
l Tbat assignment
E mination of the
b in availablecoltar it is surely a
lu ir sharessome
lrfies of ParapanI a home for it as
EE rhat Para.panrs and Poplawski,
Fr q?s at least
r longirudinally
ar sith a broad
.drher side by an
*r.
subsymmetrr furrow on only
ltr! and an asymI s conspicuously
r tb€ anterior-posldenent, which is
Itpears to lack a
Eradons, but it
Ergitr of the mid3 ro note that that
r of this element;
imargin in the po7rrsible in figures
Ege- and Ethingtognderodus arc
t Middle Ordovi
cse specresappar-
THE MAJORCONODONTGROUPS
ently containedonly two typesof slenderconiform elements, both of which exhibit a
"panderodontfurrow" (or a pair of them) on
the posteriorface.The basalcavity oftheseelements seemsto be somewhatdeeperthan that
in elementsof P. asymmelricusblot m\ch shallowerthan cavitiesin elementsofyoungerpanderodontidconodonts.
Panderodus(Fig. 5.9) made its debut a bit
later in the early Middle Ordovician (Llanvirnian or Whiterockian).Accordingto Ldfgren
(1978),the apparatusofthe earliestspecies,P
sulcatus(Fahtaeus),had a birnembrateapparatusofslender(or "graciliform") and laterally
compressed(or "compressiform") coniform
elementsthat are longitudinally striated and
havebasalcavitiesabout one third ofthe total
elementin length.Elementsof both typesappear from Liifgren'sillustrationsto be slightly
bowed,and the distinctivepanderodontfurrow
is situatedon the outer, or convex,side.
Later Ordovician speciesof Panderoduswere
determinedfrom large collectionsof discrete
elementsto have had quinquimembrateskele(Sweet,1979),and this is also
tal apparatuses
the casein fused clusters representingPanderodusunicostatus(Bransonand Mehl) from the
Early Silurian of Podolia,which were recently
describedin detailby Dzik and Drygant(1986).
Although certainof the elementsin the apparatus I reconstructedin 1979are nearly bilaterally symmetrical,they are furrowed on only
one side and were describedonly as "similiform." Jeppsson(1983)reportsthat there are
truly symmetricalforms in his collectionsof
Panderoduselementsfrom the Silurian of Gotland, and a few have also beendiscoveredby
Britt Leatham, one of my students,in the large
collectionson which I basedmy 1979reconstruction. Thus it is appropriateto conclude
that the apparatusof fully evolved speciesof
Panderodus was probably seximembrate although,as Dzik and Dryeant (1986)note, distinctionsbetweenelementsforming the S series
are commonlydifficult to make.
Panderodusrangedthrough the Silurian into
the Devonian, but information on the species
representedis sketchy.Indeed,specimensare
illustratedin only a few reportson Devonian
faunasand Silurian forms have receivedless
than adequatetaxonomictreatment.
The youngestmembersof the Panderodontidae are late Early and Middle Devonianspecies ol Neopanderodus,which apparently had
apparatuses
like thoseof better-known
species
of Panderodusbut composedof furro\ped coniform elementswhosesidesaremarkedby longitudinal striaethat are more prominent than
thoseof Panderoduselements.
The oldestrastrateelementsknown are from
early Middle Ordovician rocks of about the
sameageas the onesfrom which the earliest
Panderoduselements haye been collected.
Opinions vary as to the genusrepresentedby
theseelements.
but I regardthemasspecimens
of Belodina.Later, better-knownspeciesof ,BeIodina (Fig. 5.10)formed trimembrateapparatusescharacterizedby three differenttypes of
rastrateelements,which are separatedprimarily by differences
in their radii ofcurvatureand
the number of denticleson the posteriormargin. One elementlacksposteriordenticlesand
is formally coniform.
In 1979| describeda sequenceof speciesof
Belodina and noted that in younger and
youngerOrdovician stratadenticulaterastrate
elementsbecome more numerous and coniform ones proportionatelyless so. However,
one Late Ordovician species, B. calciprominens,is known only from largeconiform rastrate elementsof the sort onceincludedin the
form genusEobelodina.Although specimens
assignabletoBelodiru are known from rocks of
latest Ordovician age, none has yet been recoveredfrom the Silurian.
Pszudobelodina(Fig.
5.10)hasa quadrimembrate skeletalapparatuscomposedof deeply
excavated, laterally furrowed, posteriorly
"heeled," anterolaterallycostaterastrate elements.Threeofthese elementtypesare bowed
to the unfurrowedside and form a symmetrytransition series;the fourth, which may have
occupiedeitherthe M or oneofthe P positions,
is straight or is bowed slightly toward the furrowed side. The oldest Pseudobelodinaknown
to me is an as yet undescribedspeciesfrom
Middle Ordovician (Blackriveran) strata in
North America.Although elementsof this unnamedspeciesareall denticulalerastrateforms
and thus morphologicallysimilar to comparable elementsin the apparatusof Belodina species,thereis no coniforrnrastrate(or "eobelo-
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feru qcrq,rr 'oulpopg uro{ lou pue snpoap
-u0d wo$ ,([poJrp pedole^ep Durpopqopnasd
leql
lr {uql I ,,{JluenbesuoJ.slJor eruEs
^le{rT
eql ur
peluoseJdeJ eff qcrq/h ,ossatdu,toJ .B
pue stsuatoluour outpopg
Jo esorllpue sersods '
aurpolaqopnasd srrll Jo slueruele ueo^ueq uJoJ
uJ uorl"patJolur sr eleql qJrq,{\ ur suorpelloJ
uaos lou e ?q I pue ,luourele (,,r.uJoJrurp
'oEprtuopolapuedeqtJo EJouoB
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luaqroJo-olDrlsEx .0I.S ,i|!J
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r^ opro'I I'S'3!d
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r0r,$ 'spnuop
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Dlleqs ur ,(lrsje^
VJNOCONOJ AHJ
89
THE MAJORCONODONTGROUPS
versityin shallow,tropical,or subtropicalseas. 5.7 The Pdoniodontida
Dzik, 1976
Panderodusitself has a somewhatmore cosmopolitan distribution,but it seemslargelyto This major division of the Conodonta (Fig.
have been replaced in higherlatitude Ordovi- 5.1l) includesconodontsin which one or both
cian faunas by Drepanoistodus and, farther ofthe P positionsin the fundamentallysexi-or
poleward, 14alliserodus. Specialized pandero- septimembratecephalicapparatusare occupied
dontids, with rastrate-elementapparatuses, by pastinateconiform or pectiniformelements
were orimarilv low-latitude forms.
or their platformedequivalents.In a few spe-
Fig.5.ll.
Ordovician and earliestSilurian families and generaofthe Prioniodontida.Later Silurian and Devonian
reDresentatives
of the order are shown in Fis. 5.17.
s
e
a
t
Parubelodina
to
bergsand
and i-ndepenies all have
apparatuses.
I interpret it,
e
I
l
I
I
?
POL
fl\
il\
e
I
i
3
I
I
\
of heeled,
multicostate
to$ bowed tothat reptave denticles
including
mostof
ttrey may be
s
I
a
i
PERIODONTIDAE
II
AA
a
!
il
J
a
Ip
l
the
of Culumbodishedinforlit €arlierin the
America,
early part (EdContemporary
noi
.
which must
group.
panderodonts
seem to have
and di
il\
artouDnl' '(snll
) luauelo uJoJ
t9 etl) snqtDu
P m}?r?dd? Ie1
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Fd-. puE slueu
4 raqlo eql uo)
Dl o.Yrlsll oCueq
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^lInJ
rPossou 'pelues
!.oq lsnf eur ol
6 snqpuSouDnl'
snp olslo' tL6 l
'lolue^\
tr6I
ssa\f, snponqd
I '9581'ropued
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itsnlEJeddeqcns
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)e Jo
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flJeg pu? u?rcr^opJouae,{rleq
dqsuouElal eql
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VINOCONOJ
3HJ,
THE MAJORCONODONTCROUPS
6l
Eumented Lower
4pear to be anaive stocks.The
' probably seYeral
'r*J'that Sandbde werederived
lalus and probEus. As noted in
rDdae, however,
ncian and Early
rd tater Silurian
fu alything but
bs suggesteda
alstrom, 1970
rodonts with a
ieletal apparatus
Protopttoniodut
Fig. 5,13.
*rudae.
Oisaodus
Elements and apparatuses typical of species included in three genera ofthe Oistodontidae.
Drawings
of coniform elements,of v/hich one (P) is pastinate (or "acodontiform"), one (M) is geniculate (or "oistodontiforrn"),and the rest form a
symmetry-transition(or S) series,which,when
completelydifferentiated,gradesfrom an alate
("acontiodontiform") elementin the Sa position by way ofa digyrate(or "paltodontiform")
laterallyunornaSb elementto a compressed,
mented nongeniculate ("distacodontiform")
element in the Sc position. Conodontswith
such apparatuseshave been included in Rossodrs Repetskiand Ethington, 1983,Acodus
Pander,1856,TripodusBradshaw,1969,Diaphorodus Kennedy, 1980, OelandodusYan
Wamel, 1974, Protoprioniodus McTavish,
1973, OistodusPander, 1856, and probably
JuanognathusSerpagli,1974,but it is not clear
to me just how many generaare really repre(Fig. 5.12) may not have a
sented.Rossod?rs
fully developed symmetry-transition series,
henceits two known speciesseemprimitive.
(On the other hand.the "acodontiform" P elements and "paltodontiform" Sb elementsof
Rossodasmay be so similar that they havenot
beendistinguished).Furthermore,if the skeletal apparatusof the tlpical speciesof Juanognathus (Fig.5.12) includesa geniculateconiform element (Serpagliwas not sure about
this), Juanoqnathusis probably a senior syn-
onym of Rossodzs,as noted by Repetskiand
Ethington.
Tripodus (Fig. 5.12) and Diaphorodus are
basedon different, but probably congeneric.
species;hence I regard them as synonyms.
Therewill probablyalwaysbe uncertaintyasto
how Acodus (or many of Pander's genera)
shouldbe interpretedin a multielementsense,
so I recommendthat Acodusbe consignedto
the list of nomina dubia and that we go on to
more important things.
Oistodus,Protoprioniodus, and Oelandodus,
all shown in Fig. 5.13, include specieswhose
skeletalapparatusesare composedentirely of
geniculateconiform elements,which neverthelessexhibit an arrayof forms closelysimilar to
that in the apparatusofspeciesofRossadzsand
Tripodus. ln lhe Treatise Bergstrdm expressed
the opinion that furtherwork might showthese
threegenerato be synonyms,and I can seeno
very goodreason(asidefrom the geniculatenature of their coniform skeletalelements)to
refer them to a family separatefrom Rossodzs
and.Tripodw. Consequently,in the schematic
classificationin Appendix A, I group all these
generain the family Oistodontidae(which has
priority over Juanognathidae),
btt rcfer Oistodus, Protoprioniodus, and Oelandodus to a
subfamilv Oistodontinae: Rossodus and Tri-
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V.JNOCONO] AHJ
l'zpodas. It is also
&znae represents
lhent from either
bcauseit is sepa'frovician from
I gpical HistiodEi describedonly
q,dthough he now
iulate coniform
?B2ssleL1925
t just above the
'ls debutare con*ehral apparatus
r are occupied by
Enate elements
res: rhe M posiictrlate coniform
irnally a bipenit an elongate,
r pnocess;and S
Eition seriesthat
&re. alate, and
clear that
t-EDs
Prioniodus PanI fnpodus, fiorn
, in a somewhat
rarus in which
ttr out into denfr rhereis alsoa
Amo.phognathus
Fig. 5.15. Elementsand apparatusestypical ofspeciesassignedto generaofrhe Balognathidae.
greaterdegreeof morphologicditrerentiationin
the S series.
Prioniodus(Frg.5.14)developedphylogenetically through a successionof similar, biostratigraphicallyusefulspeciesin the Earlyand
earlyMiddle Ordovician,and in rocksa couple
of zones above the one in which it first appeared,spawneda new lineage,assignednow
to Oepikodus(Fig.5.l4). In speciesofthe latter,
the skeletal apparatus consists of Prionioduslike pectiniform elements;a geniculateconiform M element;and a symmetry-transition
series of quadriramateelementslike the one in
the Sd position of Prioniodus.
groups of elementsthat representa phylogenetic seriesofconodont speciesreferredto Baltoniodus(Ftg.5.l5).Species
of Baltoniodus
arc
distinguishedfrom thoseof Prioniodusby morphologicallydifferentpastinateelementsin the
two P positionsof their skeletalapparatuses.
Although this is regardedas of no more than
subgeneric
significance
by severalinvestigators,
others have pointed out that Baltoniodusappeared a bit later than Prioniodus and also
probablyevolveddirectly from a youngerspecies of Tripodtts and not from Prioniodus.
However tbis may be, Baltoniodusis the universally acknowledgedancestorof Amorphognathus, which appearedin a somewhat later
part of the Early Ordovician and ranged,
Hass,1959
5.7.3 Family Balognathidqe
lhrougha succession
of distincivespecjes,
to
Beginning in Lower Ordovician rocks just the end of that period.
above the onesin which Prioniodus is fitst recSpeciesof Amorphognathus(Fig. 5.15),perognizedand extendingupward into Upper Or- hapsthe best-knownand most widely distribdovician strata is a successionof recurrent uted membersof the Balognathidae(Balogna-
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pf,utaurur,(sE ,(1pre1elq 'purlsrp ,{ ?Jr
f?ql
tcedsns
-s?d
I
S
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eqlJo e^rpuEsrp$ueurelapeurJoJ -8oloqtuou
perdncco era,{ suo[rsod d
-l€ld oql "l
lrrg 'snqpuSol4rowv Jo asotfl o^r1 oql pu€ ^qstuauele (,,uuoJr1uopoloq,, Jo)
Icel
elquaseJ1?ql sluauale rrrJoJrursJ
epnlcul oslE alepodorUet e!?zrq eJoq uorlrsod I I oql q3rq,r
l"qt seldul?seurosu rnxxo snqpuSosapoqUJo ur 1nq 'snl?Jedde snpotuo og oql uI seceld
cusuep?J€qosluel'ueleel?urlsedeql're^e,{ioq J?IuIIs ur esoql 01 elqeJuduor slueluele rruoJ
'sluepnts Jno
rel?I polqurass€suorloelloc -ItrrEJpopnlcur suo[rsod S eql qcrq.r ur sesnl
^q
uI'stuoluele(..uuoJr1uopopque,,
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VINOOONOJ
AHJ
THE MAJOR CONODONT GROUPS
zll) symmetrical
rm") elements.In
b our students,
Ets characteristic
me samplesthat
s rhat resemble
h lack the platJthe latter.Thus
lnssibly even M)
u ar:rd.Rhodesoad rhat RhodesEbrate appara*ale
one we
rte genuswaseslea the closeregm.hus ar'd.Rhoh: 5.1I is even
rhose unimemniniscaphate eleE rom) thoseof
bded in the Balibstrated in Fig.
nhidae the mulir Ihiipfer (Fig.
d bl Bergstrdm
;mn1m of Norr( 1982).NeiE
dontina is welL
Er drscussion,alr rtan passinginrticb is known
lu *ere probably
t!- be an index to
t fruna recorded
hlctive typesof
interpreted by
G ( 1980)as two
ft. 5.15), include
rdendcuatedprofte pastinateelehed processes
in
d dolabrateand
r in M positions;
i:Ily intergradaI dolabraterami6ramidal shape.
[ic in rhe M pofrate"; even so,
rrus- Apparatus
65
stuctureis, in many respects,more like that of
Prioniodus than Baltoniodus species,but the
speciesmost proximate stratigraphically is an
unnamed representativeof Baltoniodus(Savageand Bassett,1985;Orchard, 1980).Consequently,in Fig. 5.11,Gamachignathus
(includ.ing Birl<sfeldiaOrchard, 1980)is included in
the Balognathidaeand shown tentativelyas a
descend.antof BaItoni odus.
5.7.4 Family lcriodellidae,new
I interpretlyiodella Rhodes,1953as the stem
of an important prioniodontide stock that
probably also incltdes Pedavis Klapper and
Philip, 197|, SannemanniaAl-R.awi,1977, and
SteptotaxisUyeno and Klapper, 1980.In the
Treatise all these genera were included with
Icriodus and.Pelekysgnathusin the Icriodontidae,primarily on the basisof obvious sirnilarities in the Pa elementsformedby their species.
Many of the other elementsof the apparatus
are different.however,and the differencesare
questionsasto the
sumcientto raisereasonable
relationship belween l%iodella and its probable derivatives and the primarily Devonian
gnathus complex. Thus lcriodIcriodus-Pelelqts
e//a and its kin are here set asideas a new family, the Icriodellidae,the relationshipsofwhich
ICRIODELLIDAE
are shownin Fig. 5.17.
DISTOMODON
TIDAE
Basically.the icriodellidshave a quinquimembrateskeletalapparatusin which the Pa
elementis pastinatein the apparatusofthe oldest speciesknown, pastiniscaphate
in thoseof
later forms, and stelliscaphate,
with four processes,in apparatusesof the youngestspecies Fig.5.17.Silurian
P oniodontida.
andDeyonian
known. The Pb element,in apparatuses
of the
Icriodella (Fig. 5.18) and Pedavis(Fig. 5.18)
speciesfor which it is known, is a pyramidal ate, wilh adenticulateor faintly serrateedges.
pastinateelementwith three sharp edgesthat In apparatuses of species of lcriodella (Figare adenticulateor only weakly denticulate.I 5.18)the occupantofthe Sapositionis a deeply
interpret the "coronellan" elementin the ap- excavated,pyramidal alate elementwith minparuttrsof Steptotaxrsspecies(Fig. 5.18)as an utely serratedlateral processes,whereaseleoccupantof the Pb position,primarily because mentsin otherS positionsarepyramidaltertioit is related morphologicaltyto the pastinate pedateunits that are closelysimilar to the Sa
coniform (or "acodinan")elementsassignedto element,but asymmetricwith respectto disthal position in apparalusesof various icrio- position of their lateral processes.
S positions
donts.Theapparatus
ofthe only knownspecies in the apparatusesof Pedavisspecies,however,
of Sannemanniais unknown,
are filled by elongate,prominently costate
M elementsoflcriodella, Pedavis,and prob- structuresthat apparenlly are nongeniculate
a.blySteptota-xisare bipennate or even pastin- coniform elementsin the Sb and Sc oositions
-/ a /
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lmuwsoL 7)
rxuala S /'Aorl
q Jql Jo luotu
DqSUUOJUI JO
u,Jo sasnl.Er?d
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JlqrssodJo
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JlsJrpur
I
LI
S 3ll ul
Jo slueurela rrJoJruoc ele{nrrue8uou'el?lsot"
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ol ueql peuftss?s?q(986l)Joqcsrg pu?'e?pp 'fiC) sercedsstxDtotdats uI 'uonrsod ES aql uI
-uopourolsrqeqlJo sebodssnou?Afq pourroJ sluelllele alErq?lop elelnJrluep,(lJouelsod lnq
VINOCONOJ 3HI
THE MAJORCONODONTGROUPS
'te Distomodonrsigned them to
nber of possible
mg the generaI
hiodellidae, Dis*b.e. Pedavis is
RlcriodellidaebeEtr elementsI asriions in appara. &.nnemanniais
of its only
-mro that formed
L
:Easoningis also
Irz'rol&rts to this
f its more charl-r assignedto Pel=- Non-Paelexiderably from
s of Pedavisarc
bidella.
ldements ofspeilto Distomodus
Rtorhoseformed
I lresumed occur are clearlydistEreric relationuodonts
haye
fuilarities in Pa
rhis case,the
lbn
of Pedavis
-
Rotundacodin6
Distomodus
Fig. 5.19. Elemenlsand appamtusesofspecies typical ofgenera assignedto the Distomodontidae.
ftom Distomodusor some other genusof the
Distomodontidae instead of from lcriodella.
To my mind, the prominentlycostateelements
that occupy S positions in apparatusesof Pedarls speciesareequallydifficult to derivefrom
either the Icriodella ot Distomodus stock and
may thus be regardedas the featuresthat distinguishPelavis.Thus, if one considersthe remaining elements(Pa, Pb, M), similarity is
greatest between Pedavis and lcriodella, and
that hasguidedme in assigningPelavls to the
Icriodellidae.
Broadheadand McComb (1983)arguefrom
comparisonofthe Pa elementsof Itte Silurian
Pedavisand thoseof latest Silurian-earliestDevonian Latericriodus woschmidti that the latter
may have developedfrom the former by neoteny.Althoughthat is an attractiveidea,non-P
elementsin the apparatusof L. woschmidti
(Fig. 5.20)differ substantiallyftom thosein apparatusesof Pedawsspeciesand, in the absence
of information on the ontogeneticdevelopment of the latter, it is difrcult for me to see
how S elementsof the supposedpaedomorph
(L- woschmidtl)might be relatedto earlierontogeneticstagesofancestorsin PedavtJ.Never-
theless,I indicate in Fig. 5.17 the possibility
thal Pedavis,whosefamilial assignmentis not
all that certain,might be the point of origin of
Latericriodusand the lcriodontidae.
5.7.5 Family Dislomodontidqe
Klapper,I9EI
Conodonts assembledhere and assignedto
Distomodus, Rotundacodina, and Coryssognathushaveabasrcallyquinqui- or seximembrate
skeletalapparatusin which ramiformcomponentstend to haveshort,weaklydenticulateor
adenticulateprocesses,
and the Pa and Pb positions are occupiedby stelliscaphate
and pastinate elementsor their reducedderivatives.
The latter characterprovidesa link with other
prioniodontideconodontsand suggests
origin
of the Distomodontidaein Late Ordovician
balognathids such as Gamachignathusot
Sagi odontinq.
Thereis someditrerenceofopinion in the literatureasto the apparatuscompositionof DiJtomoduskentuckyensrBransonand Branson
(Fig. 5.19),type-species
ofthe stem stock,and
this afects interpretationsof familial taxonomy and phylogeny. The apparatusrecon-
ffi
E aarppel[ /$ol
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^q
plole{s
eql qcrq { uI snleftdd?
el?Jqrueujrxes
pequJsop
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lueuele
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e qlud serceds
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egl
ur
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sl (tr6t)
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'3Jd
^qtunI
VJNOCONOJ AHJ
THE MAJOR CONODONT GROUPS
Er illustratedin
of specieswith a
l|ls in which the
biodi na-like stell)igornodus made
iDning of the Sib s'enlockian.In
cock several lintd this by charti the lower part
i bush(lhe typi
of the type
-!Lsanotherbuilt
ic
aar anticipated
i (Icriodellidae);
r along a little
Pa elements
f-d
fi.
as Hadrola.ein i1shisancestralto
-h
Jeppsson
;rcies
Epical of Dlsloae apparatusof
knt of thoseof
'i rtre Pa and Pb
E and pastinate
$. Phylogenetic
r,rhese two posipedomorphosis,
*t-\. RotundaE Silurian and is,
rest|al stock of
d
' Co4'ssognathus
d- These are refuents with a
trs of a sort that
| from the much
Endacodina duIDsson has suglsent a species
of
rnts formed by
iiscaphate,with
.ln Coryssognamd the cusp is
Itcre appearsin
Dbe a very short
Miiller lnd Miille\
5.7.6 Family lcriodontidae
1957
Included in this family are Latericriodus
Mtller, 19621Icriodus Branson and Mehl,
1938;two apparentlydistinct goups of species
assignedto Pelekysgnathw'[homas, 1949,AntognathusLipniagov, 1978;and an as yet unnamed genus, recognized on phylogenetic
groundsby Sandbergand Dreesen(1984)and
basedon a groupof late Devonianspeciespreviously refened to luiodus
Icriodonts are characterizedby a basically
multimembrateapparatusthat includesbi-, trior merely segminiscaphate pectiniform elemenls in the Pa position.If thereare elements
in other positions,they are an assortmentof
rather variable coniform or only weakly denticulatedramiform elements.
Weddige and Ziegler (1979) advocate divi
sion of Icrioduss.l. into two groupsofspecies,
Pa
one that formed bi- and trisegminiscaphate
Paeleelements,and onewith segminiscaphate
ments.For the first goup they usedthe generic
nameLatericriodus(Miiller, 1962),for the latter, Icriodus (Bransonand Mehl, 1938).I follow weddige and Ziegler in the summary that
follows but note that lcriodus in this senseis
almost surely polyphyleti.c.^fhat is, I- angustus
both wilh the segminiscaand,I. angustoides.
phate Pa elements typical of luiodus, have
beenshownby Klapper and Ziegler(1967)and
Carlsand Gandl (1969)to haveevolvedat different times from different ancestors in
Lateriuiodus.
Serpagli (1983) has demonstrated that
the apparatus of Latericriodw woschmidti,
founder of the genus,is seximembrate(Fig.
5.20). A bisegminiscaphatepectiniform element occupiesthe Paposition;apastinate,lowpyramidalconiform elementthe Pb position;a
more compressed pastinate coniform (or
"acodiniform") elementthe M position;and a
morphologically intergradational series of
alale,tertiopedate,and dolabrateelementsthe
Sa,Sb, and Sc positions,respectively.Klapper
and Philip (1971),however.did nol recognize
homologuesof the S-serieselements of .L.
woschmidtiin the apparatusof Z. Iatericrescens
(Fig. 5.20),which they concludedwasbi- or trimembrate.Thus,ifZ. woschmidtiandL. l^ter'
icrescensdid,,indeed, form part of a single lineage,a featurein the evolution of that lineage
may havebeenthe lossofS-serieselements.Al-
Fig. 5.20. Elementsand apparatusesofspeciestypical ofthe Icriodontidae
/ ! - ^/ J n
<$-<:/-{/-9,4
UV V V
Latericrioclus woschmidti
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VJNOCONO]
EHJ
THE MAJORCONODONTGROUPS
* hasbeenlittle
EI of sucha sepdberg and DreeP. inclinatus
-ol centralmemd
rian group, also
I lor "oistodon;r ro b€ reported
Ldovician.
$ har-ealsoiden{c "oistodontits of one memL- onian species
r hrr thought inio differentlinh elements,the
lese species,are
E rhree denticle
timaely be dermes for these
Bouckae odus
rrarlable for one
ls of the poorly
b- posterolateral
$ T-shapedout&s'. Origin and
I tnown, but its
: apparentlyrer hlpersalinelalfe s olked out
r earliestDevotitanjte lcriodus
[fti7) haveconil rhe history of
r) ard Weddige
Dlulionary pat4x of lciodus;
[984) have rei the history of
la-ol stratigraphE biofacies,and
r as Devonian
, prior remarks
:ms in icriodont
in to be solved.
br that either
tre elementsby
'71
Bergstrdm,
which species of luiodus have customarily 5.7.7 Family Polyplacognathidae
198t
been identified or the single-rowedsegminiscaphate elements taken as typical of Peleldys- I-ate in the Early Ordovician a speciesof the
gnathw may have been produced indepen- Amorphoqnathusgroup (Balognathidae)was
dently at different times in different lineages. apparentlythe progenitorof a stock of prionAlso, origins of the family are by no means iodontide conodonts characterizedby a recertain.
duced, bimembrate apparatus of bilaterally
that asymmetricalstelliplanateand pastiniplanate
Klapper and Murphy (1974) suggested
Icriodus sensu lato mlght have evolved from elements.Recunent groups of such elements
Pelekysgnathusindex, but they found no spec- are the basis for recognizinga successionof
imensthat would bridgethe gapin morphology speciesof Eoplacognathus(Fig. 5.21). These
that separatesPa elements of Lateriuiodus specieshave been of considerablebiostratiwoschmidti and P. index. Nicoll (1982) re- graphicutility in rocksprimarily of earlyMidgardedRotundacodinadubius(Distomodonti- dle Ordovician age.The stock seemsto have
dae) as the most likely ancestor,and Broad- died out, however, early in the late Middte
headand McComb (1983)regardLatericriodus Ordovician.
woschmidtias a neotenousderivative of PeDuring its early Middle Ordovicianheyday,
davis. Fihraeus (1984) suggestsderivation of Eoplacognathusseemsto have giyen rise to an
Pedavis from Distomodus, in which presum- important line ofspeciesthat Bergstrdm(1983)
ably he included Rotundacodinadubius.I re- refers to the genusCahdbagnathus(Fig. 5.21).
gard the neotenousroute suggested
by Broad- The bimembrateapparafisof Cahabdgnathu\
headand McComb (1983)as unlikely because lrke thai of Eoplacognathus,is composed of a
it was devisedprimarily to explain the origin of pair ofstelliplanateand a pair ofpastiniplanate
the Pa elemenlsof I. woschmidlifrom thoseof pectiniform elements.Howeyer, those of the
Pedavislatialatus,andit thus ignoredotherele- Cahabagnathusapparatusform mirror-image
rnentsin the apparatus,which are surely not pairs, whereasthose of Eoplacognathusare biOn the otherhand, laterally asymmetrical and have more simply
the samein thetwo species.
there is substantialsimilanty betweennon-P ornamentedupper sides.
ofR. dubiusandL
elementsin the apparatuses
woschmidti,as Serpagli(1983)has shown,and
the remote ancestors of R. dubius certainly Fig,5.21. Elementstypical ofthe apparalusesofspebuilt scaphateelementscomparableto thoseof ciesassignedto genemofthe Polyplacognathidae.
L. woschmidti.(Of course,the scaphateelements someof them built wereeven closerto
those of Pedavis,which I ally to Iuiodella because of greater similarities in M and Pb
elements.)
While I do not claim to have specialinsight
into the ancestryofthe icriodonts-or eventhe
clairvoyancethat would be requiredto makea
monophyleticunit out of this family-I suggest
that a derivation of Latericriodus woschmidti
from Rotundacodina dubius explains tlte char
acters of more of the Lateriuiodus apparatus
than doesa derivation from Pedavls.Pending
badly needed additional information on Iate
Silurian conodontfaunas,I submit that this is
a more straightforwardroutethan is the oneby
way of Pedavis,whoseown origins are yet to be
amDlvdocumented.
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VJNO(ONOJ IIHJ
THE MAJORCONODONTGROUPS
;Eas of generainlfueDb at top of
I- aid P elements.
species haye
-b- most authors
tiaisodus (Fig.
qls of the typeanlv lacked an
q be a reduced
E\eomultiois-
r
rFi g . 5 . 2 3 ) ,
Esj apparatusof
! llulrioislodonti-
whosebest-knownspecieshas a bimembrate
apparatus,was probablyalso derived from an
early speciesof Neomultioistodus(e.9.,N. clypaas)by simplificationof elementsin the P positionsand by lossof elementsin the M and S
positions.
Lower Middle Ordovician (Whiterockian)
strata in North America also yield two recurrent groups of dominantly hyaline elements
that are identical in plan to those by which I
recognize speciesof Pteracontiodus, Neomultioistodus, and.Multioistodus. but with shorter
cuspsand longerprocesses,
which bear numerous white denticles.Thesegroupsrecord two
closely related speciesof Paraprioniodus (Fig.
5.22).The olderspecies(P. costatus)apparently
had a geniculateconiform elementin the M position of its skeletalapparatus,whereasthe M
position in the apparatus of the currently
unnamed younger speciesis occupied by a
dolaUrate(or "cyrtoniodontiform") ramiform
element.
Pteracontiodusrznges,in the form ofat least
one unnamedspecies,into Middle Ordovician
rockswell abovethosedominatedby the other
membersofthis family, but apparentlywasnot
ancestralto other conodonts.Dzik (1983)reported that, in a manuscriptthen (and still) in
preparation,he and I might consider Pcraprioniodus, a Pteracontiodusderiyative, as the
ancestor of the important Ordovician Phragmodus.Forvaious reasons,however,that now
seemsunlikely, and Phragmodusis shown in
Fig. 5.17 as a possibledescendantofan as yet
unidentified speciesin the Tripodus compLex.
5.7.9 Family Plectodinidae,
new
In this new family I include prioniodontide
conodontswilh a sexi-or septimembrateskeletal apparatusthat includes a pastinate (or
"dichognathiform")elementin the Pa position,
an angulatepectiniformelementin Pb, a dolabrateor bipennateelementin M, and alate,di
g).rate,and bipennateramiform elementsin
the Sa,Sb,and Scpositions,respectively.
Sinceabout 1969I have assignedconodonts
with such apparatusesIo Plectodina Stavffer
and AphelognathusBranson,Mehl, and Branson,takenin a multielementsense.However,I
havealsoassignedto thosegeneraa numberof
specieswith apparatuses
in which the Pa position is occupiedby an angulatepectiniformelement (e.g.,P. tenuis,P. florida) becauseof the
often repeatedobservation that, in large samples from a succession
of late Middle Ordovician populations,the denticulatedlateral process of the 'dichognathiform' Pa elements
rotates gradually anteriorly and ult.imately
fuseswith the short, adenticulateanteriorprocessto producean elementin youngersamples
that is formally angulate.As Bergstrdmand I
noted in 1972,this sequenceof eventsresults
in a type of apparatusthat is basicto the maJor group of dominantly post-Ordovician
conodonts herein identified as the order
Ozarkodinida.
Becauseil is unfortunateto have a major
evolutionarystep such as developmentof the
Ozarkodinida buried taxonomicallywithin a
genus,I suggestthat Plectodinabereservedfor
specieslike P. aculeala(FiE.5.24)with pastinatePa elements,and that thosespecieswith angulate Pa elementsbe transfered to another
genus, which I will identify here only as
"Plectodina."
The problem with Aphelognathusis just the
reverse.That is, the type-speciesof Aphelognathushasan ang)latePa elementand is thus
appropriatelyan ozarkodinide (instead of a
prioniodontide)conodont;the two species
with
pastinate Pa elementsnow assignedto this
genus(1. gigas,A. kimmswlckensls)will need
a new genericassignment,
or can be accommodated temporaily in Pleclodina. Thtts, in
Fig. 5.11,I show the rangeonly of Plectodina
sensu stricto. Species of " Plectodina" and.
Aphelognathuswith angulate Pa elements are
included in the ozarkodinidefamily Spathognathodontidae,whose content and strati
graphic distribution are shown digrammatically in Fig. 5.36.
PleAodina s.s. (in the form of P. acalea/orZes)almost madeit to the end of the Ordovician, and specimensrepresentinga host of
variable (and largely undescribed) speciesare
common in samplesfrom low- and intermediatelatitude Middle Ordovician rocks.
The Plectodinidaeseemsalso to be the best
home for Scyphiodus(Fie. 5.25),whoseonly
known specieshas a unimembrateskeletalapparatusthat consistsof an anguliscaphate
pec-
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VINOCONOJ
IIHJ
9L
THE MAJOR CONODONT GROUPS
, end Phragmodus
tlnenrs
from the
r-!Ls
Dalota (Sweet,
irns representE clearly pastina liat lateral and
I dements reprebFnetic stages.
Ets retain only
r mark their pasir.i was apparirdontide cono(the other
rtir
r) ro develop a
and the
-trl unimembrate
..
t HasJ' 1959
crefer the genera
t to the family
tEssary because
bm genus Cyrr specimensthat
ttrugmodus cog&ble synonym
t ClroniodontiD- as subfamily
rirrin over the
family Phragmodontidaeeslablishedby Bergstrtimin 1981.
Phragmodus (Fig. 5.24), characteristic of
low-latitude Middle and I-ate Ordoyician
faunas,has a distinctiye seximembrateapparatus in which the P positionsare occupiedby
pastinate(or "dichognathiforrn")elements,the
M position by a dolabrate(or "cyrtoniodontiform") or geniculateconiformelement,and the
S positionsby the "phragmodontiform"ramiform elementsdistictive ofthe genus.The latter, which include alate, tertiopedate,and
bipennatetypes,all have a terminal or subterminal cuspwith smooth sidesor with a sharp
costaon one or both sidesand a long, arched,
denticulatedposterior processat the crest of
which is a major denticlethat may rival or exceedthe cuspin size.
In 1972 Bergstrdmand I included Piragmodus and Plectodina in the family Cyrtoniodontidae and speculatedthat these genera,
which dominate Middle and Late Ordovician
conodontfaunasin the North AmericanMidcontinent, originated rn Pfioniodus. lt now
seemsmore likely that the prioniodontideplan
of the skeletalapparatusin speciesof these
three generais indicative of a common, but
probablynot direct,origin. Thus,in Fig. 5.1I I
show Prioniodus, Phragmodus,and Plectodina
as heterochronousdeiyaliyes of Tripodus.
With somereservationsI alsoincludeBryantodina (Fig. 5.24) wtth Phragmodusin the Cyrtoniodontidae.Bryantodina has an array of
ramiform elementsin S positions that look
very much like the distinctive "phragmodontiform" elementsof Phragmodus.However,P
positions are occupiedin the apparatusesof
Bryantodina speciesby carminate and angulate
pectiniform elementsthat are quite diferent
from the pastinate P elements ol Phragmodus
and cannot be shownat this time to be either
ontogeneticor phylogeneticmod.ificationsof
suchelements.In favor of an associationwith
Phragmodus,Inote that in the apparatusofP.
cognitus one of the pastinateelementsis replaced phylogenetically by an angulate pectiniform one that retains a clue to its pastinateancestryin the form ofa riblike offseton the outer
faceof the cusp.No suchofset ornamentsthe
outer faceof either P elementin the apparatus
of Bryantodina typicalis, however; hence, if
eitherdevelopedby phylgeneticreductionofan
orginally pastinate form, that reduction is not
documentedby extantcollections.
5.7.I 1 Family Rhipid.ognathidae
Linhtriim,
1970
ln the Treatise Bergstrom assembledBelg$roemognathus,Appalachignathus,Rhipidognathus, Carniodus, ar^dHistiodella in the Rhipidognathidae,which was assignedto the Prioniodontacea (the Prioniodontida of my taxonomy). Following publication of new information on Histiodella, which I have cited in
Section5.7.1,I feelrnorecomfortableassigning
that genus to the Oistodontidae.Also, Carniodus is out of place here,becausethe seximembrateskeletalapparatusofits only known
specieshas a pastinateelementy/ith threedistinctly denticulatedprocesses
in one P position
and an alateelementwith a long, denticulated
posterior processin the Sa position. Because
neitherofthesecharacters
is a featureofthe a[F
paratusof any other rhipidognathidspecies,I
feel that Carniodus should be reassigned.The
only prioniodontidegroupin which Carniodus
could be accommodatedis the Balognathidae,
but thereis little morphologicresemblance
between most of its skeletalelementsand those
formed by typical balognathids.Furthermore,
Siluian CarnioduJ is separatedfrom the Ordovician balognathidsby a considerablestratigraphic gap. Consequently,I follow Bischoff
(1986)in assigningCarniodus1o the Pterospathodontidae(Ozarkodinida),whosenumerous
Silurianspeciesbuilt apparatuses
that included
strikingly similar P elements.
Generathat remainin the Rhipidognathidae
include specieswith tri- to septimembrateskeletal apparatusesin which P position(s) are occupied by structuresthat are fomally angulate
or carminatepectiniform elementsbut are allied morphologicallyto the "acodontiform" P
elementsof Tripodusand olher slem pfioniodontidesthroughpossession
ofa distinctrib or
costaon the outer faceofthe cusp,which may
be markedon the basalmargin by a conspicuous boss or lappet. In short, those elements
may be regardedas very simplepastinateones,
in which no sharpedgeor denticulateprocess
ever deyelopedalong the anterior edgeof the
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YTNOOONOJAHJ
9L
THE MAJORCONODONTGROUPS
long posteriorprocessthat is arched,laterally
compressed,and surmountedat the point of
maximum geniculationby a denticle that is
wider and conspicuouslylonger than the oth-
Fig, 5.27. Elementstypical of the skeletalapparatuses
of speciesassignedlo th€ PeriodoDtidae.Top elements
in each row Pa, followed downward by Pb, Sa, Sb, Sc,
and M elements.
ilEathidae. Figl.L I 1974).Saele-
t hown part of
lhr is the genus
re- so it is the
? 5.u .
fulstrom, 1970
(Fig.
arkodina
| fu Treatise to
i:donlidae, reI the same famFience with, or
E_cenus,but berr studies (LdfEcies of the two
tt€ same Ea y
d subsequently
P. aculeata,
i
4Daratus, with
ts. one of which
ftiform";
dolaments, with an
i end an S series
hts have short,
;rocesses and a
ers. The stratigraphic(and probably phylogenetic) predecessor
of P. aculeata,P. flabellum,
had geniculatecon.iform,not anteriorly denticulate dolabrateelementsin the M position,
and the ramiform S elementswere apparently
neither so profuselydenticulatednor so conspicuouslyarchedas those in comparablepositionsin the apparatusofP. aculeata.An even
older speciesof Periodon,which has not yet
in the Proteusand
beennamed,is represented
Eleganszonesof Jamtland(Ltifgren, 1978)by
adenticulatealate elementsinterpretedas occupantsof the Sa position and by adenticulate
pastinate(or acodontiform)elements.The latter, and the "tortiform" angulateelementsthat
evidently replacethem in the apparatusesof
youngerspecies,are the elementsthat indicate
that Periodon and the Periodontidae belong in
the Prioniodontida.Ofcourse,the distinctiveS
elementsofP aculeataar;ldP. Srazdri,the speciesof Periodon that I know best,are also geDerally similar to thoseof Phragmodus,and this
led me in the heydayofform taxonomyto sugEestlhat Periodon might be an older name for
Phragmodus-I no longerhold sucha view.
Speciesof Miuozarkodina, such as M. flabellum, tbe type-species,have quinqui- or seximembrate skeletalapparatusesin which the
roundedanteriormargin of pastinate(?) P elements is not developedas a processand the
short, anteriorlydeflectedlateralprocesscommonly bearsno more than a singledenticle.M
elementsare geniculateconiform structures,
and S posirionsare occupiedby alale. tertiopedate,and bipennateelementswith a long,
straightposteriorprocess.
According to the vielvs just summarized,
Periodon ernergedftom roots in eady oistodontid populations by developmentof a highly
differentiated skeletal apparatusin which processes of component elements gradually
becamelongerand more conspicuouslydenticulated. As in the presurnably ancestralOistodontidae,P elementsnever differentiatedinto
conspicuouslypastinatestructures,but S elements becamedistinctively archedand denti
culated.In Microzarkodinadevelopmentwas
apparentlysimilar. but the posteriorprocesses
of S elementsare not arched,M elementsdevelop no anterior denticles, and the feature I
interpret as a lateral processin P elementsremained short and bore only a singledenticle.
: Jql lnq 'luaur
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VJ.NOCIONOJ AHJ
8L
THE MAJORCONODONTGROUPS
t'-..".
f,\f'
+""-.-)
.//
,IDAE
flDAE
Erized
in Fig.
lE:ltuses have
rd- or reduced,
hida are charblic apparatus
! and includes
r digtrate eleUl slr or seven
Ilrs tend to be
lcrgradational
E chamcteristirhire matter is
i- ir elements
k ro the axial
, fr io diffi.rse,
fuzzy-edgedclouds that are irregularly distrib- tends downward to form a distinct anticusp
uted through cusp and denticles.Speciesof a thal bearsseveraldenticleson its upper margin
few early genera(Chirognathus,Curtognathus) and is thus a true anteriorprocess.It seemsunand of at least one Triassic one (Parachirog- necessaryto invent a new morphologicterm
nathus)formed elementswith little white mat- for suchan element,which (if needbe) might
ter and thick lamellaethat are mineralizedby be described as a quadriramate structure with
apatite crystalliteswhose prism surfacesare a short,adenticulateposteriorprocess.
oriented parallel to the direction of growth.
Ancestors of Erraticodon (and thus the
Suchelementstend to breakasif they werein- Prioniodinida)are not known, but it is surely
ternally fibrous, and Bransonand Mehl used reasonableto rcgardEqaticodon as the plexus
this feature to distinguish the order Neu- from which Erismodusevolvedin the shallow
rodontiformes,
Middle Ordovician seas of North America.
It is alsoa characteristicofthe Prioniodinida Speciesof Erismodus (Fig. 5.29) differ from
that there is little differencein size or basal thoseof Erualicodonprimarily in that the Pa
morphology betweenthe distinctive digyrate componentsof their seximembrate
apparatuses
elementsassignedto the two P positions,the lack a denticulatedanterior process(but retain
ramiform elementsthat occupy M positions, a bosslikeanticusp),andbipennateM elements
and those that make up the morphologically lack denticleson the short posterior process
conservativesymmetry-transitionseries.For (and thus have sometimesbeen describedas
this reason, diferent authors have assigned "falodontiform"). The oldest speciesof .Eriscomparableelementsof variousspeciesto dif- modus with which I am acquainted are repreferent positionsin the apparatus,with the re- sentedby typically "fibrous" or "neurodonsult that homologiesare not alwaysobvious.I tiform" elements.However, the skeletalelewill attemptto be unirbrm in the followingdis- ments of later species(e.g.,E. quadridaaylus)
cussion(and in the classificationin Appendix have more white matter and commonly break
A), but my yiews do not alwayscoincidewith like othernonfibroustypes.This suggests
to me
thoseof other authors.
that "fibrous" histologymay somehowbe relatedto the probablyquite unusualcondilions
that obtainedin the shallow,very warm MidBransonand
5.8.1 Family Chirognathidae
continentwatersin which earlyEru'smodus
deMehl, 1944
veloped.Later species,suchasE. quadridactyEqaticodon, the oldest known chirognathid /as, may representforms adaptedto the more
and the most ancientmemberofthe Prioniod- normal conditionsthat probablycharacterized
inida distinguished thus far, appearedlate Midcontinenl seas later in the Middle Orin the Early Ordovician, when it achieveda dovician.
very wide biogeographic distribulion E. patu
The bizarre Chirognathus(Fig. 5.29),repreCooper(Fig. 5.29),the oldest(andbest-known) sentedin the shallow, tropical waters of the
species,
is distinguishedfrom otherchirognath- Nofth AmericanMidcontinentby a singlespeids primarily by featuresofthe curiouselemenl cies, is allied morphologicallyto Erismodus,
I assignto the Pa position.At first glancethat from which it differsprimarily in a skeletalapelement appearsto be pastinate(or "dicho- paratuscomposedof elementswith a scarlike
gnathiform"), and its associationin the same attachmentsurfaceskewedto oneface,an alate
apparatuswith a Pb elementthat is an exten- Saelementwithout a posteriorprocess,and an
siform digyrate(or "oulodontiform") element, M elementwith denticulatedanteriorand posand thus typically prioniodinide, might be terior processes.
Cutlognathus(in which I intaken to suggestthat E. patu straddlesthe cltde Trucherognathus and Polycaulodus) is
boundarybetweenthe Prioniodontidaand the alsoa Iikely derivativeof Erismodus,butitsapPrioniodinida.Closerinspection,however,re- paratus has not yet been competely reconvealsthat the distinctivePa elementof E. palu structedand, like Chirognathus,it is not likely
is also basicallyan extensiformdigyrale ele- to be on the "main line" ofprioniodinide evoment, but the anterobasalpart of the cuspex- lution. Thesetwo generaare significant,how-
|f,! lrr"unuop eql
E aqr uI 'uoqec
4J alull dre^ Jo
q JrDolur uEunl
a eqr trrJeJ(snpo
pPQ ro snpolno
lrnFs lEre^es
'JSprurporuoud
€uotaq ,{Feelo
qtJeddE uI Pol?l
PO el?'I Eu€uer
t|mqpuSotsrtd
Drq ueDr^opro
Dsl lEql sercadso
I|s ueunlls
^[Jeg
Er sr lueudola^
t()-r.rp ur pes?eJc
r'lEl eql ul seuoz
F: tlllt\r\ '$IJ?rx
I uoos aJe.{\serc
Dpro elpprr\l ar€l
gorlrpuoc snoJo
'wtn.b snpolno
q" rqnoPel I eq
pJrlDrtg Puesnp
r" ilJeolc sr .tnpo
dasJo -rxeseqlJo
I'mpolno Jo sal.
I uJe ed sql pue
aruoJrurporuoud,,
a e^vf snporust
$urs 'pezl?rcods
puas o rcqt tsee
trssod ? Jo (€861
qq{o lseequou
Duepr^a
^ddeJcs
n aql ur lugurele
lIrE u?Jo peelsuo
E! Jelleur elrq.?t\Jo
I qloq ur slueulele
P rruoJrsuelxo3JE
r ? Jo snlElBdde
pu? qneluQqrssu
q ot f,lleelcsursos
f9 EtJ) snpolno
4td:rllu0l
08
'pequcsepuoeqlou sE.q.oaproqqlHJo ]lua:uliele
ed eq.L slueuolo ed pue'qd 'I I'rs 'qs
uel
^q
eql pJeAol po.rolloJ'tqBu eq1le ore sluoEole eS sasnlEredd€lls ul'(o aprDqqtH'Dutpoluoltd'snPol O) eeprutpor
-ioi1pue(tnqniion4S'mpowsug'uopocrpll3')oepqleuSorqJequoIecrd,{lsorcedsJososnlEreddv'62'5'8!tr
tttt.
^"t.tt^,^
"nqteu6otlr.lJ
slppteqqtH
389
euuoJrluopojn0N
mu aqt snql puB
;Jql Surequr 'JaAa
THE MAJORCONODONTGROUPS
v
tratar) and PrionEoEed loward the
8l
ever,in beingthe primary "fibrous" conodonts platformedpectiniformelementsin P positions
and thus the nucleusof Bransonand Mehl's and startedto experimentwith ontogeneticor
phylogeneticreductionin the developmentor
Neurodontiformes.
mineralizationof elementsin various apparatus positions,the Prioniodinidaremainedcon5.8.2 Family Prioniodinidae
Bqssler,1925
servativeand developmentproducedno starOulodus (Fig. 5.29), the oldest prioniodinid, tling modifrcations.
seemsclearly to have evolved from Erismodus,
Devonian workershave not yet dealt in deas Schijnlauband I suggested
in 1975.In the tail with the Prioniodinida,but Klapper and
apparatrusof O. serralrs, the type species,there Philip (1972)have demonstratedthat the apare extensiformdigyrate(or "oulodontiform") paratus of Hibbardella angulata (FiE.5.29),
elementsin both positions,only minor patches type-speciesoflllbbardella, differs from that of
ofwhite matterin any element,and a dolabrate Oulodus primaily in that the alate Sa element
(insteadof an anteriorlydenticulatebipennate) of the former has a denticulateposteriorproelementin the M position.Thesefeatures,and cess,whereasthat of Ou/odrs hasa stubbyposscrappy evidencefrom Ordovician rocks in tenor processthat lacksdenticles.For this reanortheast Oklahoma (Amsden and Sweet, son the assemblyof elementsfrom the upper
1983)ofa possiblyslightlyolder species.
sug- Middle Devonian of Manitoba that Uyeno
gestthat O. seryatusmay have been somewhat (Norris, Uyenoand McCabe1982)hasrecently
specialized,sinceall potentialancestorsin Er- referredto Oulodusmay representa speciesof
ismodus have an extensiform digyrate (or Hibbardella, instead.
"prioniodiniform") elementin the Pa position,
Murphy and Matti (1982) have established
and this patternis repeatedin all youngerspe- the genusErika (Fig.5.30) for a single Early
ciesof Oulodus.However this rnay be, the plan Devonian species,which had a seximembrate
ofthe sexi-or septimembrate
apparatusof Oul- apparatuswith extensiformdigyrateelements
odusis clealLyanticipatedin thoseoftnJmo- in both P positions,a breviform digyrateeledus and.Erraticodon.its ancestor.and therecan ment in the M position,and a symmetry-tranbe little doubt about closerelationship.
sition seriesthat includesalate.extensiformdiOulodus qutcHy adaptedto the probably rig- gyrate,and bipennateramiform elements.The
orous conditions in shallow subtidal parts of apparatusof Erika divarica, the only species
late Middle Ordovician seas,in which its spe- known, is sirnilarin many waysto that ofboth
cies were soon establishedas ecologrchall- Oulodus and.Htbbardella; however, the M elemarks. With enlargementof those marginal ment is not much like that formed by species
zonesin the later Ordovician,Oulodusalsoin- of either genus,and the alateSa elementlacks
creasedin diversity, and its evolutionaryde- a posterior processand thus difers from any
velopmentis recordedin Late Ordovicianand other prioniodinideconodontwith which I am
Early Silurian strataby a succession
of chron- familiar.
ospeciesthat is of considerableutility in Iate
Accordingto Nicoll (1980),the apparatusof
Ordovician biostratigraphy (Sweet, 1984). Late Devonian specresof Apatognathus (Fig.
Pristognathus,representedby a single,short- 5.30)is also prioniodinide, and I include the
rangingLate Ordovician species,is closelyre- genuswith the Prioniodinidaein Fig. 5.28.The
lated in apparatusarchitecture to Oulodus and elementtypical of Apatognathusin form taxclearly belongs with that genus in the onomy was apparentlyan occupantof the M
position.
Prioniodinidae.
SeveralSilurian speciesassignedeither to
Sparling( l98l) has also concludedthat the
Oulodusor Delotaxis(which I include in Ozl apparatusofat leastoneMiddle Devonianspeodus) cany the Prioniodinida through the Si- cies of Prioniodina (P. tortoides) (Fig. 5.29) is
lurian into the Devonian but proyide evidence closely comparableto those of Oulodus and
of very little changein basic plan or diversifi- Hibbardella in element tlpe and number and
cation. In the Early Devonian, when sonleof in morphologicfeaturessuch as style of denthe dominant ozarkodinidesbeganto develop ticulation. Sparling'sdiscussionsuggeststhat
suaurelo ES (eepru
FsrrrelS 'Ic'g '8rd
@tl? SnUeE rxJoJ
peloJ
IrelJ
[I pu? ^q
u?rrllled
1 tuaurols ol"J^t
dsnJ erll Jo as?q
ql qll\r A lcull
pue
^lpJ?^\u^\op
a sTlpoluottdorpJ
p ueql reSuol eq
Daq ol puel luaur
Torlrsod qd oq1
lp eql sr slueuod
EsgruErs, ll?rced
Ereds 'Uoqs fue^
rruoJrrusr moJJo
Pve '.(snpotuol.td
furpnlcur) slueur
rll qloq uI sluelu
[f,ullsrp epnlJur
3re u,t\ou)l serc
Pls aqlJo slueur
Frmu E ur parr g
+|Jqr Pu?'(tI6I)
5 tuslueleulnuj
mpotuoLtdorpl
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$rlr?Jerp osp sr
decxa ,r\eJP
ds pspnlcur ^luo
osle
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tDue elerpeuur
potlo ueeq e^eq '8961
oql tse8itnsot procor peqsllqndeq1ur enlrt sr pu? (9161)
^qroN ^q
ejeql 'eJeqdnoJ8srql o1 1ruSrsseI pue '?pIuI 'peorxsid snqnu?DpqruDT
Jo serJeds ruJoJ
slueuolE elu?u elqelr"^ElsePlo
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eqt eq ot suees '896I 'peolxeu snlpuSopDlx
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pJo sertedstuau
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suoursod qS aql ueql selr,edssnqpuSopDlx Jo -etuoc luosoJdal lEr{l sdnoJEluarlnceJ ITJJoJ
e^eqol ourpouoSrT pue 'snporuoudoaN 'o apnqqlH
sJsnleJedde
eql ur suonrsod
d pardnJJo
r(lelrt ejour ere D aDpIuSoW Jo selcedsuuoJ 'D aDplruBDI4[ 'snqwuSopDlxJo sercodsuloJ
s€ pezruto3eJ,(lsnor^oJdslueruelalDqt lsoE8ns s? pegrluopr ,{Fnoherd stueuolo (@tddlssrs
I tnq 'sortods.tn?truSopDtxJo sasnlz:eddeeq1 -sr] I) ueuelsoqc1?q1suodel (186I)p?orxau
'?purpoIUoud
Jo uonJrulsuoJJJsrq ul sluauroled ezIuSoJeJ
lou pp (1861)peoJxed snpoluottdoaN
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pu? u"ru?^[^suu0d
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ruo+ atu?J trl{datrlElls
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pJluournJop? s?q leql ptutpoluouda^tlJull
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eq
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^
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puEJspJoeql roJeprurpoluoud
erpJ
aseql teql 1csJsql alldsep 'elu e?prurPoruorJd
^le^n?IeJ
,{Oueunuo
sIqJ pos?qir\ouar? suj"u oqt tursn ,{q oseceql eq tq8ru sql l?ql
ot IBcrSoI
suaas
e^eqI pu? 'esuesluaueleqFur ? uI
sznedssnqnuSop0lx gcrq^\ uo sdnoJBluounc peledrcDu?
-er oql ol suo4rppelsuuelod se(1861)p?orxel{ uorlducsep[zr,l.eDurporuoudJo serJedsJeqlo
1286I)Ilrew
puer(qtuny{ urorJslueuoler{lgJo seqflels stueuele€d puE'qd 'I{ 'cS 'qS,tq Uolot po/$olloJ'lq8u uo sluaurolo
pS(oeplurporuoud)
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sorceds
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ff-m
VJNOCONOJ AHJ
THE MAJORCONODONTGROUPS
Sa
Itio odinidae).
Bftom \'lurphyand
ftiotrs to the relgn zrlrus species
iendv logical to
E relalively rare
ler components
ristically signifiEoostruction of
? i,il.
*ment
Kladooccupied
by
$
poslenorproVibbardella;
the
oplmon,
not
Ii
mnrs rypical of
r ard (probably)
rition by bipend m form taxEls df multielebped dolabrate
led as form sped I l98l) did not
rEonstructionof
Ls species,but I
$- recognizedas
; are more likely
r the apparatuses
the Sb positions
d them.
*ment
Klado,of the Prioniodmp here.There
d ro suggestthe
immediate ancestryof Kladognathus, bnt spe- time regardedsuch"enantiognathiform"digyciesofthe genusappearto havebeenmembers rate elementsas occupantsof the Pb position
of nearshore,shallow-waterassociationsthat in the skeletal apparatus of Ellisonia and,in
also included speciesof Cavusgnathus.
With thoseofa succession
ofspeciesof multielement
only a few exceptions,this ecologicpreference Xaniognathus and.Cypridodella. Hence, occuris also characteristicof the Prioniodinidaand rence in the Pb position of ttle ldioprioniod.us
may help confirm the assignment.
apparatusof elements reminiscent of these
Idioprioniodus(Fig. 5.31)wasdiagnosedin a later Paleozoicand Triassic specimenspromultielement sense by Merrill and Merrill yides strongsupportfor the origin of Ellisonia
(1974),and their reconstructionhas beencon- and. Xaniognathus in ldiopioniodus or a
firmed in a numberof subsequent
studies.Ele- closely related genus.And, becauseI regard
mentsof the skeletalapparatusofthe two spe- Xaniognathusas the most generalizedcompocies known are large, mostly hyaline, and nent of the famity Gondolellidaeand E1lr'sozla
include distinctive digyrate pectiniform ele- as the centralradicalofthe Ellisoniidae,strucmentsin both the P positions;dolabrateM ele- tural similarityof Pb elementsplaysan imporments (including the tlpe form speciesof Nea- tant role in my ideasconcerningthe origin of
prioniodus);and a symmetry-transitionseries thoselineages.
offour ramiform elementswith largecuspsand
very short,sparselydenticulatedprocesses.
EsLindstrdm,1970
peciallysignificantin this arrayofskelelalcom- 5.8.3 Family Bactrcgnathidae
ponentsis the diglrate elementthat I assignto In the Treatise Atstin and Rhodes assernbled
the Pb position. Lateral processesof this ele- the distinctive Mississippian generaBactroment tend to be quite short (althoughthey may gnathus, Doliognathus, Dollymae, Eotaphrus,
be longer than those of other elementsof the Scaliognathus, and Staurognathusin a family
Idioprioniodus apparatus),and they project Bactrognathidae.
However,theywereunableto
downwardlyand anteriorlyso as to form a dis- suggestthe relationshipofthis obviouslyintertinct V with the inflated, but undenticulated, relatedstockto any ofthe othergroupsofconbaseof the cusp.A closelysimilar type of di- odonts describedin the Treatise.
gyrate element is common in collectionsof
In 1985 Kad Chauff demonstrated(to my
Permian and Triassic conodontsand was re- satisfaction,at least)that Bactrognathus,Dofened by Clark and Ethington (1962) to rhe liognathus,and,Staurogna.thus,
the centraleleform genusEnantiognathus.I have for some mentsofthe Bactrognathidae,
haveskeletalapFig. 5.31. Elements and apparatuses typical of species assigled to Kladogaathus and ldtopioniodus (pironiodinidae). Sa elementson right, followed ro left by Sb, Sc, M, and P (or pb and pa) eremenrs.
qr ilqsqord s?r\
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5 JO -IXOS E SI
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sauesI J pu€ (S Jo) uoDrsue4-tuleuu^s
uI sluourelouroJrrrr?JJo ,{eJ" u? e^eq (Zt S
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,89
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t8
THE MAJORCONODONTGROUPS
liplanate and thus similar in their generalorganizationto thosedevelopedat varioustimes
in unrelatedprioniodontideand ozarkodinide
stocKs.
apparatus of
;i<re
r :rE indistinguish-
duded that the
i a groupofconC4ptotaxis that
! represented
in
nd While this
lion- I wonderif
r boking in the
oftheir incepr- prioniodinide
lost diversetaxd biogeographFironments. EpIoropes seemto
srccessfully, if
Ghore habitats.
asr sdmespecies
an sucha stock.
ddirional signifhioniodinida ber corectly, they
lb.tform rnaking
m15. As in other
platformed elec- Doliognathus,
7 in the curious
t modificationof
aions.It may not
slite their origin
lrres. theseplatcame to be stel-
85
tina(FiE 5.33),ktown thus far only from a distinctive Pb elementfrom the I-eonardianHess
Limestoneof Texas.Later Permiancollections,
describedin detail by Wardlaw and Collinson
(1986)and Croft (1978),includethe complete
array of elements,which indicatesth^t Swee5.8.4 Family EllisoniidaeClark,1972
lin a had a seximembrateapparatuslike that of
The first representatives
ofthe Ellisoniidae,in- Ellisoniabuthad,an elementin the Pb position
terpretedby von Bitter and Merrill (1983)as that is reminiscentoflhe triradiateextensiform
two speciesof Ellisonia, appearedin the Ato- digyrate element of Ordovician Erraticodon,
kan (Pennsylvanian).The youngest species the earliestprioniodinideknown.That element
known are of Late Triassic(mid-Carnian)age, has one long and one rather short lateral proand the type-species(-E lrlaJsicd) is Early cessand a stubby posteriorprocessthat bears
Triassic.In architectureof their skeletalappa- only one or two denticles.
ratuses,speciesof Ellisonia (Fig. 5.33)are simSpeciesof Merrillina (Fig. 5.33), another
ilar to speciesof ldioprioniodus,from which Permianellisoniid,developedskeletalapparatheyalmostcertainlydeveloped.The apparatus tusesthat were closelysimilar in their conforis a sexi- or septimembratecomplex, with mation to thqseof Sweetinaspeciesbuthad exmorphologically conservative alate, digyrate, tensiform digyrate Pb elementsthat lack a
and bipennateramiform elementsin the S and postenor process. In successiyelyyounger
M positions,an extensiformdigyrateelement Permianpopulations,the short lateralprocess
in the Pa position and, in the Pb position, of Pb componentsbecomeseven shorter,and
another digyrate element, with processesre- in the youngestPermian collectionsI have
flected sharply downward and anteriorly. I seen,the short processis largelyvestigial.Spehaveearliercommentedon the significanceof cies of both Sweetina ard, Merrillma were eyr
these "enantiognathiform" digyrate elements dently restrictedto the Permianand were apin establishing relationships within the parentlybestadaptedto very shallow,agttated,
probably hypersalinewater. Commonly, their
Prioniodinida.
Obviously only minor morphologic ditrer- largeelementsarethe dominant (or only) comencesseparatethe apparat.uses
of Ellisonia spe- ponents of collections in which they occur;
ciesfrom thoseofpartly contemporaneous
spe- however,thoseelementsalso tend to be fragcies of Idioprionioda.r.Such featuresinclude mentary,soreconstructionofapparatuses
is difthe tendency for processesof Ellisonia ele- ficult and only a few speciesare known in
ments to be longer and more profuselyden- detail.
ticulated than those of Idioprioniodus and.for
In the Early Triassic(Scythian)E/llsoziawas
the attachmentsurfacesof E/ftsonraelements joined (or replaced)in shallow-waterenvironto be composedof smallbasalpits borderedby mentsby speciesof Furnishius,Hadrodontina,
prominent zonesof recessivebasalmargin.In and Pachycladina(all illustratedin Fig. 5.33),
some IdioprionioduJ elements the attachment which formed seximembrate apparatuses
surfaceexhibits featuressimilar to thosejust whosecomponentelementsare distinguished
described,but in nonethat I have seenare pits by wide zones of recessivebasalmargin and
so small or recessive-margin
zones so broad whosealate Sa elementslack a posteriorproand conspicuous.It might also be noted that cess. Pa elements are extensiform digyrate
the M elementin apparatuses
of ldioprioniodus structuresin apparatusesof speciesof both
speciesis a distinctive dolabrate structure, Furnishius and Hadrodontina, blot extensiform
whereasthe one in Ellisonia apparatsesis bi- Pb elementsdevelopan additionalrow ofdenpennateand readily confused(by me, among ticles on the anterior face. In Pb elementsof
others)with elementscommon in Sb positions Hadrodontinathe supplementalrow joins the
in the apparatuses
ofother species.
main denticlerow at both ends.In comparable
Early in the Permian a speciesof Ellisonia elementsof Furrrishiusthe supplementalrow
wasprobablythe ancestorofa speciesof Swee- joins at only one end and additional denticles
a
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VJNOCONOJ:lHl
98
THE MAJORCONODONTGROUPS
Fig. 5.34. Iareral and undersideviews of a Pa element
of Gladigo ndoIella (Ellisoniidae).
ThaynesFormation of Utah, and one ofthose
collections also contains a number of platformed pectiniform elementsthat are gradational in morphologywith the unplatformedPa
elementsof.E. triassica,and.are also identical
in form with the elementson which Huckriede
(1958) based Polygnathustethydis,the typespeciesof Gladigondolella Mnller (1962) (Fig.
5.34).Paull (1983)noted the samefeaturesin
her large Triassic collections and describedthe
Lower Triassicspecimensasrepresentatives
of
G. meeki.
A distinctiye featureof gladigondolellidPa
elementsis the fact thal narrow platform brims
completelysurroundboth anterior and posterior processes,
hencethe elementsare formally
anguliplanate.Comparable elements of the
gondolellids, which also occur in Triassic
rocks,are segminiplanateand thus are readily
separatedfrom those of Gladigondolella.
I
9
5.8.5 Family Gondolellidae
Lin^ftam, 1970
The late Paleozoicand Triassicconodontsassembled by Treatise afihorc in the families
Gondolellidae and Xaniognathidae form an
obviouslyinterrelatedstockthat is bestconsidered as a singlefamily. The name Gondolellidae,coinedby Lindstriim in 1970,is the oldest
one available for the combined family.
Gondolellid conodonts formed a basically
seximembrateskeletal apparatusdistinguished
primarily by angulatePa elements,breviform
digyrate Pb and M elements (the former "enantiognathiform"), and a slrnmetry-transition
series that includes alate Sa elements with a
Es rhe Sa element
I ross indicate that
lndcles in either
i- E. triassica, is
tkrions I made
Lo*er Triassic
I
long.profuselydenticulated
posreriorprocess;
extensiformdigyrateSb elements;and bipennateScelements,commonlywith long,delicate
anterior and posteriorprocesses.
In the apparatusesof generalizedgondolellids,the postenor processof Pa elementstends to be short
and fragile. whereasthe anterior processis
stout and is commonly surroundedat midheight by a well-developedmidlateral rib. In
gondolellids,however,the posterior
specialized
processof Pa elementsdisappearsand the segminate structuresthat result may also develop
platform segmentsthat are confined to the
sidesor joined aroundthe posteriorend ofthe
cuspto form a continuousbrim.
For many yearsit has been my contention
that when gondolellid conodonts developed
segminateor segminiplanate
elementsin the Pa
positionthey alsoceasedmineralizationofelements in other positionsin the apparatus.Accordingto this view, the Gondolellidaewould
include specieswith both unimembrate and
seximembrateapparatuses.
This interpretation
ofgondolellid organizationis basedon my observation that collections made from rocks
within the range of the Gondolellidaecommonly conlain either the segminatePa elements or an assortmentof ramiform and pectiniform elementsfrom which it is possibleto
assemblea completeseximembrateapparatus.
Only rarely do collectionsinclude a balanced
assortment of platformed and ramiform
elements.
For a number of gondolellidspecies,particularly Triassiconesreferredto Neospathodus,
Platyvillosus, Epigondolella, Mosherella, and
Misikella (seeFig. 5.35),I believemy original
idea holds. That is, those speciesformed unimembrateapparatuses-or at leastonly those
elementsin the Pa positionsweremineralized.
However,evidencefrom a coupleofnatural assemblages [one almost certainly coprolitic
(Rieber,1980)lindicatesthat othergondolellid
specieswith segminateor segminiplanatePa
elementsmay have had complete seximembrate apparatuses.Fot example,Pseudofurnishias (Fig. 5.35)and at leastsomespeciesof Gar?dolella (Fig. 5.35) and,Neogondolella(Fig. 5.35)
appearto have had apparatusescomposedof
segminiplanateelementsin Pa positions and
ramiform elementslike those tvoical of Xan-
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VJNOCONOJ:IHJ
THE MAJORCONODONTGROUPS
?s.udolutnishtus
be [n rheapparatuses
Sb.5c.M, Pb,andPa
E badmultimembrate
L rom thosein the
l egminate or segrcloped from time
re sp€cializedspeled only by substigminiplanate) ele:!'r-ative angulate
h orhers, developdniplanate Pa eleEcompanied either
n ofthe apparatus
d€ments in M and
Origin of the Gondolellidaeis obscure.Ap- dle Ordovicianage;the youngestare from earparatusesof Xaniognathusspecies,the most liest TriassicstEta.
generalizedgondolellids, are very close in
Except for a few generatransferred from the
structure to those characteristicof Ellisonia Prioniodontida,my conceptof the Ozarkodispecies,and it is likely that both stocks ap- nida is essentiallythat of the superfamilyPopeared at about the same time in Atokan lygnathaceair the Treatiseclassification.In
(Pennsylvanian)seas. I have already com- Fig. 5.36 (and in Appendix A) I divide the
mented on the likelihood that Ellisonia devel- Ozarkodinida into 12 families. Treatise auoped from the prioniodlnid ldiognathodus, and, thors recognizedonly 6.
the gondolellidsmay havehad a similar ances- The cenlral family in the taxonomicscheme
try. This suggestionreceiyes some support of Fig. 5.36, the Spathognathodontidae,
infrom the fact that PennsylvanianXaniognathus cludesan assortmentof ozarkodinidegenera
arld Gondolella characterizea relatiyely deep- that is superficially quite heterogeneous.
It
waterbiofaciesthat overlapsone in which ldi- could be made much lessso by includingEogoprioniodus is also very cornmon. Ellisonia nathoduswith the Polygnarhidae;by erecting
and its kin, of course,are especiallycommon separatefamilies for the compactbut distincin rocks that represent very shallow-water tive stocksnow represenled,by Ancyrodella, by
environments.
Amydrotaxis ar.d Ancyrodelloid.es,and by PoMy views on relationshipsbetweenmajor lygnathoideq and,by further revision of Bispageneraof the Gondolellidaeare summarized thodus and,reassrgnmentof the resulting parts.
diagrammaticallyin Fig. 5.28,and apparatuses In effect,that lype of raxonomictrimmingwas
characteristicof typical gondolellidspeciesare begunin the Treatiseby crealingseparatefamillustratedin Fig.5.35. 11 is my suspicion,llies for Kockelella and,Ancoradella and. for
however,that formation of segminate(or seg- Pterospathodusand.its kirr.
miniplanate) elements in Pa positions (and
I have resistedthe temptation to erect the
commonly concurTentapparatus reduction) severalnew generaand familiesjust suggested
happenedrepeatedlyin the history ofthe Gon- on the groundsthat sucha task is bestunderdolellidae,perhapsasan expressionofiterative takenby someonefar morefamiliar than I with
adaptationby successive
membersof the Xan- all the speciesinvolved. On the other hand, I
iognathus-Cypridodella stock to similar envi- have not hesitatedto referAlternognathusand.
ronmentalconditions.If that werethe pattern, Siphonodel
la to the famity Elictognathidae,
bethe taxonomysummarizedin Fig. 5.28is vastly causespeciesofboth arewell setofffrom those
oversimplified,ofcourse,althoughit is difrcult retainedin the Spathognathodontidae
and auto imaginea defensiblealternative.
thors who havecontributedmost to unraveling
the evolutionary history of the two generic
stocksagreethat.despiteresemblance
in some
morphologicparticulars,this stock originated
5.9 The Ozarkodinida
Dzik, 1976
in the Spathognathodontidae
independentlyof
The basic skeletalapparatusof ozarkodinide the Polygnathidaeand at a much later time in
conodontsis sexi-to septimembrate,
with P po- the Devonian. Those authors may not, howsitions occupied by carminate and angulate ever, be too happy about my choice of
pectiniform elementsor their platformedana- Elictognathidae.
logues.In ozarkodinideconodontswith fully
I have also divided the Idiognathodontidae
developedappantuses,elementsin S and M of lhe Treatiseclassificationinto Gnathodonpositions are commonly generalizedin mor- tidae and Idiognarhodontidae,
becausesuch a
phology and, from published descriptionsat division recognizes
a naturaland highly signifleast,are much the samefrom one speciesto icant interval in the mid-Carboniferous
history
the next. Pa elements,on the other hand,r/ary of both groups.The Palmatolepidaeis newly
greatlyand haveprovidedthe principal means establishedfor a phyletic lineagepreviously
of taxonomic differentiation.The oldest ozar- submergedin the Polygnathidae,and the Dikodinideconodontsarefrom rocksoflate Mid- plognathodus-Sweetognathuslinea$e is re-
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06
YJNOCONOSAHI
THE MAJORCONODONTGROUPS
!
6
6
I
s
t
aftoAE
g/ l
6/ |
A'@HfIDAE
aEalhus;Pgn. = Pro!obrious.
batidae Hqss, 1959
nls are a Yariable,
d.rp. as befits their
)zarkodinida. The
sexi-or septimembrate
skeletalapparatusis basically simple, with bladelike carminate and
angulatepectiniform elementsin both P posi
tions;a dolabrateor bipennateM element;and
an undistinguishedsymmetry-transitionseries
that includes alate, diglrate, and bipennate
elements in the Sa, Sb, and Sc positions,
respectively.
The alate Sa elementsof most of the earliest spathognathodontids(e.9., "Plectodina,"
posterior
Aphelognathus)
lack a well-developed
process,and breviform digyrateelementsoccupy the Sb position.A few, however,haveSa
elementswith long,profuselydenticulatedposterior processes
and at leastonespecieshad developed extensiform digyrate Sb elements by
early in Cincinnatian(I-ateOrdovician)time.
The Spathognathodontidaeare recordedfirst
near the baseof the Nonh American upper
Mohawkian (i.e., the Rocklandian) by conodontswith severaldistinctive types of apparatus. One, typified by the specimensBergstriim and I (1966)referredto "Bryantodina"
abrupta (bat probably referable now to Yaoxianognathusabruprrs),has a simplecarminate
Pa element,an angulatePb element,a dolabrate(or "cyrtoniodontiform") M element,and
a symmetry-transitionserieswith a bipennate
Sc element, an extensiform digyrate Sb element,and an alateSaelementwith a long,denticulated posterior process.Bergstriim and I
( 1966)referredthe ramiform assemblyto a species we named Plectodina? posterocoslatatI
transferred it to "Bryantodina" abrupta in
t979.
A secondtype ofearly spathognathodontid
is
represented
by the LexingtonLimestonespecimens from Kentucky and Ohio that Bergstrtim
and I (1966)made the typesof Bryantodina2
stauferi. '[he apparatus of B-? stauferi includes only slightly arched, angulatePa elements,conspicuouslyarchedPb elements,and
an alate Sa elementthat lacks a denticulated
posterior process.Other componentsof the
skeletalapparatusof B.?stauferi haye yel to be
identified. 8.2 stauferi ranges into only the
early part of the Late Ordovician (Cincinnatian), but a specieswith morphologicallysimilar
P elements, "Ctmognathus" pseudofissilis of
Lindstrdm (1959), may have continued this
line into evenyoungerpartsofthe Ordovician.
The third and by far the most common type
9l
of early spathognathodontid
is representedby
speciesthat have been included in Aphelognathus and. ln Plectodina.The type-speciesof
Plectodina,however,
hasa pastinateand not an
angulateor carminatepectiniform elementin
the Pa position and is thus a typical prioniodontide. The type-speciesof Aphelognathus,
and all other specesofthe genusexceptMiddle
Ordovician A. kimmswickensisand.A. gigas,
has a pair of typically spathognathodontidP
elements.Thus, in this volume I will assign
species such as Pleclodina tenuis, P. florida,
and P. bullhillensis to "Plectodina," for their P
elementsare angulateand carminatepectiniform elements,not the pastinateand angulale
structurestypical of Plectodinas. s. Similarly,
Aphelognathus gigas and A. kimmswickensis
will have to be included for now in "Aphelognathus,"in recognitionof the prioniodontide
structureoftheir Pa elements.
" Plectodina"(Fig. 5.37)evolved from P/eclodina, it the late Middle Ordovician, an event
documentedin greatdetailin sequentialcollections from the L€xington Limestone of the
Cincinnati Region.Aphelognathus(Fig. 5.31),
which wasalsocommonin the late Middle Ordovician,cameto be evenmore widespreadin
marginalpartsofthe lowlatitude seasthat covered North America in the Late Ordovician.
The youngesl "Pleclodind" speciesare evidently the Earty Silurian "P." hassiand "P."
oldhamensis,which have beenreferred in most
studiesto Ozarkndinabut evidently represent
"Plectodina," becausetheir skeletal apparatusesseemto hayeincludedbreviform digyrate
(or "zygognathiform")Sb elements,insteadof
the extensiformdigyrate(or "plectospathodontiform") Sb elementstypical of Ozarkodina.
The three typesof spathognathodontid
conodonts describedthus far from Ordovician
rocksdiffer from one anotherin severalparticulars and probablyrepresentdiferent lines of
development.That therewere also other lines
is indicatedby discoverya few yearsagoofdistinctive but as yet undescribedforms with extensiform digyrate Sb elements in early Late
Ordovician rocks in both Kentucky and Oklahoma and by the sporadicoccurrencein late
Ordovician strata in Europe of more or less
typical representativesof Ozarkodina (e.g.,
Ozarkodinan. sp. of Bergstriim),whoseP, M,
and Sc elementshave been known for more
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VJNOCONOJ AHJ
z6
THE MAJOR CONODONT GROUPS
dorinellinaMiller and Miiller, 1957(Fig. 5.38),
which Uyeno (in Norris, Uyeno, and McCabe,
1982)has shownto be closelysimilar in apparatus construction to Mehlina Youngquist,
'aa :rosl widespread
l. x- \1. Pb, and Pa
Ers, dolabrate or
r Eansition series
I ror "plectosparor "hindeodelSbard Sc posiISa
element of
t
hnculated poste*r Devonian and
Fcies with triraL position to Par-
,as O:arkodinaad
.lL andPaelements.
hasa welld€vel-E.:
91
(Fig.5.398,5.39C,and 5.39D);in the EarlyDevonian of Ozarkodinaseffi (Fig. 5.39G), the
shortliyed speciesfrom which Lane and Ormiston (1979)derive both Eognathodus(Fig.
(Fig.s.a0).
19as
5.39H) and Polygnathus(Fig. 5.39I) of the
Unfortunately,the completeapparatusesof Polygnathidae;and in the late Middle Devospecies nian, when Ozarkodinasemialternans(Wirth)
only a handful of spathognathodontid
havebeenworked out, so it is not alwayspos- spawned" Polygnathus" latilossdtusWirlh and.
sible to decide if Ozarkodina or Mehlina or the Palmatolepidae.
Evidently,as Bransonand
Pandorinellinais the appropriategenericas- Meht noted long ago,the notoriouslyunstable
signmentfor many Devonian and early Car- basal cavity of spathognathodontidPa €leboniferous forms or which generic stock is mentsexpandedand migratedtowardthe posinvolved at various evolutionary junctures. terior end for the last time in the early MissisSandbergand Ziegler (1979) have somewhat sippian.Speciesproducedin that episodeand
simplified the problem by restricting Pandoi- their successors
herein the famare assembled
nellina Io those spathognathodontidswhosePa ilies Anchignathodontidae,Sweetognathidae,
elementsare morphologicallycloseto thoseof Cavusgnathidae,
and Mestognathidae.
Carminate
P. inslla (Stautrer),the type-species.
In a secondtype ofdevelopment,sidesofPa
Pa elementsofthe latter havea high finlike an- elementsexpandedat or near midlength to
terior processthat exhibitsa slightrightlateral form lobulate lappets on one or bolh sides
offsetat itsjunctionwith the posteriorprocess. above the central part of the basal cayity. Not
which is built of much shorterdenticles.
uncommonly,the upper sidesof theselateral
Althoughtherearecertainlywell-markeddif- extensionsgrew longer phylogeneticallyand
ferencesin other elementsof the spathognath- cameto supportone to seyeralnodelikedentiodontid apparatus,dividing the family into cles,commonlyarrangedin singlerows.Pa elesubordinatetaxonomicgroupsand tracingde- ments modified in this way characterizeearly
(Fig. 5.39F) in the
velopmental history has been accomplished stagesof Pterospathodus
primarily throughstudiesof Pa elements.This early Silurian; Amydrotaxis and. Ancyrodelstructureexhibitsgreatplasticity,as I attempt Ioides (Fig. 5.39E, 5.39P,and 5.39Q) in the
to showdiagammaticallyin Fig. 5.39.
early Devonian; and Ancyrodella(Fig. 5.39K,
In typical representativesof Ozarkodinq and 5.39L)much later in the Devonian. Each
(Fig. 5.394'),the basalcavity of Pa elementsis oflhesemodifrcationepisodesrepresents
a geoa groovelikeindentationalong the lower mar- logicallyshort excursionby spathognathodongin that expandslaterally to form a cuplike tid ozarkodinidesinto the never-neverland of
structure at about midlength of the element platform building in the Pa position.Although
and tapers toward the processextremities. Trealrseaulhorsidentifiedthe platform-buildFrom time to time in the long history of the ing excursionthat Ied to Ptercspathodusand its
Ozarkodina-Mehlina-Pandorinellina plexus, allies as a stock worthy of recognitionat the
however,and possiblyalso in the still-obscure family level, the otherswere not so identified
Early Silurianhistory of "Plectodina"and spe- and I will not tamperwith this situation.
cies derived frorn Aphelognathrrs,the laterally
In speciesof Silurian Polygnathoidesand
expandedportion of the basalcavity cameto mid-Devonian Torlodus, Pa elements were
occupya longerand longersegmentof the un- modified in yet a third way. That is, riblike
dersideof the posteriorprocess.Elementslike thickeningsof the sidesof basicallycarminate
this (e.g.,Fig. 5.39.Gand 5.39H)have a ladle- spathognathodontidelementsdevelopedinto
with the unmodified smooth-surfacedbrim- or shelflikeplatforms
or scooplikeappearance,
anteriorprocessservingas handleand the pos- alonga segmentof the element,commonlythe
teriorly widenedbasalcavity as scoop.An ex- posterior process(e.g., Totlodus, Figs. 5.39J
ercisein basal-cavitymodificationlike this ap- and 5.41),or around the entire element.The
parently precededdevelopmentin the early latter type ofdeyelopmentcharacterizes
the bi(Ftg.5.40),which
Silurian of Kockelellaand the Kockelellidae zarreSilurianPalygnathoides
ffi
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VJNOCONOS
t6
IIHJ
THE MAJORCONODONTGROUPS
denticleson the upper surface.One denticle
row, the primary one,extendsfrom one end of
the elementto the other.At the anterior end it
forms a single-rowed"blade," which is bordered on either its right or left side by the
shorter secondaryrow. Numerous modifications ofthis basicpatternleadto ozarkodinide
groupsidentified here as the farniliesGnathodontidae and Cavusgnathidae.
History of the Spathognathodontidae
was
marked by severalintervals of accelerated
innovation that were expressedprimarily in
modificationsof Pa elementsalong lines just
discussed.
In the late Llandovery(Silurian),for
example,migrationofthe expandedpart of the
basal cavity toward the posterior end initiated
morphologythat cameto be typical of Pa elements in the apparatusesof speciesof KockeIella and,the Kockelellidae(Fig. 5.39B,5.39C,
and 5.39D).At aboutthe sametime, lateralexpansionofthe centralportion ofthe basalcavity, eventually to form denticulated platform
elements,establishedthe pattern characteristic
of Pa elementsin apparatusesof speciesof
(Fig. 5.39F) and other memPterospalhodus
bers of the short-lived Pterospathodontidae.
The Early Devonian included two closely
spacedepisodesof innovation in the spathognathodontid stock. In the first, shortly after the
beginning of the Devonian, Pandorinellina
(Fig. 5.38)separatedfrom contemporaryOzarkodina populations (Fig. 5.38) by developing
Sa elementswith a denticulatedposteriorprocessand distinctively finned Pa elements.At
about the sametime, iniiation of Amydrotaxis
(Fig. 5.40) andAncyrodelloides
(Fig. 5.40)was
heraldedby rapid, lobulate expansionof the
centralpartsofbasal cavitiesin Pa elementsin
segmentsof widespreadpopulationsof Oz4rkodina remscheidezsrJ.
In the secondEarly Devonian episode,posteriorwideningofthe basal
cavity in Pa elementsofsom€whatlater populations ofthe sameOzarkodinastockled to O.
selrt(Fie. 5.39G)then, promptly, to early representatives of Eognathodus, with doublerowedPa elements(Figs.5.39Hand 5.41),and
Polygnathus, with triple-rowed Pa elements
(Fig. 5.39I). Although Klapper and Johnson
Fig. 5.40. Elements and appamtuses of sp€ciestypical of genera of the Spathognathodontidae. The complete aFF
palatrrsesof AmydrcIaxis, Polygnathoides, and Mehlina ate shown, but only the P elements of Ancyrodelloides ite
illustrated. The M and S elementsofthe latter are no1 known.
Rq
Fo€trathodontidae
;@r (C), (D) various
a6hui (J) Tortodus;
)l\atr'cEd B$palho-
lFratuses in speGnothodus (Figs.
Eronian and Carfrhodus and its
r 5.39M, 5.39N,
G of thesespecies
Iions similar to
J inclinata (Fig.
)bur difer in gradtbparallel rows of
Ancy.ode
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ueluo^o(I el?'I eql ur seEeNI?re.\os1€suon"ln
-dodur stueruelopdJo suort?cgPou ',(UeuIC
'oeprluopoqleu8oql?ds
eql ur
osl? 1r spnlcurI 'u?rus"Jceql Jo Pu3 eql l?eu
enssrlnoqqrnpe4dxee^eq01srEaddP
lPql at?a
-ur[ cueueSouour'altuls ? ur popnJcur,(Fpeor
sr esoreleql lcols eqt osnEceq'pu"(0t S tIC)
sapolppo.thuv Jo tueurdole^ep ol pel leql
eq1 sIFJer eposldasrql
euo usruo^e(
^lleg
',{llecrSolorldrol
l lcors (It'S 'Brl) o apor{ruv
VjNO(IONOJ
AH]
THE MAJORCONODONTGROUPS
Iorphologicalty,
J Devonian one
Anatodelbides
d that aroseis
mogeneric lin:d sithout issue
I include it also
fuenrs in popr l:te Devonian
B 5.i8) led to a
>rosed forms by
rirs of ScaphigEr- vanousspefrtathus. Mehfuvrhodus (Ftg.
lhdonddae and
Sulus initiated
!I Devonian,to
ln. sratigraphic
Carboniferous.
fuionodella are
Edir \) asparts
lnarhidae. Simfitgnathus arc
f rEpresentatrves
frlloe that pracBlsrtomferous
-r.{.
Fhogllalhodonrs sirh both sinI elemenls are
Eirarive reports
| ,{usdn (1974);
l[ Evenif 8. stcrrr single-rowed
lde Sa elements)
:ft long-ranging
ac reassignedto
fuatidae, BispaEI surgery.That
l4ler, Sandberg,
lare that th€ "8.
t r" branchesof
Eed from .8. s/ar |le "aculeatus"
E4lnJ, the type-red in Bispathorould be needed
r- branch.There
f familial assign-
me']tfor Bispathodusand the new genus.Since spathognathodontid species such as Ozarku
the situation with Early Deyonian Eognatho- dina abrupta(Aldridge, 1972)beganto widen
dus (Fig. 5.41)seemsalmost preciselyparallel, toward the posterior and also to expand
it seemsto me that thebestsolutionis to assign conspicuously to the sides, giving rise to carbroadly conceivedand unrevisedBispathodus miniscaphateelementswith a relatively long,
[and its obvious later Carboniferous deriva- even-crested
anteriorprocessand a rathershort
tive, Rhachistognathus(FE 5.41)1,to the postenor processsurmountedby small dentiSpathognathodontidae.
clesthat declinein sizetoward the processtip.
Finally, Bispathodusstabilis, with conserva- Near the end of the Llandovery, continued
tive, single-rowedPa elements,wasapparently modification resultedin Pa elementswith an
the stock from which speciesassignedto Psea- even shorterposteriorprocessand a large,aldopolygnathus(Fig. 5.al) developedat leasr mostposteriorbasalcavity whosebasalmargin
twice, oncein the late Devonianand a second hasan almostcircularprofilein superiorview
time in the early Carboniferous.The complete (Fig. 5.398).Pa elementsofthis sort characterapparatusof Pseudopolygnathas
has yet to be ize the apparatus of Kockelella ranuliformis
described,but Pa elementsof primitive species (Walliser),which alsoincludesangulatePb and
in both the Devonian and Carboniferousiter- bipennate (but neverthelesspick-shaped)M
ations of the genuscombine open, Bispatho- elements and a symmetry-transitionseries
daslike basalcavitieswith at leastrudiments composed of bipennate Sc, asymmetrically
of three (not 2) denticlerows.Later speciesin alateSb, and symmetricallyalateSa elements,
both lineagesaredistinguishedby carminiplan- the latter lacking a denticulated posterior
ate Pa elements,with a largebasalpit that is process.
surroundedby a zone of recessivebasalmarSubsequentdevelopment of the Kockelella
gin. In this respect,Pa elementsof Pseudopol- lineageinvolved litlle apparentmodificationof
ygnalhusarehomeomorphicwith thoseofPo- elementsin the M or S positionsbut substanIygnathus,znd,the two heterochronouslineages tial changein those in the Pa position. That
now includedin the genusv/ill probablymerit changeis markedfirst by developmentofshort
sepamtegenericand familial identity sometime processes
on one or both sidesofthe cusp,with
in the future.At present,it seemsbestto retain no conspicuousmodification in profile of the
Pseudopolynathusand," Pseudopolygnathus"in basalmargin. The basalcavity of Pa elements
the Spathognathodontidae
with the ancestral in samplesfrom slightlyyoungerpopulationsis
BispathodusstabiI is stock.
laterally restricledbeneal.ha posleriorprocess
that is somewharlonger than in specimens
from older populations;and, in the Pa ele5.9.2 Family Kockelellidae
Kkpper, 1981
ments of the youngest speciesof Kockelella
Toward the middle of the Llandoyery (early (Fig. 5.aD, lateral processes
lengthenand also
Silurian),the basalcavity of Pa elementsin a bifurcate,and the basalcavity beneaththem is
Fig. 5.42. Elementsand the apparatustypical ofa speciesof(ockele/1a, typical genusofthe Kockelellidae.
FrEdde lelele)s
[[nJ ueeq
EJ
aqJ
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pue pEorxad'3'e) ecueuncJo-oo
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-edsDulpoqozo ,fuHoduatuoc Jo sasntHedd? -"ruJoJsueJleslexeq'DllaplJox IJro+ pe^lo^e
eql ur seJnFruls olqeJ?durooueql eJolu serrods D apDloruv teql [pelou o^eq (9161) red
snpou.toisreJo sesnt?Jeddeeql uI asoql elqures -d?I)I pu? {olr?g sel slsa8EnsAIuI?lec sluoul
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Jo
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eql Jo Ir/t\oDl sI elou tuqloN
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uI ul3l"ru J?s?qo^rsseceJJo auoz peoJq P
pus {crueg ,{q tq8noqf slueurole 'eJns 0q oI
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pepunolns sI qtlq,rr' '1rddsn3qns? ol peJnp
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Jo lcols ? s€ e?p4uopoqledsoreld eql pJeEeJ sr acEyns lueurqcql? oql lnq 'JosITl?A\s41g
I,{q asuesluetuelall -D!to^ ofiap\)ox Jo seuo elEqdecsqlalsaql Jo
I '(2861)utel{ pue
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lnru ? ur paqursep
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$ 1nq 3^oqe
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^llcuqslp
reEuol ou q uIEJeu
uo{J pea\er^ sz JBIncJIc
ps"q aquo olgoJdeql l?qt os 'pepulser requru
tltutDf
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€ 6'I
'oepnuopoqledsoreld eqt Jo ereuoBJo sorJedsJo lecrd,{l sasntEreddEpue slueurolg
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YJNOCONO"
86
IIHJ
THE MAJORCONODONTCROUPS
I
lntidae.
k
Cooper,)977
riq between Pa
Hus amorphodspecies of lnfred in a mula Uatti (1982),I
E a 5asto cko f
r Silurian prion,.i the Treatise.
t$ Barrick and
b d V and Sp o Iro of the three
ls (Fig. 5.43) re= of Distomodus
.!fructures in the
O::nrkodina sper between diagand specimens
ats of Pterospantrrmed by pub:g.- Rexroad and
per. 1976),and
rments becomes,
for me, a more compellingreasonfor reclassi- earliest speciesknown by carminiplanatePa
fication ofthe family.
elements,angulatePb elements,dolabrate(or
The Pa position in the apparatusof Pteros- bipennate)M elements,and an S serieswith bipathodus speciesvtasoccupiedby stelli- or pas- pennateSc, extensiformdigyrateSb, and alate
tiniscaphatepectiniformelementswith simple Sa elements, the latter with a denticulate
or bifurcate lateral processes
that may be ad- posterior process. Pa elements formed by
enticulateor may beardenticleson oneor both early speciesof the stem Eenus,Polygnathw
sides.It seemsobviousthat suchstructuresde- (Fig. 5.a4) (e.9.,P. pireneae,P. dehiscens,P.
veloped from Ozarkodina-Like
carminateele- gronbergi), are carminiscaphate; however, in
mentsthroughgraduallateralexpansionofone slightlyyoungerspecies(e.g.,P. lalicostatus,P.
or both sidesat midlength(Fig. 5.39F),just as inversus)the undersideof Pa elementsis planPa elementsof Amydrotaxis (FrE.5.40) andAn- ate, a condition that was attainedby develop(Fi!.5.40) arethoughtto havede- ment of a wide zoneof recessivebasalmargin
cyrodelloides
velopedin the Early Devonian. The Pb posi- at progressivelyearlier ontogeneticstagesand
tion in the apparatusof the type-speciesof consequentrestriction of the inilially broad
Pterospalhoduswas occupied by a distinctive basalcavity to a pit in forms representing
later
angulateelement,which bearsat midheight a stagesofeither growthor phylogeny.The pit is
stout lateral rib or ridge that projectsout and commonly elongate,situatedat elementmiddownwardlyon the outer side to form a chev- length,and enclosedwithin a keellikeridgethat
ron-shaped lateral lappet. An element of projectsbelow the lower surface,is narrowly
almost preciselythe sameconformationoccu- groovedon its under edge,extendsthe compied the Pb position in the apparatusesof pletelength of the element,and represents(as
various speciesof Early Devoniat Ancyro- doesthe pit it encloses)the under edgeof the
delloides.
element prior to the stage at which the basal
The stelliscaphate
Pa elementsof Astropen- margin beganto recede.
tagnathus(Fig. 5.a3),Aulacognathus,and ApIn Pa elementsof a few species(e.g.,Polysidognathus(Fig. 5.a3)are only slightly differ- gnathusspicatus) thereis no discemible keel in
ent in plan from those of Pterospathodus,and, the immediatevicinity of the pit, which is surall three genera were included vith Pterospa- roundedinsteadby a flat areaof varyingwidth
thodusin the Treatise concepl of the Pterospa- that mimics a structuretermed a pseudokeel.
thodontidae.However,if elementssuchas the Suchstructuresare common to the undersides
onesWalliser(1964)refened to the form spe- of Pa elementsof Siphonodellaspeciesand
ciesAmbalodus galerus and Pygoduslyra werc may,in mostcases,be usedto distinguishthose
also components of the Apsidognathlrs appara- structuresfrom comparablePa elementsofPotus, as has been suggested
by severalauthors, Iygnathw species.
then all bets are ofl at least for Apsidognathus.
Soon after the character of Polygnathus beUnfortunately,I have no other idea about re- came establishedin populationsof Early Delations or alternative classification of Apsido- vonian conodonts d escended,
from Ozarkodina
gnathusand corrlinueto showit in Fig. 5.36as selfi l-ane and,Ormiston ( I 979),three main lina member of the Pterospathodontidae.
Car- eageswere founded. Theseare identified in Fig.
niodus,as rcconstracted
by Bischof(1986)and 5.45 as the linga{ormis, robusticostatus,and
others,is also assignedto the Fterospathodon- costatusslocks.The natureofthese stocksand
tidae and not to the prioniodontide family their many branchesand the subsequent
Early
Rhipidognathidae as was done in the Zreatise. and Middle Devonian history of their compoThe apparatusof the type speciesis illustrated nent specieshave been explored in some detail
in Fig. 5.43.
by Weddige (1977) and Weddige and Ziegler
(1979),who drew Fig. 5.45and madethe interpretationsthat I summarizein the following
5.9.4 Family PolygnathidaeBassler,1925
paragraphs.
The few polygnathid ozarkodinides that have
The three rnain stocksrecognizedwithin Pobeen fully reconstructedhave a seximembrate lygnathus by Weddige and, Ziegler are distinskeletal apparatus characterizedin all but the guishedin the fossil record by the outline and
JOUoISOOAql Ol ?uIl?C eql enu4uoc
01 eires
-Iep orlJ 'suollcefloc ur ..luerJoq? J?odde ' '
qc
qlplr,\pur
pu"
pelou s?
,(q
ua{orq
erP
slueu
reltev
eftppei[
'GL6t)
'sapou
",
te
l&\
^q slueuelo
l?ql
-el} ed uadooJ d Jo ..en8uol, Jouetsodeql 'pu" azrs leoJ8 qceeJ
^Iuoururoc
'\Jols
uo
Ed poluou?uJo ,(lesJ"oc
squ
esJa^susrl
sna-toll8
'Je,/l.a^roH
1c?duro3 'lnols a^eq
-r{ aqlJosJqJuEJq
snous ur sJrJedsJo
sluau '(LT,
st's pue'gz'svs 'se'sn9 'Ez'9n'9'BIJ)
snql puB 'pua rouel pu"q raqlo eql uo 'qct{erq smButdpelleJ-osaqt
-eI5elq?J"durorolquroseJ
pu" Jo sorcedsluJoJteldeqtJo puo Jouolsodeql te
-sod eql le ..en8uol,,pepelJep,{lpJ?ndu^rop
f,lpret"l ? a^"q 'culauul{s?
eJe5lcols ,.en8uol,,paqqu
suell 'pedole^eplle\ e
^Ilqftls(6/6I)
pu?
srql
pJluec^lesJe
ur
serceds
eql
eq
01
relS
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01
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reqlro
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er?
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'Z'Sn'9'l'SVS trC)seqru?rqeerwslrpw lcols l]lxo"ts
snlDlsosrlsnqot
eql uI sarcads
JelelSuuueseJd seroedsJo
sluetueleed 'soEpu
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snutpDs etll Jo sorcodsJor o pue snuuotas d ao"Ins Joddnosoq^\,.en8uol,,? u oJ ol
pelceuepsl^Fou
Ed SaZ apeJEqJ /fulelulu,{Se SUOllS -elsod pJe/iru,$oppue
leql
JO SIUOIuAIO
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3 [luo sJeoq ruroJtqd e dole^ep pu?
Fculauflu,{s?
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rqr augrpqrrq/A
'(87'9n'9
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"d
'3rg)
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r"edd?
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{co1s
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ocqjns-Jeddn
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Ie€pldeloreurled)
pue (eeprqtetJ8(lod) srywu8qod 01 peuSrss?serjedsJo lecrd^l sesnlepdale pu? sluoruolg 'tt'9 '3!d
Er aqr ur pogrluspl
r-Erol. 'St'S'3!d
FOnPar euoceq
{d '(rt st's pu?
aq muol.taFalz
I aqt ol spuetxo
E pue .,en8uol,,
td I"Jutoruu^s
fE,lPJdde uadooz
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E IJ 3ql Jo pue
VJ,NOCONOJ AHJ
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
3 linguiformis
irnadves (fig.
rhich define the
r -rongue" that
ri[es- whereas
ah elements
of
d the serotinus
l
aniJrst icostatus
C i.15.1,5.45.2,
f tt'1., 5.45.33,
bse platforms
doped on either
t all rhe way to
ra- Polvgnathus
hddige and ZieI Secres in this
la\-e a laterally
Gu.-' at the posomparabte elerhes of the /in rsr-erse dbs on
antperi Pa ele!'nodes, which
b rhe posterior
tZ \Stl'.uaddop!'d
bFl'.oto8u DsoSM
gFo\aaUt OSOpOul
+talqJs otnutw d
1d d (Z):olarod
z \Ei 'oz) :1ulwrat
F d $l):wtaqks
I l8l sDupMw^sD
..snqtou3/qod,,
'F.L1 pue sulsH
lFrZ
Fqralg 'rt'S '8Id
ueeq ,{FoururoJ
a^?q
qu^l
srorllo
es
1"ql
FJe
Ielelels eql 1noq3atu 01 elqElr"AeuorlsrrrroJur
pue
regle8or '(tt'g 'Art) stdappupd
'(LV
ou f,lueJJncsr aJeqI'sn)upurutsD srxDrcsam
S
'8tg ur pe1a1sn11r Ue) sKDpsaW 'DutJaddolx
su q3nsa"prdolol.eurledeqt Jo roqurou e uro+
'snLlwuSolptuqJs ol pautrss?,{nuaunc sarceds sdeqrodro (S?'S '8ld) r{laorssnJolsu snqpuStl
-od erll uI rolsecue u^roDlun la,{ s? euos ruo{
aau 'aDpgalopwlDd ttllu0f
9 6'9
uEruoAJCJlpprhl lselel Jql ur pedole^op'sarJ
-eds ueluo^eq elrl e ururlsJpJo uorssecJns
e,{q pelueserdar'(9t'9 '8lC)snqpuSotl)uv
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,(qpelueseJdeJ
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e^?q o,$teql (586I 'euq puz reddelt) ](ltuec
sr ll pu?'(1961 'sulleH) ll?lep
ur
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snqt
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pre^es
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luasaJdal
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Jo) snoJeJruoqJ?J
Je^{o'I eql olur anunuoJ
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t ro episodically
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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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V.LNOCONOJ lHJ
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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VINOCONOJ 3HJ
90r
THE MAJORCONODONTCROUPS
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
$qL
bJ&
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
blJ;ds u?ruo^
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{reds)
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Erss€sluopouoJ
0.tl!u.ro!
t 69
x rql Jo pue eql
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6
lt
61
81
VJNOCONOJ
80r
:IHJ
THE MAJORCONODONTGROUPS
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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(t) (086I) rolSorzpue ,tuoqpues,oue-I,{q peuturr}.p suor1e1e1
eepsuoloqieugeqllo fusiolfq4 169.9.a11
6
I
N y, A/ V]1 ',t,( SN rlJ d
YINOCONOJ
AHJ
0l I
THE MAJORCONODONTGROUPS
D
t
b
r:!
(1)
I Zegler(1980).
frti.16) P. [email protected]$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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VJNOCONOJ
zl I
AHI
THE MAJORCONODONTCROUPS
ll3
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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VJNOCONOJ AHJ
tu
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-
d puB e^nourl
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VINOOONOJ
9I I
IIHJ
THE MAJORCONODONTGROUPS
h- and Aethotaxis
ts. P elementsof
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
J:-r.FJurlsp
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VJNOOONOC AHJ
8ll
THE MAJORCONODONTGROUPS
l&menrs
ig',trrs
on right,
have not
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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VJNOOONOJ:IHJ
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
t2l
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
AHI
zzl
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,
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d specific level.
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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
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233-246 i\ Index
E \I . Shimer and
'aL
ft. R. (1983).Paea frmily Icriodonlifiw. Fossilsand
*t'oniar. Icriodus
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fuatigraphie und
es der dstlichen
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conodonts from
51,'772-796'ar.
[tte multielement
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ria.n crisis and its
coDodont taxont7-158. Clark, D.
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gudies 9(2), 102teqstrom, S. M.,
Rhodes,F. H. T.,
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Cooper, B. J. (1975). Multietement conodonts
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(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
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Dzrk, J- (1976\. Remarks on the evolution of Ordovician conodonts. Acta Palaeont. Polonica
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(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,
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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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u?ruJr^u?n pu? ue€rueJv (g16l) 'V uuoJSJq.I -uoceJJo uorlEcgrssplcl?rllru?C 'QL6l) VJNOCONOJ AHJ
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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brforsch. 41, Lt- s'. (1986).Paf rhe Phosphoria
'our. Geol. 24(2),
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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EVOLUTIONARY PATTERNS
+ht also have
r and post-Misbnra. Logs conr series-by-series
Fnera also show
r of diversity cyFpiled recently
rsit-v of Ordovi:cies on a zoneiin this chapter
b evolutionary
Isent three logs
rs{evel diversity
cmire late Camhte Conodonta.
il. 6.2,and 6.3,
t fuctuations in
b, which Clark
k ofevolution.
Lof 4 first-order
'&hed line) for 40
blog indicatetimes
r separating diver-
v t-* _ v
_+
_* _
r-'
\v,'.-.,,
rdm.|..n
- oEvoxratl
l3l
x _r_
',//
I
\,,"."'
x[ _* _
i7
xf i-
-1__,.\-
"-
v|-
Mlssrsst?flar{
PEI{I{SYL
Fig. 6.2. Species-diversitylog (upper solid line) and log of origination-extinctionratios (lower dashedline) for 64
Devonian and Carboniferouszones.Stippled segmentsoforigination-extinction log indicarc limes when more speciesevolved than becameextinct; asterisksildicate times of sigdificantextinctionieparating diversity cyclesidintifred by Roman numeralsVI to XIII.
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.
x l -* -
xtv - * -
xv -*-
xvt_*-xv _*_xvt *xtx *_
I
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EVOLUTIONARY PATTERNS
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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VINOCONOJ AHI
,tl
EVOLUTIONARYPATTERNS
.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, Pelel<y* collectively for a Frasnian high-diversity epignathus, and luiodus (in a very broad sense). sodethat numberedat its peaksome53 conoThat burst introduced a short interval of high dont species.
diversity in the late I-ochkovian during which
Ziegler and Lane ( 1987)recognizean abrupt
the number of conodontspeciesbuilt back up decline in speciesdiversity at tlte end of Cycle
to about 26. Although temporarily vigorous, VIII and attributeit to a sharpextinctionevent
Ihe Amydrotaxis and Ancyrodelloides stocks just before the end of the Frasnian which, like
were fairly shortJived dead ends,and their dis- the one that terminatedOrdovician CycleIII,
appearance,
alongwith that ofseveralsimilarly is commonlyincludedin the shortlist of major
shortliyed speciesof Icriodus s. 1., produced Paleozoicrnassextinctions.They note that "at
the "extinction event" that broughtCycleVI to this positionalmostall previousspeciesofpala close.
matolepids,ancyrodellids,ancyrognathids,
and
Cycle VII, like its immediate predecessor, polygnathidsbecameextinct." The tabulationI
representsyet anotherepisodein iterative de- presentin Fig. 6.2 alsoshowsa rapid declinein
velopmentof the Ozarkodinida,reinforcedby speciesdiversity in late phasesof Cycle VIII,
burstsof vigorousspeciationin shallow-water but not one that is as abrupt as suggested
by
environments by the lTiodus, Pedavis,and Pe- Ziegler and Lane's comments.Although this
Iekysgnathusstocks.Particularly significant de- disparity may resuli from the fact that I have
velopments frorn Ozarkodina (Spathognath- assembledranges on a biostratigraphic scale
odontidae) were the Eognathodus lineage, that is cruderthan the one usedby Zieglerand
which is confinedprimarily to CycleV and, a Lane, it may also suggestthat the speciesto
bit Iale\ the Polygnalhus stock,which roserap- which they refer disappearedmore gradually in
idly to importance in the offshore envrron- the lateFrasnianthan hasbeenassumed.Ifthis
menls that becamemore widespreadwith the werethe case,conodontsmight provide some
advance of Kaskaskia transgression.In the support for the idea expressedby Johnson,
high-diversity interval of Cycle V, which Klapper,and Sandberg( 1985)that at leastpart
peakedin the Emsianand Eifelian,at least36 of the end-Frasnianmassextinctionofbenthic
conodont speciesprowled Devonian seas-a animalsmight havebeenthe cumulativeresult
sixfold increasefrom the low-diversity interval of threeearlierFrasniandeepeningeventsand
not the result of a single sudden shallowing
of CycleVI.
The high-diversity interval of Cycle VII was event.That is, the drowningofreefsand reducterminatedin early Givetian time by the dis- tion in extentof marginal.shallow-water
enviappearance
ofall but two ofthe lineagesofPo- ronmentsthat followed suchdeepeningevents
cycles
6.3.4 Devonianand Carboruferow
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YJNOCONO]
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9EI
EVOLUTIONARYPATTERNS
bthodontidae),
b Spathognathnar-r/ eYents,
u. tt the Polygmisian high-di
It more than 50
fted at a hUh
im by developf earl;-'speciesof
r prioniodinide
lnl short-lived
',futtrognathus,
Staurognathus).
fr rhe conspict the evolutionte Visean and
rfosippian) segcorresponds in
bse of Sloss's
t- The end of
imus declinein
Ie primarily to
ination in la1e
ryhodus, PseuEl stocksas the
lch they were
tea- However,
G a-ndincrease
Erzed by much
rtle salinity and
q)porrumty to
ih and priond and thereby
rr1 in specieslbdiversity in,lbgelgnathus,
erolved from
tte innovative
iat a\d. VogelI this cycle,but
lrir ed through
hanian.
pecies-leveldiion of Cycle XI
ment of Cayusdinction in the
dus. Lochriea,
os-el.er,during
dnian) interval
C!'cleXII, pop-
137
ulations of idiognathodontid conodonts de- also provide in Fig. 6.3 a zone-by-zone
log of
scendedfrom highly variableGnathodusgirtyi the ratio of origination to extinction of conogradually segregated
into discreteand some- dont species.That log may be used (with the
what lessvariablestocksidentifiedas Decllno- customarycaveats)as a crude index to temgnathodus, Idiognathoides, Neognathodus, poral oscillationsin eyolutionaryvigor.
Idiognathodus,and'streptognarhodus.
RadiaAlthough at least lg5 conodontsfeciesorigtion ofthesestockskept the evolutionaryindex inatedin the permian-Triassicinterval,speciis
above the maintenancelevel of 1.0 through diversity hoveredaround l0 at any one time,
mostofthe Morrowanand promoteda slowre- dropping to a low of 5 in the earliestTriassic
coveryin species-level
diversity,which peaked but reboundingrapidly in the sameinterval to
in the succeeding
Atokan epochwith addition a post-Mississippia
nhigh of 22.
of the first representatives
of Diplognathodus Fig. 6.3 also ouflines four second-ordercyand the Gondolellidaeand gradual phyletic cles in the permian and another four in the
evolutionofthe/ry'eognathodusltneage.
Triassic.
Extinction of Neognathodus and probably
The earliest of the Permian cycles,Cycle
alsoof Rhachistognathusin tl|'elate Desmoine- XIV in Fig. 6.3,followeda long interval in the
sian (Pennsylvanian)markedthe end of Cycle late Pennsylvanianand Asselianwhenthe evoXII, which is also shownin Fig. 6.2 to be ter- lutionary index was consistentlybelow1.0and
minated by a rapid declinein species-level
di- was provoked by the appearancein the Sakversity. Continuedgradualphyletic speciation marian of an early speciesof Neogondolella(N.
in the I di ognathodus-St rept ognathodus lineage, bisselli), the first speciesof Sweetognathus(5.
togetherwith the appeaftnce of Aethotaxis, EI- merrilli), and a distinctive speciesof DrploIisonia, and new speciesof Gondolella, Diplo- gnathodus (D- sakmariensis). N. bisselli is
gnathodus, and Hindeodus, signaleda brief in- thought to have evolved ftom N. praebisselli,
terval of innovation in the eady part of the which precedesit stratigraphically.However,
mid-PennsylvanianMissourianEpochand the thesetwo speciesare separatedby much ofthe
somewhatlater peak in species-level
diversity Pennsylvanianfrom N. clarki, the earliest spethat marked culmination of Cycle XIII. Note cies with elementsmorphologicallyassignable
in Figs. 6.2 and 6.3 that in the latter half of to Neogondolella.For this reason,I suspectthat
CycleXIII the evolutionaryindex was contin- the N. praebisselli-N.bissellilineageis an inuouslybelow1.0and that speciesleveldiversity dependent Permian development from the
declinedgraduallyto 15in the Virgilian and to long-ranging Xaniogna.thus-Cypridodellastock
l0 in the succeeding
Assel.ian
Stageofthe Early which, as I speculatedin Section 5.8.5, was
Permian. Although speciesof Hindeodus, Di- probably the progenitor of several Gondolella
plognathodus, Streptognathodus, Idiognatho- lineagesin the Pennsylvanianand earliest
dus, Adetognathus, Gondolella, and.E llisonia Permianand the ancestor,aswell.ofnumerous
survivedthroughmuch or all ofthis long inter- additional lineagesof Neogondolella, Neospaval ofsorely restrictedevolutionaryvigor, only thodus, and, Epigondolella later on in the
the Hindeodus, Diplognathodus, gondolellid, Triassic.
and Ellisonia lineagessurvived Clark's (1972)
SecondaryCycle XV follows the early ArtinEarly Permian crisis to provide the basis for skianextinction of the typically Carboniferous
secondarycyclesin later Permianand Triassic genera Streptognathodus, I diognathodus, and
seas.
Adetognathus. Following a brief interval of
slightly lowered diversity, a short innovative
burst
resulted in the appearanceof the earliest
6.3.5 Permianand Triassiccycles
speciesof .ly'eostrep
tognathodus,Sweetina, and
Fig. 6.3 is a zone-by-zonesummary of con- Merrillina. Neostreptognathodus apparently
odont speciesdiversity through the Permian representsa modification of the Diplogndthoand Triassicperiods,which togetherrepresent dus-Sweetognalhus
stock, whereasI have inter78 million yearsof geologictime. As in sum- preted,Sweetinaand.Merrillina jn Chapter 5 as
mary logs for earlier parts of the Paleozoic,I successive
stagesin a prioniodinidelineagethat
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EVOLUTIONARY PATTERNS
d almoslincesE of secondary
b to -Anisiandenia and perhaps
Ents elsewhere
l Chirodella and
l_r very tiny eleim (or part o0
I Xaniognathustine, several
the type, M
frg
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rily- becauseno
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Dd phyletic exrc evolutionary
Ioq'ewer, Clark
r39
)r
I
=
z
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o
x
I
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{
\
x\
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400
75
Fig. 6.4. Survivorship curves(left figure)for cohortsofconodont genera(e.g.,all generaoriginaling in Ordovician)
and curve oflongevity ofconodont generaofall ages(on right). Redrawn from Clark (1983).
(1983,1987)hasrecentlyfocusedparticularattention on events in the late Paleozoicand
eady Mesozoichistory ofthe phylum that precededits extinctionat the end of the Triassic.
In his 1983study Clark presenteda seriesof
survivorship curves (Fig. 6.4) for Cambrian
through Triassic cohorts of conodont genera
and a summary curve for all conodont genera
listed in the Treatise(Clark el al., t98l). Clark
noted that thesecurves indicate that the ayeragesurvival of systemiccohorts was 109 my
and that meansurvival ofconodontgenerawas
about 30 my. Thus, as Clark pointed out, an
observerin the late Paleozoicwith accessto the
samerecordswould not have foreseenthe endTriassic extinction of the phylum but would
probably have predicted surviyal into at least
the Jurassic and possibly into the Early
Cretaceous.
The record indicates that the Conodonta did
not survive into the Jurassic or Cretaceous,
however, but that the phylum becameextinct
in the latest Triassic. Conodonts were not the
only groupto becomeextinctor to suffermajor
declinesin diyersity at this time. Indeed,Sepkoski and Raup (1986),summarizingthe opinions ofothers,note that "the I-ateTriassiccontains one ofthe five largestmassextinctionsof
the Phanerozoicmarine record." The data they
present, however, like those for conodonts
shown graphicallyin Fig. 6.5, show generally
high extinction intensities throughout the Late
Triassic,with a maximum in the latestNorian
(or Rhaetic).This suggests
to me, as it apparently did to Clark (1983),that the end-Triassic
extinctionofconodontsmay havebeenthe cumulative result of severalfactors.not of a single catastrophiceventthat terminatedthe phylum "well beforeits time."
Farly in the Permian,at the endofsecondary
CycleXIII, a majority ofthe typically Carboniferous stocks disappeared,following a long
late Pennsylvanian and early Permian interval
during which species-level diversity plummetedand the evolutionaryindex was consistenfly below the maintenancelevel. Permian
recoyery was surely not spectacular.It primarily involved a coupleofbursts of speciationby
the gondolellid stem group, one eachby lhe EIIisonia and.the Diplognathodusstocks,but very
little action on the part of Hindeodus, which
merely "showed the flag" through most of the
Permian.Note in Fig. 6.5, however,that, despite these fitful attempts to restock Permian
environmentswith new and vigorously successfulconodont stocks,extinction intensity
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tc
r smple number of
rin eachzone,and
I d|Iough the Early
lG plo$ per capita
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rditions proba: Triassicby a
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t4 I
groupsof conodonts,particularlythe Ozarko- gin iterative deyelopmentfrom basic lineages
dinida and Prioniodinida.A similar patternis is also an attractive explanation.Plectodina
apparent,however,in at leastsegmentsof the and.Phragmodus,for example, appearde novo
Panderodontidaand Prioniodontida, and it in the early Middle Ordovician, and their spemay be generalin the Conodontaasa whole.
cies forrned skeletal apparatusesthat are, in a
As shown diagrammaticallyin Fig. 5.36, nurnberof ways,closelysimilar to thosebuilt
which summarizesmy viewson the phylogeny by severalspeciesin the Multioistodontidae
of the Ozarkodinida,elaborationof the sides (e.g.,Paraprionioduscostotus\.Becausethe latand surfacesofPa-elementcrownswascarried ter precedePhragmodus and,Pleclodina slratiout repetitively in spathognathodontid
popu- graphically,they havebeenconsidered(by me,
lationsat severalwidely spacedintervalsin the among olhers) as attractiye candidatesfor
Silurian and Deyonian.As a consequence,
ap- ancestors. However, multioistodontid eleparatusesofthe speciesthat developedin each mentsare mostly hyaline,and thereis little to
ofthese iterativeburstsare closelysimilar. For recommendthem in other morphologicparticexample, Ancyrodelloides of the early Devo- ulars as hallmarks of the ancestorsof either
nian has Pa and Pb elementsthat are very sim- Phragmodusor Plectodina. I now suspectthat
ilar to those of early Silurian Pterospathodus those two lineagesdevelopedindependently,
and also essentiallyidenticalin morphologyto and at slightlydiferent times,from generalized
those of later Devonian Ancyrodella. The spe- prioniodontide ancestors in Tripodus (Oixocies of early Siltian Kockelella experimented dontidae).Although one may distinguishreadwith elaborationof Pa elementsin much the ily betweenrnany of the elementsin apparasamemanneras did early DevonianEognath- tuses of Phragmodus and, Plectodina, others
odus, pitnitive representatives of somewhat (includingcertainPa elements)are closelysimyounger Polygnathus, and, a number of later ilar. Thus homeomorphymay not be as great
Devonian and early Carboniferousspeciesas- as between Ancyrodelloides and,Ancyrodella,
signedto Scaphignathus(Spathognathodonti- but it is nevenheless
substantial.
dae)and the Cavusgnathidae.
And iterativedeI suspectthat iteration alsocharacterized
Sivelopment of the Protognathodus-Gnalhodus,lurian developmentofthe prioniodontidefamDiplognathodus, Lochriea, and,Rhachistogna- ily Distomodontidae,but this may be very dif/lius lineages from B ispathodus (Spathognath- ficult to establishobjectively,and there are
odontidae)producedsuch a welter of species surely other possibilities,as indicated in Secwith morphologicallysimilar Pa elementsthat tion 5.7.5.Ifthe basicapparatuspatternofthe
phylogeniesare notoriouslydiffi- distomodontsincludedunplatformedpastinate
supraspecific
cult to reconstruct or defend.
elementsin the P positions,then iterative deIn Section5.6I havealsonotedmy suspicion velopmentmay have beenresponsiblefor in!
that Parabelodina denticulata, a panderodon- tiation of stockssuch as thoserepresentedby
tide specieswith well-developed rastrate ele- Icriodina, Hadrognathus and, ultimately, even
ments in its skeletalapparatus(and thus simi- some or all of the icriodonts.Chattertonand
lar to speciesof Belodina and,Pseudobelodina), Peny (1977)seemalso to have suggested
that
evolved directly from a conlemporaryspecies iteration might have been the active pattern in
of Panderodus(P.bergstroem), not from some icriodont evolution when they wrote that
earlier speciesof Belodina or Pseudobelodina.
Other evidence,mostly unpublished,suggests speciesthat could be or have beenassignedto the genus
that Panderodusitselfincludes a number oflin- Pelekysgnathus, as it is broadly conceived, may have
eages,eachprobablydeservingof independent been derived from other genera during four diferent
generic(or even familial) designationand rep- evolutionary events(origin of"P." inder in the Late Silurian, origin of I hadnagyi in the Lochkovian, origin
resenting chronologically distinct iterations of of
lsteptotaxis'l Iumishi group in the Eady Devonian
stocks characterized by skeletal elements of and.oigin of "P." comn r/rrs in the late Devonian.
closelysimilar morphology.
Within the Prioniodontida rhere are quite a The recent suggestionby Sandbergand Dreenumber of"cryptogenic" stocksfor whoseori- sen(1984)that a groupof LateDevonianspe-
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VINOCONOJ:IHI
zbI
EVOLUTIONARYPATTERNS
bletla lireages.
qponed by furd phylogenyof
bsic overhauls.
nmarized comI to each of the
flo*1y evolving
rans of conserEi from which
'fud-ended linb closerelationI irErative bursts
sive-regressive)
ing of major ina provided by
tepening cycles
k for iterative
r lineage of the
:reas shallowing
' ptases of those
mnities for elabrc-Icriodus and
r Late Paleozoic
ment of gondo: a.ndof ellisony indicate a simEtic events and
r-Yof the Prionof responsibility
iral extinction of
143
6.6.2 Apparatusreduction
by the fact that there is a stratigraphic reduction in the ratio betweenramiform and pectinIn species of a few Ordovician genera iform elementsin severalmajor groups (Fig.
(e.8, Eoplacognathus,Polyplacognathus, Scy- 6.1).
phiodus) and in many post-Ordovicianones,
Merill and Powell (1980) suggestthat elethere appearsto have been a concened,if only mentsin the S and M positionsmay havebeen
superficial, effort to undo the hard work of mineralized early in the ontogeny of certain
rnorphologic diferentiation carried out by their species,
that P elementsmay not havebeenmipredecessors.That is, speciesin these stocks neralized until later, and that the early-formed
apparently formed apparatusesthat were re- S and M elements may have been partly or
duced just to P elements,or apparatusesin wholly resorbed by maturity. This intriguing
which elementsin the P positionswerethe only hypothesis is supported by evidence from the
onesmineralized.This conclusionis suggested Drum Limestoneof Kansas,in which succesFig. 6.6. Frly diversification of the two basic corodont lineages,the Proconodontida on the right, the protopand€rodontidaon the left. Numbers in circlesindicate number ofelement$ in appamtus.Note tha,earliest members
of both lineageshad only a single element type, but that dive.sification produced appantuseswith two to four
element lypesby early in the Ordovician. Redrawn from Sweet0985).
z
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nnl
AHJ
EVOLUTIONARY PATTERNS
...
im rtrrough Triasmany post-Sigroups.Also,
nic
from those
a.lions.
-rent
Ents in which
to go around.
rirely, by P elephosphate-poor
ineralization of
6 by resorbing
dored in those
145
specieswith platformedpectiniform elements tial ecologic controls in the materials they
in the Pa position.The Prioniodinidadelayed study.
developmentof platformed P elementsuntil
The large, stmtigraphicallywell-controlled
the Mississippian,when the curious, short- collectionsayailableto studentsof conodonts
lived Bactrognathidaeinvented thern. Later, should also be well suited to studiesof develhowever, in the Pennsylvanian(and increas- opmental, or evolutionary, strategy.Because
ingly in the Permianand Triassic),prioniodi- well-madecollectionscommonly include elenide stocks came to be characterizedby plat- ments that representvarious stagesin the onformed pectiniform elements in the Pa togenetichistoryofthe speciesthat contributed
position.
to thosecollections,it shouldbe possibleto reIn the absenceof firm information as to the late ontogenetictiming ofthe morphologicfeafunctionofelementsin any positionin the con- tures recorded to developmenlal strategiesin
odont apparatusit is diftcult to relateevolu- the lineagesthat formed them. And, if collectionary elaborationofP elementsto specificen- tions are carefullyrelatedto the petrologicand
vironmental or behavioral factors. However, other biologic properties of the rocks from
from the fact that such elaborationcharacter- which they vvereobtained,it should also be
ized someor all of the lineagesin nearlyevery possible,in time, to determinethe natureand
major stockat onetime or anotherin their his- extent of any relationship betweendeyeloptory, it may be concluded that platform elabo- mental strategy and environmental paration was surely of positive adaptive sig- rameters.
nificance.
Althoughinformation on the ontogeneticdeNicoll (1987),who suggests
tllat the skeletal velopment of various skeletal elementshas
elementsofconodontsmay havebeeninternal been provided by various authors,I am not
supports for ciliated tissue in the food-gather- awarethat anyonehas yet documentedstages
ing systemof generallymicrophagousanimals, in the ontogenyof a conodont speciesfrorn
concludesthat P elementswere opposedand studiesthat involve all elementsof the skelelal
operatedin sucha fashionasto roll, bruise,and apparatus.Consequently,in the following seccrushparticulatematter caughtbetweenthem. tions I summarizepertinentinformation from
Phylogeneticelaborationof P elementsmight a few recentstudiesofPa elementsthat sugg€st
thus have beenrelatedto changeswith time in that developmental strategies may have difsize,shape,and bruise-or cmshabilityof food fered greatly from one group of conodonts to
particles.If P elementsfunctionedas teeth,as the next. We can only guessat the extent to
Jeppsson(1979, 1980)and othersconclude,a which additional strategiesmay be suggested
similar relationshipmight alsobe postulated. by future studies of complete apparatuses.
I P positions
t ard developirus lineagesa
boration of peciions. In a maritions in even
, were occupied
ments; in only
dae. Polyplacorments formed
ian. platformed
rdl-v in P posiE Spathognath.nd Icriodelli
later in the
locks included
6.7 DevelopmentalStsategies
6.7.1 Recapitulqtion
Gould (1977)hasprovidedsignalserviceto the
biologic community by emphasizingnot only
that "parallels between ontogeny and phylogeny are produced by heterochrony" but also
that theremay be a continuumofrelationships
between ecologic factors and the life-history
patterns that may result from alterations in the
timing by which various somatic and beharrioral features develop. In short, Gould expandedon Van Valen's(1973)somewhatearlier observation that eyolution might well be
the control of development by ecology.Paleobiologists are thus encouragedto seek informatioD on developmentalpattemsand poten-
Heterochrony involves "changesin the relative
time ofappearanceand rate ofdeyelopment for
characters already present in ancestors"
(Gould, 1977)and may be effectedeirherby accelerationor retardation.Ifa characterthat appearsin the adult stageofan ancestorappears
at a youngerstagein developmentofa descendant, accelerationis the processinvolved and
classicalrecapitulationthe result. Ii on the
other hand, a character that appears in the
youngstageofan ancestorappearsin, or is retainedinto, the adult stagesofa descendant,
retardation is the processand paedomorphosis
the result.
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V.LNOCONOJ !IHJ
9br
EVOLUTIONARY PATTERNS
E fPrioniodonir (Prioniodonl:ted by paeDren!, of the
ar While steps
|P Postulateare
ttive evidence,
I Inmary anengts of Pedavis
sbfiantial difG ln other api Section5.7.6,
a. Icriodus may
m Pelekysgnadubius,
-dina
distantly re4Elities are conbern involved.
lat the Pa elespeciesof Epr:rain juvenile
Er ontogenetic
d this paedoh-v. In a more
El more extenI has substantiirr at least two
r relatedvein, I
ebeeninvolved
dh from Pseut of Misikella
fulel Ia or those
il ancestorsin
d of the Conlhat of a classirhich therewas
loration of the
dablishment of
the subsequent
i:al fluctuations
des-level diverMer
eustatic
t a one-for-one
ts conserYatlve
@es of repeatrgh iterative,or
repetitive, development of generally shortlived lineages that were strongly homeomorphic with their predecessorsin featuresof
their skeletalapparatuses.
From the l-atePennsylvanian on, however, speciesJeveldiversity
was low, and populationsappearto have been
at or below the maintenancelevel for long periods of time. With only a brief interruption in
the Early Triassic, extinction intensity increasedfrom the Early Permian to the end of
the Triassic,when the phylum wasreducedto
a few species,probably representedby populations of small sizeand provincial distribution.
Although studiesof generic-levelsurvivorship
would not have led Permian observersto predict the extinction of conodontsbefore some
time in the Jurassic,the phylum did not survive the Triassic.Reasonsfor extinction are
not at all clearbut may haveincludednot only
thosebuilt into the long post-Iate Pennsylvanian decline,but also a dollop of bad luck in
the Late Triassic when surviving basinal populationsmay havebeenunableto adaptto conditions producedby a worldwide drop in sea
leYel.
References
Bergstritm, S. M. (1983). Biogeography,evolutionary relationships, and biostratigraphic significance of Ordovician platform conodonts.
Fossilsand Strata 15, 35-58.
Broadhead,T. W. and McComb, R. (1983). Paedomorphosis in the conodont family lcriodontidae and the evolutiolJ of lcriodus. Fossilsand
Strq.ta15, 149-154.
Chattenon, B. D. 8., and Perry, D. G. (1977).
Lochkovian trilobites and conodonts from
northwestern Canada. I Paleont. 51, 7'72796.
Clark, D. L. (1983). Extinction of conodonts."/.
Paleont.:57, 652-661.
(1987).Conodonts:The final fifty million
years. Pp. 165-114 Palaeobiology of Conodonts (ed. R. J. Aldridge). Ellis Horwood,
Chichester,180 pp.
Clark, D. L., Sweet, W. C., Bergstriim, S. M.,
Klapper, G., Austin, R. L., Rhodes, F. H. T.,
Miiller, K. 1., Zieg)er, W., Lirdstrijm, M.,
Miller, J. F., and Harris, A. G. (19E1).Conodonta. In Treotise on Invertebrate Paleontolof? (ed. R. A. Robison). Pt. W, Suppl. 2. Geol.
Soc. America and Univ. Kansas,202 pp.
t47
Gould, S. J. (1977).Ontogenyand Phylogeny-The
BelkDap Press of Harvard Univ. Press,Cambridgeand tondon, 501pp.
Jeppsson,L. (19'19).Conodont element function.
Lethqia 12, 153-l'll.
(1980). Function of the conodont elernents. Lahaia 13, 228.
Johnson,J. G., Klapper, G., and Sandberg,C. A.
(1985).Devonian eustalic fluctuationsin Eurameica. Bull. Geol.Soc.Am.96,567-587.
Lipps. J. ( l98l ). What. if anlthing. is micropaleo\toloqy? Paleobiol. 7(2\, 167-199.
Merrill, G. K., and Powell, R. J. (1980). Paleobiology of juvenile (nepionic?)conodontsfrom
the Drum Limestone(Pennsylvanian,Missourian-Kansas City area) and its bearing on
apparatus ontogeny. J. Pqleont. 54, 1059L0'74.
Mosher, L. C. (1970). New conodont speciesas
Triassic guide fossils.J. Paleont. 44,'137-742.
Nicoll, R. S. (1987).Form and function ofthe Pa
element in the conodont animal. Po. 78-90 in
Palaeobiologyof Conodonts (ed. R. J-.Aldridge).
Ellis Horwood, Chichester,180 pp.
Orchard, M. J. (1983). Epigondolella popllations
and their phylogenyand zonation in the Upper
Triassic.Fossl/sand Strata 15, l'77-192.
Sandberg,C. A., and Dreesen,R. (1984).Late Devonian icriodontid biofacies models and alternate shallow-water conodont zonation. Geol
Soc.Am. Spec.Paper 196, 143-178.
Sepkoski,J. J., Jr., and Raup, D. M. (1986). Periodicity in marine extinction events.Pp. 3-36
in Dynamics of Extinction (ed. D. K. Ellion).
Wiley, New York, 294 pp.
Sloss,L. L. (1963). Sequencesin the uatonic interior of Nonh Ameica. Bull. Geol. Soc. Am.
74(2),93-t14.
Sweet, W. C. (1970). Uppermost Permian and
Lower Triassicconodontsofth€ Salt Ranseand
Trans-lndus ranges.west Pakjstan. Ppi 207275 h Stratigrephic boundary problems, Permian and Triassic of llest Pakislqn (ed. B. K.ummel and C. Teichert). Univ. Kansas, Dept.
Geol., Spec.Publ. 4.
(1985).Conodonts:Thosefascinatinglittle
whatzits."L Paleont. 59(3),485-494.
Vail, P. R., Mitchum, R. M., Jr., and Thompson,
S. III (1977). Seismic stratigraphy and global
changesof sea level, pt. 4: Global cycles of
relative changesofsea level. Pp. 83-95 in,Sedmic Stratigraphy-Applications
to Hydrccarbon Exploration (ed..C. E. Payton). Am. Assoc.
Petrol. Geol.. Mem. 26.
Van Valen, L. (1973). Festschrift. Science 180,
488.
Walliser, O. H. (1983). Geologic processesand
g.lobalevents. Terra Cognita 4,l'l-20.
(1984).Pleadingfor a natural D/C-Boundary. Pp. 241-246 it The Devonian-Carbonifer-
uD 'rrou4ertuel
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.V9I-LiI ,EI
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-ue8tuopouoc u?ruo^oo elpptur ur suJe ed fue
-uo4nlo^ll (6/6I)'A\'JelSerZ pu?')'o8rppeiA
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9t-). Cycles in
Eian to midblaeobiolog of
Ellis Horwood,
7. PALEOECOLOGYANDPALEOBIOGEOGRAPHY
7.1 Introduction
understood as a preamble to the following discussion.Obviously,until we have satisfiedour
curiosity and reachedsome agreementas to
thesematters,conodontswill be lessusefulin
reconstructingconditions in past depositional
environmentsthan are featuresof the rocks
formed in thoseenvironmentsin interpreting
the conodonts.Thus I approacha discussionof
conodontecologycautiously,urith initial emphasis on featureson which there is general
agreement.
For many yearsaftertheir discoveryit wasgenerally held that conodontswere cosmopolilan
in their occurrence within the marine realm
and that most specieswould turn out to be
worldwide in their distribution. However. as
more and nore information was assembled,
it
gadually became clear that conodont species
behavedlike the speciesof most otheranimals.
That is, there were lateral limits to their geographicdistributionand decided,ifbroad, ecologic factors that determined where they lived
within Paleozoic and Triassic seas. Further?.2 Modeof Life. or Habil of Conodonts
more, in 1958 Huckriede suggestedthat the
developmentof Triassic conodontsfollowed It is common knowledge that elements repredifferent courses in various biogeographic sentingthe same or closely similar conodont
provinces,and in 1959Sweet,Turco, Wamer, species may be collected from sedimentary
and Wilkie concluded that the distribution of rocks that representa wide variety of deposi
conodontsin rocksoflate Ordovicianageout- tional environments.In addition, many conlined two distinct faunal provinces.
odont specieshave beenshownto have had a
Neither the ecologynor the biogeographyof very broad geographicdistribution, and a few
conodontscould be studiedin a very meaning- may have beencosmopolitan.Thosefacts,toful way, of course,until a biologicallysound gether with the elongate, wormlike body and
taxonomy had been developedand fleshedout distinctively finned tail of the few complete
systematically.Nevertheless,the repeateddis- specimensknown, support the generallyheld
covery between 1926 and 1966 that conodonts conclusionthat the animalswereprobablynekare exceptionally useful biostratigaphically tic in habit and that the distribution of most
stimulated the assembly of comprehensive, speciesmay have beenlargelyindependentof
geographicallywidespread collections through conditionson the bottom. Suchan interpretathe known range of the group. Thus, with the tion seemsto be supported most strongly by
developmentof the multielementtaxonomy I the common occurrenceof conodonts in cerhaveoutlinedin Chapter5, much of the infor- tain typesof black shalesthat yield fossilsonly
mation neededfor both paleoecologicand pa- ofpelagic organismsbecausethey accumulated
leobiogeogaphic interpretation of conodont under anoxic bottom conditions that inhibited
faunashasfallen into place.
developmentofa benthicfauna.
Becauseconodonts are extinct and have no
Conodonts occur in rocks with corals,brachvery closeanalogsin the living biosphere,fea- iopods,echinoderms,and cephalopods,
whose
tures of their biologic associationsand lithic living representativesare all marine, but they
occurrencesmust be usedto reconstructtheir havenot beenreportedasautochthonouscomhabits, habitats and ecologicpreferences.Infor- ponents of rocks known to have accumulated
mation derived in such indirect waysis always in nonrnarineenvironments.Thus,asat leasta
subject to debate, revision, and periodic rein- first approximation, conodonts may be interterpretation, of course,and this must be firmly pretedas pelagic,probablynektic,marine ani149
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YJ-NOCONOJ IIHJ
0sr
PALEOECOLOGY
AND PALEOBIOGEOGRAPHY
bn from various
rd fiom different
med Pa elements
ts the elements
ir the skeletalapbat this disjunct,
badsto the comn over-represenm late Paleozoic
skeletal apparamt speciesmay
Aom one ontotbat such differ: from a pelagic
retic stagesto a
rin later ones.
b some or all of
E mode of life, it
d study to deterries represented
cttobenthic nibmore at home in
ove the deposirYe had diferent
r itr their life histbs€rvation that,
le water column
tauon, rn a neariphering the life
qreciesbecomes
hl it is a game
lattention to the
of conodontsto
sitional basinas
I arrangement of
mts.
heses,have been
rlbution of condc marine rocks.
bl, outlined in
tan with the ase pelagic and attion for puzzling
iistribution. The
brnes and FihI, attempts to exiF of conodonts
lateral distribu-
++++++++++
+++++++++++
++++++++++
-a-
---
---
a
-OrD
a
-ar-
--
rD
-
'a
--af
'a
r.)
Fig.7.1. Schematicexplanationofthe depth-stratificationmodel ofseddon and Sweet(1971).Diagram showstwo
species;one indicated by crossesis uniformly distributed in a surficialwater stratum; the other, indicated by filled
stratum. Elementsofboth sp€cieswould accumulatelogetheron
ellipses,is uniformly distributed in a deepe!-wa1er
the seafloor,butbetweenA andB thoseofthe "cross" specieswould dominate,whereasbetweenB and C the reverse
would be tlue. Redr-awn,with omissions,from Seddonand Sweet(1971).
tion in sedimentaryrocks.To Bamesand Fahbioraeus,the existenceof laterallysegregated
facies was taken to indicate that most
conodontswerebenthicor nektobenthicorganisms, whereasSeddonand Sweetfound that
laterally segregatedbiofacies (which are real)
required a special explanation if conodonts
were pelagic. Although the two models seem
very different-and they certainly began with
very diferent premises-neither has been generally acceptedor rejected.They both bear
summarydiscussionbeforewe go on to more
concretematters.
7.3.1 The depth-strattficationmodel
Seddonand Sweet(1971)pointed out that the
Ordovician and Devonian conodonts they
were studying in eastem North America and
Australia, respectively, characterized distinct,
laterally segregatedbiofacies instead of being
more or less uniform in their lateral distribution, as one might expect the remains of truly
pelagicorganismsto be. To explainlateralsegregation of conodontsin the benthic record,
Seddon and Sv,/eetproposed that, in their lifetimes, different conodonts inhabited diferent
levels in the sea,in the manner of living chaetognaths.A schematicillustration ofthe depthstratificationmodel is given in Fig. 7.l.
Figure 7.1 considersjust two pelagic conodont species,one representedby crosses,the
other by filled ellipses.The crossspeciesis assumed to have been widely and uniformly distributed in a stratum nearerto the surfacethan
the ellipse species,which was also uniformly
distributed, but was confined in its lifetime to
a deeper-u,aterlayer. As individuals of eachof
thesespeciesdied,their heavy,phosphaticskeletal remains would haye settled to the bottom.
We would expect to find fossil representatives
of the crossspecieseverywhereon the bottom.
However, skeletalelementsof the ellipsespecies would accumulateonly seawardof the
level at which the top of its depth zone
"dragged bottom." Consequently,there might
have been a tract of the bottom to the left of
point A in Fig. 7.1 on which no specimensof
the ellipsespeciescameto restand in which the
fossil record was exclusively of the cross species. On the other hand, the ellipsespecies,a
relativelyrare componentof colleclionsmade
from tlre rocksthat accumulatedbeneathpoint
A, would pradually increase in abundance
relative to the cross sp€ciesin samplestaken
fron rocks that were formed at greater and
geater depth. At point B in Fig. 7.1, for example, the two speciesmight be representedby
approximately equal numbers of specimensl
farther to the right, at point C, specim€nsof the
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VJNOCONO]
EHJ
a9I
PALEOECOLOCY
AND PALEOBIOGEOCRAPHY
Fcies wereinterrrized by a very
csseDtiallithofadal apparatuses
Its. The lateralnhesis has been
n of the simplicm of the distribra4 deposits.As
rgh- as Klapper
rn- the laterally
t car in no way
: or nektobenthic
t disfibution are
lals were nektoI of the vagrant
lhculr to explain
distribution of
m occurrenceof
rt shales,which
rr anoxicbottom
beeD hostile to
tinglJ compelling
; to abandonthe
t conodontswere
cnthicand to use
m the mode-ofEcords with the
rt Ecology
c generalizations
I conodonts as a
:ilcludes a numrich it tnay evensrch principles.
nake generalizaX-- I sumrnarize
nine if the conEst any common
of Cincinnati
abundant in the
tEr OrdoYician
lncinnati Region
ia (Fig. 7.2). The
J-f
=r-
f---t---t--,1_
i
-
- :-
7- - !- - - - - - -
153
---:
\
Ldno
- ;' "' t
?--';:,--:-*=!!
::;:
.-_-i-_---i_
_ - ____10cs
r{
ocs:::
t::::::::
4--...
Fig. 7,2, Situation ofthe Cincinnati Region in the Middle and I-ate Ordovician. The Cincinnati RegioDis th€ tristatetract in the centerofthe map. In general,it was part ofa broad carbonateshelfto which terngenousmaterial
was introduced from the southeast and that was bordered on the norlhwest by the distal extremity of a narow,
deep sag(white) that permitted connectionwilh cold, phosphate-richwatersto the south. Approximale positions
of l0'and 20'S paleolatitudesindicated by heavy lines; Iight arrows suggestprevailing-wind direction; hea\rier
arrows in sag suggestwind-driven circulation pattem permitting upwelling of phosphate-rich water in Cincinnati
Region.Adapted from Crcssman(1973).
speciesrepresentedare quite well known taxonomically, and their stratigraphic and geographic distribution has been worked out in
considerabledetail. Somewhatlessdetailedinformation is availableon the petrologyof Ordovician rocks in the Cincinnati Region,and
questionsaboutthe
thereare someunanswered
environments in which they accumulated.
Nevertheless,
enoughis known to supportgeneralizationsabout at least some of the factors
that may have controlled distribution of the
conodontsin the warm, low-latitudeseasthat
covered this area in the Middle and Late
Ordovician.
The Middle and Upper Ordovicianrocksof
the Cincinnati Region accumulatedon a cra-
tonic platform that was situated a few to as
many as 20 degreessouth of the Ordovician
equator. Lithofacies are various mixtures of
shalederived from orogenicsourcesfar to the
east and highly fossiliferouscarbonaterocks
generatedon the spot, primarily from skeletal
matter furnished by echinoderms, brachiopods, and bryozoans and reworked and fragmented in some environmentsby wave- or
storm-generated
currents.Fossilsin mostofthe
carbonaterocks haye been transported,but
probably not too far frorn their living sites.
The conceptually straightforward deposi
tional frameworkofthe CincinnatiRegionwas
complicatedby differential uplift and subsidenceof segmentsof the platform along pre-
orIJ
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VJNOCONOJ
3HJ
b9l
PALEOECOLOGY AND PALEOBIOGEOGRAPHY
t ddepositional enEeristic conodonts,
d below.
ryesent Phrag'r few are assign' dozen species
E shallow end of
Eel,! glauconitic
hite, with ripr the upper part,
bial corals.The
rr deposit that
tl shallowly subr ma-v have en! on the Drakes
fumed in water
fflatfleld, 1968).
lla-kes are quite
a least half the
erable to RhipEeas the other
Hognathus and.
r rn rhe Drakes,
&ost any speHorician of the
p here and there
rllemes marked
ler and Drakes
nops of facies,
f the conodonts
155
they contain but are mapped as differently dominated.by Phragmodusundatu4 have only
named formations or membersbecausethey minor nurnbersof elementsassignable
to Oulare composedof carbonatesand shalesin dif- odus or Aphelognathus,and,never yield speciferentproportionsor because
they developpar- mens of Rhipidognathus- ln Clays Ferry and
ticularly conspicuousinternal structures.
Fairview strata, on the other hand.,PhragmoIn the late Middle Ordovician, before the ar- dus, Plectodina, Oulodus, and,Aphelognathus
rival in the Cincinnati Region of much terri- are almost equally represenled,specimensare
genousdetritusfrom the southeast,phosphatic characteristically larger, and total collections
limestonesassigned
to the Grier Memberofthe are smaller. Furthermore,Rhipidognathusis
kxington Limestonewerethe chief depositin represenledby an occasionalspecimenin samoxygenated,
well-lit watersomewhat,to consid- ples from the Clays Ferry (but not from the
erably,shallowerthan that in which the Logana Fairview).
Member was deposited.Cressman(1973)esti
The CallowayCreek facies,entirely of Late
matesthal the Grier accumulatedin water less Ordovician age, accumulated in probably
than l5 m deep, but the range may have been somewhatdeeper-water
settingsthan the mudmuch greaterthan this for at the basethe Grier flat environmentsin which the Drakes Forgrades both laterally and vertically into the mation was deposited,and in quieter-water
deeper-,quieter-water l-ogana and, at the top environmentsthan thosein which the conspicinto the Brannon or Tanglewoodmembers, uously cross-beddedTanglewoodMember of
which formed in very shallow, more highly ag- the Lexington Limestone was deposited. In
itated water. Furthermore, the lower Grier both the Calloway Creek and Tanglewood,
shows a dominance by Phragmodusundatus Oulodus and.Aphelognathusare the dominant
nearly equal to that of the I-ogana, whereas conodonts; Plectodina is common, and PhragPlectodina dominates in the upper Grier. The modus is reducedto about 25 percentof the
high phosphatic content of the Grier, a matter fauna.Specimensof Rhipidognathusmay conof specialinterestto Bourbon whiskeydirtill- stitute as much as 5 percent of the collection
ers, suggeststhat water above Grier deposi- from someunits. The Grant Lakeand Ashtock
tional sites was supplied by upwelling with formations were deposited,in Late Ordovician
more phosphatethan could be abstractedby time, at various sites slightly seaward of the
plants.This addsto the suspicionthal at least Drakes flats; the Grant Lake representswavesome of the carbonaterocks now included in agitatedshell-heapshoals,betweenand among
the Grier were depositedbelow 15 m depth, which various faciesof the Ashlock (or rnemperhapsnot too far aboye the compensation bersofthe Grant Lake)weredepositedin sheldepth.
tered lagoons. Grant Lake conodonts are
Late in the Middle Ordovician considerable mostly referable to Plectodina, Oulodus, and.
quantitiesof silt and clay spreadinto the Cin- Aphelognathus,
but specimensof Rhipidognacinnati Region from the southeast.Much of thus arc also common and, locally, may make
this bypassedor was washed through bathy- up as much as 10percentofa collection.Rl,rpmetric highs to settle in thick beds in topo- idognathus may be as important in samples
graphically lower portions of the seafloor (the from the Ashlockas in samplesfrom the very
elongate,southwestward-opening
sagshownin shallow-waterDrakes, but the proportion of
Fig. 7.2). Thus the broad regioncharacterized Plectodinais comrnonly greater.Otherwise,the
in the Middle Ordovicianby carbonatesofthe two formationsresembleone anotherlithologGrier Member was partitioned in later Ordo- ically and faunally.
vician times into deeper-waterareasin which
It shouldalso be noted that the Loganaand
the very shalyKope and lessshalyPoint Pleas- Grier membersof the Lexington Limestone,
ant formations accumulated, and somewhat and the Late Ordovician Kope Formation,
shallowerareasin which the much less shaly grade laterally into dark, fissile shales comClays Ferry and (later on) Faiwiew formations monly loggedby drillers as the "Utica," but is
weredeposited.
nowhere exposedat the surface in the CincinThe Kope and Point Pleasantfaciesareboth nati Region. We have retrieved only a few con-
E p€orq eql Jo
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YJNOCONOJ AHT
9 SI
PALEOECOLOGYAND PALEOBIOGEOGRAPHY
Eton Limestone)
dow the oxygenbt .lmorphognai ttre cool water
I irself.
qFclmens repre'k maximumin
Epreted as the
rimati Region.
rine1on, thought
rr accumulated
qnh. exiibits the
'P. undatusof all
E in Fig. 7.3, I
Fl inhabitantof
r the colderbotE- of the more
tidal-flat areas
fddle and Late
fugnathus, Ould the ozarkodirens of collec4ion lithofacies,
r importance in
rl shallownearincreasein relaf closelyrelated
73 | show PlecladLJ (an almost
ter). as inhabit My of water,
Modus progesa onshoredirecr norcd, wascerrt in tidal-flat
ti Regrbn.I show
re is no compelI data ofthe sort
irn rocks of the
r profound conEn water depth
rolled distribuE west(north in
Eti Region, spenively moreimblomitic Upper
Whus a d Oulbrodontide conmon in stratain
t5 7
.,
;y .4s
/-s','tl';
Fig, 7.4. Location ofthe DeseretBasin and adjacentfeaturesofthe Mississippianof the westemUnited States.
Redrawnand simplified from Sandbergand Gutschick (1984).
northern Wyoming thought to have accumulated at sites marginal to those in which evaporites formed in the Williston Basin (Sweet,
t979). These featuresin the distribution of
thoseconodontssquarewith the onesdeduced
from the record in the Cincinnati Region but
ofer little more as to the actual ecologic
controls.
western
7.4.2 Mississippianpaleoecology,
UnitedStates
Sandbergand Gutschick(1979, 1984)provide
accountsofthe distributionofconodontsin the
Mississippianrocksof westemUtah and adjacent Neyada that are rich in detail and of exceptionalvaluein building up a generalpicture
The rocks in quesof conodont paleoecology.
tion weredepositedon and alongthe westedge
of the broad carbonate platform of the conti-
nental interior, and in the Deseret Starved
Basin, which was separatedfrom the Antler
Flysch Trough on the west by a narrow submarine rise, or sill. The generalarrangementin
the Deseret Basin and adjacent platform areas
of Mississippianrocksof the Anchoralis-Iatus
Zone is shown in Fig. 7.4. From the distribution of conodonts in the several lithofacies
shown in Fig. 7.5, Sandbergand Gutschick
(1984) have also inferred the habitat of conodont speciesthat represent the genera indicated in that figure.
Dark brown to black mudstones,phosphorites, and phosphaticshalesare the deepestwater depositsrecognizedin the Mississippian
transect summarized diagrammatically in Fig.
7.5. Sandbergand Gutschick(1984)conclude
that thesedark-coloredrocks formed in cold,
oxygendeficientmarine water at teast300 m
deep along the axis of the DeseretStarved
Basin.
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The deep-waterdepositsintertongue east- the reconstructedforeslope,including its preward with thicker accumulationsof dolomi- sumably dysaerobic,deep-watertoe. The abtized biosparitethat include encriniteswhose senceof BaclrognarrrJsspecimensin axial detextures proclaim their formation as debris posits of the DeseretBasin, coupledwith its
flows. Theserocks, thought to have accumu- occurrencein samplesfrom both poorly and
lated on the foreslopeof the broad cratonic well-oxygenated
slopeenvironments,suggested
platformjust eastofthe DeseretBasin,proba- to Sandberg and Gutschick (1984) r}|lat Bacbly weredepositedin the murky but well-aer- trognathuswasa swimmer,but that it confined
atedwaterof the aphoticzonelthe encrinites, its swimming to the aeratedwater just basinof course, are built of material that moved ward of the slope floor. In other words, Bacdown the slope from the shelf edgeor from trognathus, like its close relatives, Scaliognasiteshigheron the slopeitself.
thus and.Doliognalhus,was mesopelagic,but
Eastof the platform margin, rocks of the An- its habitat may have huggedthe bottom aboye
choralis-LatusZone arc thick-bedded,light- the oxygen-minimumzone.
colored, bioclastic limestonesthat are replaced
Gnathodus and Pseudopolygnathusare also
farther eastward by more thinly bedded lime- represented
by specimensfrom upperand midstones.These,in turn, gmdelaterally into do- dle slope deposits, but by very few from rocks
lomites. Sandbergand Gutschick infer that the that representslopeenvironmentsinterpreted
latter rocks should merge even farther to the to havebeenbelowthe oxygen-minimurnlevel.
eastwith sandydolomites,perhapswith anhy- Because
of the latter fact,it seemsunlikely that
drite or other evaporites.Thosefacieshave not individuals of either genusever strayedmuch
yet beensampled,however.In general,all these abovethe bottom or out into water abovethe
facies representdeposition in the relatively deepestparts of the basin. In short, speciesof
shallow-waterenvironmentsof the carbonate Gnathodus and. Pseudopolygnathusmay well
platform. All those environments were cer- have been nektobenthic,with habitats that
tainly within the euphoticzone.
"bottomed out" basinwardagainstthe oxygenRepresentatives of Bispathodus utahmsis minimum layer.
Sandbergand Gutschick, 1984, and PolygnaEotaphrus is confined to rocks that formed
Ihus communisBransonand Mebl, 1934,have high on the slopeand in adjacentparts ofthe
beenrecoveredfrom all but the mostnearshore shelf edge; Hindeodus is commonly reprefacies.This near-ubiquityof occurrencesug- sentedin strata depositedon the outer shelf;
geststhat both specieswerepelagicand inhab- Pandorinellina appears to have characterized
ited the euryhaline environmentsof the eu- the inner shelf, in environmentsaboye wave
photic zone.
basefand.Mestognathus,a closerelative ofboth
Scaliognathus and Doliognathus, curiously Pandorinellina and the cavusgnathids, may
specializedand biostratigraphically very useful well have beenadaptedto hlT,ersalinelagoonal
prioniodinidesof the family Bactrognathidae, environmentsalong the innermost margin of
are representedin the deepest-waterdepositsof the platform. Thereis little in the reporteddisthe DeseretBasin as well as in the much shal- tribution of conodontsassignableto any of
lower-waterstrata of the foreslope(Fig. 7.5). these generathat proyides a definitive answer
Such a distribution logically suggeststhat indi to their mode oflife. Their absencein slopeor
viduals of these generawere also pelagic, but deep-basin
depositssuggests
theymay not have
liyed at greater depths than did Bispathodus been petagic, in the manner of Bispathodus
utahmsis andPolygnathuscommunis. In shor1 and Polygnathus. Possibly they were also
speciesof thesetwo generawere probably me- nektobenthic.
sopelagic.Their depth zone may well have
It is always risky to generalizefiom obserbeen defined by the well-oxygenatedbut still vations made in limited areas,no matter how
rnurky conditions of the dys- and aphotic sophisticated
the studiesor convincingthe conzones.
cfusions. However, it may be noled, that PanBactrognathus,a distinctive prioniodinide, is dorinellina is the acknowledgedancestorof the
representedin sarnplestaken from all parts of Cavusgnathidae and. that Cavusgnathus(and,
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PALEOECOLOGYAND PAIEOBIOGEOGRAPHY
T
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5
"0
r the U.S. Midcotr-
in Merrill and
n governed the
rbdon of these
are outlined
r- in comprehenr|s by Heckel
=mann (1975),
E misleadingto
lrtemEnt on all
y of Pennsylvadearly basicdifmatters.Not all
r occasionalstrirent would suglion of informaqrorts proYides
r and habitats of
Fobably also of
E
midcontinental
in a chain of
rere situated in
b equator.This
l6l
areawas borderedon the north and northeast middle limestoneis followed by a meter or so
by the relativelystablecratonicinterior and on of abundantly fossiliferousgray-brownshale,
what were then the south and northwest sides which is typically (but not invariably) introby active rangesoffolded or fault-block moun- ducedby (or includes)a distinctivecomponent
tains. The southernmargin and the southeast- offissile black,phosphaticshale.In this "coreem end of the foreland-basinchain received shale" interval, conodont diversity may drop
great quantities of terigenous detritus from off, but frequency of occurrencecommonly
streamsdraining the Appalachianand Oua- reachesa rnaximum and preservationis genchita-Marathonfoldbelts. This spreadout to erallyoptimum. Representatives
of Gondolella
form a complex of laterally shifting alluvial are almostcompletelyrestrictedin their occurplains and coalescingdeltaic systems,which rence in Kansasand Iowa to the core-shaleinwereonly occasionally(andthen briefly)trans- terval, and specimensof the prionidinide 1d,gressedby offshore marine environments. Far- oprioniodusare more numerousin this facies
ther westand north, the basinchain was more than in any of the othersin the Kansas-Iowa
persistentlyoccupiedby the sea,sedimentsare cyclothem.
offiner grain,and the recordofconodontsand
In the basatpart of the "upper limestone,"
other groups of fossils is more nearly con- which succeeds
the coreshaleofa Kansas-Iowa
tinuous.
cyclothem,maximum conodontdiversityis reIn parts ofthe foreland-basinchain that were establishedand specirnensof the Idiognathomost persistently occupied by the sea, the dus-Streplognathodusplexus continue to domPennsylvanianrock record is made up of a inate in frequency.Adetognathus,represented
successionof cyclothems. These are repeated by few, if any, specimensin the lower part of
setsof distinctive rock types,which record a upperJimestoneintervals, tlpically becomes
number of transgressive-regressive
cycles that more abundantlyrepresentedupward in those
are comrnonly regarded as the results of eu- intervals and may regain a dominance in the
static rise and fall of sealevel. Heckel and Bae- uppermostpart that carriesover into the lower,
semann(1975)and Swade(1985)have related marine part of the superjacentoutside-shale
the distributionand frequencyofconodontsto unit.
the lithic componentsofa numberofthe PennAs indicatedin Fig. 7.7, the typical Kansassylvaniancyclothemsin Iowa and Kansas,with Iowa Pennsylvaniancyclothem is viewed by
the generalresultssummarizedin Fig. 7.7.
mostgeologists
as the recordofa transgressiveNote in Fig. 7.7 that sandy"outside shales" regressiveevent. Thus, from the relationship
at the baseofa typical Kansas-Iowacyclothem betweenconodontsand the lithofacies that repinitially recorddepositionin nonmarineenvi- resenl vanous stagesin thal transgressive-reronmentsbut, in their upper parts, pick up a gressivehistory,it shouldbe possibleto reconfew fossilsthat indicateshallow,nearshorema- struct a general ecologic model. Such a model
rine environments.The first conodontsto ap- hasbeendevelopedby Heckeland Baesemann
pear in theserocks representAdetognathusarLd, (1975)and Swade(1985)and is showndiaEllisonia, but others referable to Hindeodus, gammatically in Fig. 7.8.
Aethotaxis, and. the ldiognat hodus-Strepto- Heckel(1977)concludesthat the phosphatic
gnalhodusplexusfollow shortly. In the "mid- black shalecomponentof the core-shaleinterdle limestone,"just abovethe "outside shale" val accumulatedin anoxic bottom water bein a typical cyclothem, the inyertebrate fauna neath a thermoclineand thus representsthe
becomesmore diverseand is much betterrep- deepest-waterenvironment recordedby any of
rcsented..Adetognathus may continue to be a the lithofacies in a cyclothem of Kansas-Iowa
prominent memberofthe conodontfauna,but type. From this conclusionit followsthat condiversity commonly is greaterthan in shaty odonts commonly representedin core black
rocksbelowanddominancebeginsto be shared shaleswerenot only pelagicbut werealso segwith conodonts of lhe ldiopathodus-Strepto- regatedin depth zones in life. That is, Gozgnathodus plexus.
dolella, Idioprioniodus, Neognathodus, and
In the typical Kansas-Iowacyclothem,the various members of the ldiognathodus-Strep-
.t? topuo9 ': tl
:i i :: ::i i :: :: ::i
-__
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YINOCONOJ AHJ
PALEOECOLOGY
AND PALEOBIOGEOGRAPHY
don tS
i
*"
Si
(aQ
o
.9 *
{
**
o
ll
lr
It
I
I
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
J,a ;l sh,ffi ;
)\
:l-;;.;
* _:
El
/
s k e l e t a lc al c i l ut i t e - c al c a r en r te
(below/above wave base)
rdmembels and
:--===-'-2
-
- t' r a cx
G r e e n - g r e ys h a l e
snal e
D i pl ognathodus
rd if the areal
rf.oed to the
ls of the coret Gondolella's
rd b-vthe cold,
Ine. Depth per
lar controlled
LI.as probably
'tte several pam of the cold,
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YINOCONOJ
:IHT
v9l
PALEOECOLOGYAND PALEOBIOGEOGRAPHY
nd commonly
conglomerates
r maryinal enr black shales
: rnterpretedas
rposits. Heckel
lansgression of
rigenousinflux
r['trt have proNronment unbr- so that the
er-e foundered
@uent deepb offshore,an['fine-grained
I|c matter setrded by superble directlyon
rb situations
Ie explanation
r insistingthat
r inrerpretedin
ized in Section
rencesas to the
t can be made
Jationship beEtary rocks in
:- *'as irnplied
r and lateralrd by Seddon
and Fihraeus
re also learn
Ering distribu;dures of both
hion that the
t have yaried
nhic. On the
i is rarely posrnical, or biotsolled the disr example,that
lhids were, for
lftalrs of shalHts that were
zted temperalinit_v, but it is
icron of those
165
harsh environments controlled the distribu- Idioprioniodus and, Gond.olellain Pennsylvations obserYed.
nian cyclothemsin Kansasand Iowa suggests
that speciesof those generawere confinedin
life to a relatively deep part of the water col7.5.1 Depthasafactor
umn and also that their depth zone may haye
Many of us who have written about conodont been defined by the cold, dense dysaerobic
paleoecology
havediscussed
the distributionof water ofthe thermoclinethoughtto be responvariousspeciesin termsofwater depth,not be- siblefor anoxicconditionson the bottom. Uncausedepth pel Je is a significant ecologic fac- fortunately,from studyofthe skeletalelements
tor, but becausea senseof at least relative of cold-waterOrdovician and Pennsylvanian
depth may be derivedfrom the textures,struc- species,I havebeenunableto isolatemorphotures, and other fossils in the sedimentary logic featuresby which other cold-waterforms
rocks from which our conodont collections might be identified.
havebeenmade.Althoughdepthitself may not
have exerteda primary influenceon conodont
distribution, factors such as temperature, light 7.5.3 Nearshoreand ofshorefaunas
penetration,lieht intensity, turbidity, energy, All threeof the modelssummarizedin section
salinity,and water densityfluctuatedirectly or '1.4relatethe distribution of conodontsto oninverselywith depth,and oneor a combination shore-offshoretransectsreconstructedfrom the
ofthese may haveexertedthe direct control on distribution and nature of the sedimentarydistribution. I suspect,for example,that tem- rock record.In all threemodelsthereis reasonperature,not depth, was the important factor ably clear distinction between an offshore
governingdistribution of speciesof Amorphu group of speciesthat may be relatedto stable
gnathus and, related balognathids.That is, temperatures,"normal" marine salinities,and
those speciesare representedin highlatitude limited turbidity; and anothergroup that apOrdovician rocks that formed in cool, rela- parentlythrived in nearshoreenvironmenls
tively shallow water, whereas at lowlatitude characterized presumably by harsher, more
sitestheir remainsare largelyrestrictedto the varied environmentalconditionssuchas great
depositsof deeperwaters in which tempera- fluctuationsin temperature,salinity, and turtures may have been comparableto those bidity. It is ofinterestthat conodontstypical of
nearerthe surfaceat high latitudes.
the offshoreand nearshoreendsofthe environmental spectrumin all three models have a
numberof featuresin common.
7.5.2 Temperatureas afactor
In all threeofthe situationsdescribedin SecDiferences in temperature were probably re- tion 7.4,speciescharacteristicof nearshoreensponsiblefor development,in the Ordovician, vironmentsbuilt apparatuses
of relativelylarge
ofdistinctive conodontfaunasin the high- and elements, with a reduced nurnber of discrete
low-latitudemarine realms.This conclusionis denticlesof circular crosssectionand little, if
suggestedby the fact that rare invasions into any. white matter.I alsohavethe impression
higher latitudes of speciescharacteristicof low thal elemenlsof nearshorespeciesare more
latitudes were accompanied by accumulation variable in morphologicdetail than those of
of vaughnitic carbonate rocks that indicate offshore species,and their apparatusesare
concurrent invasions of warmer than usual more variable in composition. Furthermore,
water.Also, as noted in Section7.5.1,the few note that Ordovicianand Triassicrocksrepretypically highlatitude speciesthat migrated senting nearshore environments have profrom time to time in the Ordovician into the duced all of the "fibrous" elementson which
low-latituderealm becameestablishedtherein Bransonand Mehl (1944)basedtheir concept
deeper-waterenvironments that were probably of the suborderNeurodontiformes.Finally, it
similar thermally to the high-latitudeonesin is clear that at any time speciesdiversity was
which the species
weremostwidely distributed. considerablylower in nearshorethan in ofAs notedin Section7.4.3,the distribution of shoreenvironmentsand, becauseit is my ex-
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VJNOOONOJ :IHI
991
PALEOECOIOGYAND PALEOBIOGEOGRAPHY
ied denticles
ner. Their dehdovician, apt apparatuses
res" Developinilar history.
i:s are repreI and the eleershore featte primitive
,dina," clearly
mr-rronments.
F ous to use
r elsewherein
iks to the enE OrdoYician
rlusion from
Etioned that,
I paleoecology
rI To be sure,
--<)r eYenthe
I a generalaffie. shallowei'en at this
teep in mind
hioniodinida
ue explorers
mid-Pennsyld nearshore
Ere frrmly esed Ellisoni-
r I notedthat
rs of Triassic
rctively, had
hde that dur:n history the
rts may have
) s.rggestions,
ilerable skepE Eell-estabfrough it has
= h their hismmopolitan
4rh) is cural circlesbe-
causeofthe light it may shedon the positionof
major crustalplatesin the past.For this reason,
a number of defailedstudiesof conodontpaleobiogeographyhave been published in the
lastfew years.Most ofthesehayebeenbrought
up to datefairly recently(€.9.,Clark,ed., 1984),
so I provide only a brief summaryhere.
7.6.1 Late Cambrianand Ordoyicqn
paleobiogeography
t67
ley Province in the late Ordovician of the
wann-water realm. tateral differencesin contemporaneousconodontfaunasalso permitted
recognition of British, Baltoscandic, and Mediterranean provinces in the cold-water marine
realm, in which conodonts may have lived in
subpolarseaswell above 60 degreessouth of
the equator.
I-ate in the Ordovician, as noted in Section
6.3.2,conodontspeciesdiversity declineddramatically, and this seemsto have led, by the
end ofthat period,to virtual eliminationofthe
fauna characteristic of the cold-water realm.
Stocksthat survivedinto the Silurian (i.e..the
Icriodellidae, Prioniodinidae, Spathognathodontidae, and Panderodontidae)are either
onesthat werewell establishedin low- to midlatitudeseasofthe Ordovicianor aresurviving
relatives(e.g.,Distomodontidae)of onesthat
were.
Miller (1984) notes that both classesof conodonts appear to have originated in the Late
Cambrian in warm, low- to midlatitude seas.
By earliestOrdovician time, however,representativesof both groups had spread into
higherJatitude seas,and Cordylodus (a cavidont) had becomeessentiallycosmopolitan.A
pair of major eustaticeventsin this Late Cambrian-earliest Ordovician interval are thought
to hal/e been influential in directing the development of the Conodontaso that, by late in
and Triassic
the Tremadocian, very different faunas had 7.6.2 LaterPaleozoic
paleobiogeography
evolvedin low- and higherlatitudeseas.
The faunaldifferentiationoutlinedby Miller Reported Silurian conodont faunasare surpriscontinued through the Ordovician, when the ingly cosmopolitan.
However.46new species
manne realm was clearly divided into two are described from the Lower and Middle Simajor biogeogaphic units (Sweetand Berg- lurian of New South Walesin a recentreport
strijm, 1974,1984).I now preferto regardthese by Bischof(1986),which may suggestthat Silmajor units as warn- and cold-water realrns, urian cosmopolitanismmay be partly a result
althoughthey were originally describedas the of faulty knowledge of Silurian conodont
North Atlantic and North American Midcon- faunas.It is alsoofinterestto notethat Silurian
tinent provinces.Descendantsof the pioneer conodonts have been collected largely from
lineageof the Conodonti,the Teridontusstock, rocks thought to have been depositedequatorfigured prominently in faunas of both the ward ofthe 40th parallels(Charpentier,1984).
warm- and cold-water realms and, as BergKlapperand Johnson(1980),whosecomprestrdm and I pointed out in 1974,there was a hensive review is authoritative, confirm that
not-surprising parallelism in development. conodontsappearto have beenrestrictedin the
Only a few cosmopolitan species existed, Early and Middle Devonian to more or less
mostly representativesof Drepanoistodus and, tropical areas.Early in the Devonian, endePanderodus,but also including the cavidonts mism was relatiyely high, but later in the peDapsilodus, Walliserodus,ar:.dAnsella.
riod, with increased rnarine transgression of
By the Late Ordovician,and possiblybefore, cratonic areas, endernism decreasedand the
distributionofconodontshad cometo defrnea nurnberof cosmopolitanspeciesincreased.Alnumber of biogeographicprovinces in the though Klapper and Johnsonnoted that most
warm- and cold-water realms. For example, a conodont genera were uniformly distributed,
combinationofR- and Q-modeclusteranalysis they loggeda number of differences,at the speof data from 26 sites in North Arnerica (Sweet ciesleyel,betweenthe conodontfaunasofvarand Bergstrdm, 1984) permitted discrimina- ious epeiric seasof the Early and Middle Detion of an equator-straddling
Red River Prov- vonian.It wastheir choicenot to formalizethe
ince and a soinewhat higherJaiirde Ohio Val- endemicunits asproyincesbecause
sucha pro-
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VJNOOONOJ
IIHJ
89r
PALEOECOLOGY
AND PALEOBIOGEOGRAPHY
lion and one of
r as a feature in
f an1-'age. Sublrd a Tethyan
tDgaea, a Gerilce in central
Pro\-ince on the
:aner's (1984)
I endemism did that Middle
Ecupied by inmt faunas. Be! Triassic have
E specieslevel,
ilentifications)
t role in supprincialism or
bm. Until there
B roday on the
E I doubt that
Eraphic distri-
E (1975).Provpoposed nektoDdontophorids.
d Middle Silulm New South
t Senckenberg
E (1978).Late
aian history of
c, central KenI Soc.Am. Bull.
i. (1934).Coneuti Studies 8,
E-246 h Index
. W. Shimer and
L
odo[ts through
nodoot provinr 196, lL-32.
rt biofaciesand
Aemoir 196, Ltrarigraphy and
' the Lexington
rtral Kentucky.
t68. r-59.
[-v and paleoe| (Cincinnatian)
169
in Indiana, Ohio, and Kentucky. Geol. Soc.
Devonian and Mississippian rocks along the
America Spec.Paper 95, l-34.
Wasatch Front and Cordilleran hingeline.
Heckel, P. H. (19'1'7).Origin of phosphaticblack
Brigham Young Univ. Geol. Studies 26: tO7_
shale facies in Pennsylvanian cyclothems of
133.
mid-continent North America. Am. Assoc.pe- (1984). Distribution, microfauna, and
trol. Geol.Bull. 6l(7), 1045-1068.
source-rock potential of Mississippian De[e
(1980). Paleogeographyof eustatic model
Phosphatic Member of Woodman Formation
for deposition of mid-continent Upper pennsylvanian cyclothems. Pp.l97-215 in Paleozoic
paleogeography of the west-central united
States(ed. T. D. Fouch and E. R. Magathan).
Rocky Mountain Sec.. Soc. Econ. Paleont.
Mineral.
Heckel, P. H., and Baesemann,J. F. (1975). Enton Group (Ordovician) in New york, southern
vironmental interpretation of conodont distriOntario and Quebea.N. Y. State Mus. Sci. Seru.
bution in Upper Pennsylyanian (Missourian)
Bull. 405, l-1O5.
megacyclothemsin easternKarlsas.Am. Assoc.
Petrol. Geol. Bull. 59, 486-509.
Huckriede, R. (1958). Die Conodonten der Medirerranen Trias und ihr stratigraphischerWert.
PaLiont.Z. 32, l4l-175.
Klapper, G., and Johnson, J. G. (1980). Endemism and dispersalofDevonian conodonts.I
Paleont. 54, 400-455.
Geol.Survey,Tech.Information Ser. 14, l-:-l.
Kozur, H. (1976). Paleoecologyof Triassic con- Sweet, W. C. (1979.).Conodonts and conodonr
odonts and its b€aring on multielement taxonbiostratigraphy of post-Tyrone Ordovician
omy. Geol.Assoc.CanadaSpec.Paper 15,313rocks of the Cincinnati Region. U. S Geol.
324.
Sum.Prof,.Paper t066-G, ct--G26.
Merrill, G. K. (1968). Allegheny (Pennsylvanian) (1979). Late Ordovician conodonts and
conodonts. Unpubl. Ph.D. thesis, Louisiana
biostrati$aphy of the western Midcontinent
StateUniv., Baton Rouge, 184 pp.
Proince. Brigham YounR t-lniv. Geol. Studies
Merrill, G. K., and Martin, M. D. (1976). Envi
26,45-85.
ronmental control of conodont distribution in Sweet,W. C., and Bergstriim, S. M. (1974). prothe Bond and Matoon formations (Pennsylvavinciatism exhibited by Ordovician conodont
nian, Missourian), northern lllinois. Geol
faulnas.Soc. Econ. Paleont. Mineral. Spec.publ.
Assoc.Canada Spec.Paper 15,243-27L
2r, 189-202.
Merrill, G. K., and von Bitter, P. H. (1984).Facies (1984). Conodont provinces and biofacies
and frequencies among Pennsylvanian conof the I-ate Ordovician. Geol. Soc. Am. Spec.
odonts; Apparatuses and abundances. Geo,/. Paper I 96, 69-87 .
Soc.Am. Spec.Paper 196,251-261.
Sweet,W. C., Turco, C. A., Warner, E., Jr., and
Miller, J. F. (1984).Cambrian and earliestOrdoWilkie, L. C. (1959).The American Upper Orvrcran conodont evolution, biofacies,and prodovician Standard.I. Eden conodontsftom the
vincialism. Geol.Soc.Am. Spec.Paper 196,43Cincinnati Region of Ohio and Kentucky. "/.
68.
Paleont. 33, 1029-1068.
Nicoll, R. S. (1976). The effect of Late Carbonif- yon Bitter, P. H. (1972). Environmental control
erous-Early Permian glaciation on the distriofconodont distributon in the ShawneeGroup
bution of conodonts in Australia. Geol. Assoc.
(Upper Pennsylvanian) of eastern Kansai.
Canada Spec.Paper 15,273-278.
Pojeta, J., Ir. (1979). The Ordovician paleontology of Kentucky and nearby states-Introduction. U.
^S.Geol. Surv. Prof. Paper 1066, AlA.48.
Rhodes,F. H. T. (1952).A classificationofpennsylvanian conodont assemblages.J. Paleont.
26,886-901.
Sandberg,C. A., and Gutschick,R. c. (1979). graphischeBedeutung.Abh. Hess.Landesamtes
Guide to conodont biostratigraphyof Upper
Bod.enforsch.
3t, 1-l 66.
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THE PHYLUM CONODONTA
bt evidencein
Fosition ofthe
ndonts is prod assemblages
r on the shape
dized elements
r4000speciesis
I discretespecib€ rest is really
tretation.
rScottish specIghtly different
t leasttwo difrions in which
lreserved as a
rith overlying
csecompressed
mains of elonl-bodied organo 1.8mm wide.
qreted as a ced- bilobed,and
iDaturalassemlt are oriented
r to that of the
site end, interI more or less
rs to i blunfly
d setsofshort,
inpressionsinurrounding the
Gterior part of
l secrmen may
ttenorly openracterize what
of ar least the
nens.Although
riately cautious
hrcs with the
ruscleblocks of
kh. Aldridgeet
111
al. (1986)describethesefeaturesas "somites" nonmarine. The first true conodonts are from
without further justification and this evidently Upper Cambrian rocks, the last from strata of
figured prominently in their ultimate determi- latest Triassic age. Conodontshave been renation that the affinitiesof the conodontsare coveredfrom virtually all types ofsedimentary
with the chordates.
rocks. and records of conspecificspecimens
Also indicated with varying degreesof fidel- frorn rocks representingmany diferent depoity on all four Scottishspecimensis a pair of sitional envionmentsare common. Especially
subparallellongitudinal lines, which extend significant(to me, at least)is the fact that most
from a point (in one specimen)not far behind conodont generaare representedsomewherein
the cephaliclobe to a point near the posterior the world in black shale deposits, which comend of the tail. Aldridge et al. (1986)point out monly lack remainsof benthic organismsbut
that theselines might represent". . . a septum in many placesinclude well-preservedspecidividing the body somites longitudinally, a mensof pelagicones.
mesentery separatingcompartments of the
body cavity, a major longitudinalblood vessel,
as Ilvertebrates
a stiffeningrod or notochord,a nerve chord, 8,3 Conodonts
the gut." They note that the gut is the one of In the 130yearssincetheir first formal recogthese structuresmost likely to be preserved; nition, the conodontshave beenreferredat one
however,they alsonote that ifthe pairs oflon- time or anotherand with variousdegreesofsegitudinal lines do representthe outlinesof the nousnessto most ofthe major phyla of invergut, the fact that they extendalmostto the pos- tebrateanimals.The hypotheses
represented
by
terior end ofthe tail in two specimenssuggests many ofthesereferralsareeasilydismissed,but
that the animal lackeda postanalsegment,or someothersare not so readily discountedand
true tail.
should at least be mentioned in this ooinion
As noted in Chapter 2, the intemal structure polt.
of conodont elementsindicatesthey are composed of two parts, crown and base, which
must have been completelyenvelopedby (or 8.3.1 ArthropodandAnnelidconnections
embeddedin) secretorytissue at least during EvenbeforePander'soriginaldiagnosisofcontimesof growh. Apatite crystallitesin lamellae odontswaspublishedin 1856,Murchison,Barof crowns are oriented with their prism sur- rande, and Carpenter had registeredthe opinfacesparallel to the direction of growth. Bases ion that the tiny fossils were the tip-ends of
are less densely mineralized, and the frame- some parts of tlre trilobite carapace,and thus
work of organic material may be thicker. In a were arthropod, not chordate, fragments.And,
few specimens,
the basalportion may includea not long after Pander'smonographappeared,
plug of bonelike material v/ithin which Bar- Owen (1860) suggested
that conodontsmight
skov, Moskalenko,and Starostina(1982)have be analogouswith the ". . . spines,hooklets, or
identified structuresinterpreted as osteoblasts. denticles,of nakedrnolluscsor Annelids."
Among the other charactersthat havo been
Although no one seemsto have followedup
consideredin speculations
on the biologicaffin- on the interpretationof conodontelementsas
ities ofconodonts,I shouldalso mention their parts ofa trilobite carapace,two authors (Hargeographic and stratigraphic distribution and ley, 186l; Simpson,in Newberry, 1875) sugthe cosmopolitanoccurrenceof many species gestedthat conodontelementsmight b€ crusin rocks representing a wide variety of sedi- tacean remains. These suggestionshave not
mentary facies.Conodonts are common indig- proved particularly appealing, largely, I susenousfossilsin rocksthat containthe remains pect, becausethe variety of elementtypes of
of animals such as brachiopodsand echino- most natural assemblages
is not duplicatedin
derms,which are exclusivelyrnarineat present the array of spines that rim crustacean
and apparently always have been. They have carapaces.
been found only as reworked or redeposited Owen'ssuggestionthat conodontsmight be
specimensin rocks that might be interpreted as a group of the Annelida received seriousatten-
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THE PHYLUM CONODONTA
csed the possinight represent
lalso raisedthe
E Edular teeth
I srggestionhas
blogists, but it
rberry, 1875),
5). all of whom
rs morphologic
rents of living
irm conodont
3 the argument
be the radular
: most types of
Dlogic counterf living gastronot found in
mdanl gastro: conodont eleE the radular
rts are entirely
t radular teeth
in sl mmetrical
$ow considerr conodont eleft et at., l98l)
lrt punch,sugEt most living
Fologically to
d henceare untoup to which
Edif (1986)haye
rodonts might
lrn-t featuresof
rcimens invite
f caudofoYeate
rh a comparihm that living
oe size as the
hts: that in
some AplacoIinro two syrnI grooveofliv:nral suture of
itions reministtish conodont
I ttrechevron-
t7 f
shapedimpressions,which are such strikine and mollusks, which have already been disfeaturesof the Scortishspecimens.
mighrcorl cussed,these include the lophophorata (or
respondeither to the dorso-ventralmusclese- tentacutala) and such disparate groups
of
ries of the Solenogastres
or to the chewonlike wormlike animals as various aschelminthes,
arrangementof dermal spiculesthal character- gnathostomulids,and chaetognaths.In fact,
izesthe dorsalsideofother aplacophoranmol- about the only major invertebrate phyla
that
lusks. Accordingto Tiltier and Cuif, features have not beensuggested
as the parentalgroup
interpreted as fin rays in the tail region of two for the conodonts are the Archaeocyathida
Scottishspecimensmight alsobe duplicatedin (which were largely or entirely extinct
before
certarnprochaetodermatids
by very long sp! conodontsappeared),and the porifera, Bryculesin the posteriorregion.
ozoa,Echinodermata,and Hemichordata.
Tillier and Cuif further suggestthat the ceLindstriim (1973) and Conway-Morris
phalicapparatusofthe Scottishconodontsmay (1976) have either directly
or indirectly sugbe analogousin arrangementand errenrn mrn- gested that conodonts might be an extinct
eralogyto the radular teeth and, particularly, group of the invertebrate superphylum termed
the buccalmandiblesofthe Aplacophora.They Irphophorata or Tenkculata by various
aunote,asI havein a previousparagraph,that the thors and composedat presentof the phyla
principal reasonfor rejecting a rnolluscanaffin- Brachiopoda,Bryozoa,
and phoronida. Conity for conodontsis the argumentthat mollusks way-Morris's suggestionis basedon his interare incapable of secretingphosphatic hard pretation of a curious little fossil from
the faparts.However,Tillier and Cuifhave subjected mous Middle Cambrian
Burgess Shale of
the radular teeth and buccal mandibles of British Columbia as a vagrant tophophorate,
Chevrodermaturnerae (a prochaetodermatid and on his interpretationofimpressionsin the
caudofoveate) to both X-ray microdiffraction supposedlophophoreof that oryamsmas conand microprobeanalysisand have discovered odont elements.Unfortunately,the conical
elethat both teethand mandiblesincludesubstan- mentssupposedto havesupportedlobes
ofthe
tial amountsofcalcium phosphate,as is prob- lophophore of the creature Conway-Morris
ably also the casewith the radular teeth of at named.Odontogiphus omalius have all been
least certain scaphopods.(Gastropodradulae dissolvedaway,and the fossilis from rocks
far
studied thus far seemto lack phosphate,and older than thoseyielding the oldestspecimens
the external spiculesof the Aplacophoraare I would regardas conodonts.Thus,in addition
aragonitic.
)
to questioningthe realsignificance
ofa vagrant,
The featuresjust summarizedled Tillier and or swrmming,lophophorate,I disqualifyOdonCuif to commentthat if conodontswereapla- togriphusas a seious contributor to our undercophoranmollusks,it would be normal for the standingof conodontaffinities.
phosphaticbuccalmandibles,at least,to be fosLindstrijm (1973), on the other hand,
silized even though the aragonitic dermal pointed out featuresthat suggestconodont
elespiculesmight recrystallizeand be unrecogniz- ments were internal (not external)structures;
able as conodontelements.They thereforere- noted a generaltendencyin the evolution of
gard the Conodontaas a potentialclassof the conodonB toward an increasein
surfaceareaof
Mollusca, most closelyrelated to one or an- elements;and suggested
that pits in the upper
otherofthe groupsincludedin the paraphyletic surfacesof certain conodont elementsmight
Aplacophora.
have been the sites of muscle attachment.
Theseobservationsled him to suggest
that conodont
elements
might
have
served
as
intemal
E.3-3 Connections
with otherinvertebrates
supportsfor a muscle-operated
tentacularapMiiller (in Clark et al., l98l) lists a largenum- paratus, or lophophore, and this led to the furber ofadditional invertebrategroupsthat have ther suggestion
that
beenconsideredappropriate,for one r€asonor
another,as taxonomicreceptacles
for the con- anceslors of the co[odont stock and those
of the braodonts.In addition to the arthropods,annelids, chiopod stock were closely related at some relatively late
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tL l
THE PHYLUM CONODONTA
frata- (A) BranErioe (RhynchoE {B) atrd(C) re-
3 aITOWWOrInS
d this zoologiE the category
1982) demonr structure bed early Ordor assigned to
tEs of Sagitta
L I s'as partict tbat the conEDic and early
Eognalha bed I had used
thar group in
lin anomalies
dOrdovician
, srrh the apt study, many
lscovered the
rs. however,
t rle Chaetoirbs in distrid structure of
lls
Phakelodusand,Sagia4 elements.I am not c€r_ rmens and
concluded that they were ,,. . . frag_
tain that any oftheseproblemsis fatal,but thev ments of the
hard, crustaceousends of the
all diclate caurion. For example.the graspin! segmentsof
some trilobite.,, In apparentdef_
spinesofall living chaetognaths
areentirelyor_ erence to that opilrion, which he did not at_
ganic in composition, whereasall specirnens tempt
to refute, Pander remarked somewhat
that I regardas skeletalelementsofconodonts wistfully that (my
translation)... . . Complete
are thoroughly phosphatized.Furthermore, al_ conviclion
that theseremainsarereallyteeihof
thoughBengtson(1976)has outlined the steDs extinct fishes
could be reachedonly if similar
by which the transition from organismswiih structures could
be demonstratedin living an_
elementslike those of phakelodusto thosewith imals of the
sarneclass."
elements with the internal structure of all
known conodontsmight havebeenmade,that
transition has not yet been objectivelydocu_ E.4.2 Newberry,Hinde, Huxley and Myxine
mented.Thus it is my perhapsultraconserya_Newberry (1875),
on his own account. and
trye conclusion that phakelodus and other or_ Hinde (18?9).
on the adviceof Huxlev.congamsms with skeletal elements of similar cluded
apparentlyfrom shapeand lusteralone
lntemal structure(the paraconodontidaof the that the
conodont elements they examined
Treatise classjfic.ation)are not conodonts. If were so
closely similar to the teeth of the hae_
that is the case,then identity in structureof frsh,Myxine (Fig.8.2).
as to be indistinsuishaPhakelodus and Sagi a elementshas no bear_ ble. In
fact. Huxley is reported ro have in_
lng on the question of conodont affinities.
althoughit may suggesrthal rootsoflhe Chae_
tognathaare to be found in the paraconodon_ Fig. 8.2.. Myxine glutinosa,
a hagish (o, slime eel),
tida, a goup formerly assignedto the Cono_ reprcsents the chordate division to which many believ;
donta on the basis of gross morphology of conodonts to be mosl closely related. Redrawn from
Miiller (1836).
skeletalelements.
8.4 Conodontsas Chordates
Only a few studentsofconodontshayetakena
firm stand regarding the biologic affinities of
condonls.and most of them favor an assign_
ment rn or nearthephylum Chordata.It is thus
ofinterest to look criticaltyat the evidenceon
which that assignmenthasbeenbased.
8.4.1 Theopinionsof pander
Pander(1856),who coinedthe nameand pro_
vided the first diagnosesand descriptioni of
conodonts,included those diagnosesand de_
scnptrons rn a monogaph that dealt Drimarilv
with fossil 0shes. Throughout rhai pioneei
monograph,Pandertreated conodontsas the
hard parts, possiblythe teeth and jaws, of a
group of extinct fisbes, prirnarily, it seemsto
me becausethat is what they looked like to
him. Panderdid not seriouslyconsiderassign_
rng the conodonts to any group other lhan the
Chordata, but he noted that Murchison, Bar_
rande,and Carpenterhad examinedsomesDec_
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911
THE PHYLUM CONODONTA
ber ditrerent
lErs on con1986).
kr
rpressed the
ah and derI necessarily
osever, that
nrily (if not
riS' in exters the teeth
the teeth of
n frnd no evI of the sublructure deil by Pander.
d basalbony
rt elements
rbrns in the
lone of censome of the
|t tbe baseto
in color and
derm bone.
@inion that
have been
: he was unD include all
r later, Bar1 (1982)deel ptatesof
drs- a genus
dy in the
rs together
rrespond in
I Osreocyres,
tov and his
rE bone and
I remalns.
rred in the
tqts as the
r€d to have
never pre6 Kirk had
I did many
yearslater),that many ofthe elementsthey collected from the Middle Ordovician Hardins
Sandslone
oIcentralColoradowereattached
ai
the baseto a substancethat ".. . appearsbony
but does not have the structure of ordinary
bone." Bransonand Mehl's (1933) comment
that ". . . the materialsto which the conodonts
are attachedcould not be from annelids or
from anlthing but vertebrates..." takes on
considerableimportance becauseit was the
conclusionof a pair of trained vertebratepaleontologists;but it can hardly be acceptedas
evidenceofaffinity in the absence
offurther description, illustration, or histologic analysis.
Possiblythey werereferringto materialsuchas
that describedfrom Siberiaby Barskov,Moskalenko,and Starostina(1982),but this will alwaysbe uncertain.
t't7
Ulrich and Bassleron little evidenceother than
comparativegrossmorphology.
8.4.7 Interpretations
of the Scottish
Carboniferousspeci mens
The most recentdiscussionsofconodont amnities (Briggser al., t983; Aldridge er al., 1986)
are concemedprimarily with appropriate interpretation of the four Scottish Carboniferous
specimens,to which I have alreadymade repeatedreference.Briggs et al. comparedthe
first-discoveredand best-preservedScottish
specimen with the cephalochord.ateBranchiostoma(FiE.8.lA),the ammocoetelarva of lampreys, and arTowworms of the invertebrate
phylum Chaetognarha
(Fig. 8.lC). Their conclusionwasthat neithera chordatenor a chaetognath model ". . . provides a satisfactoryanswerto the questionofconodont amnity."
8.4.6 Compositionand growthof etements
Following description of three additional
Ellison (1944),and Hassand Lindberg (1948) specimensin 1986,however,Aldridge
and his
clearedup long-standingconfusionabout the colleagues
concludedthat conodontsrepresent
chemical composition of conodont elements a group ofjawless,craniatechordates
seDarate
and pointed out that the phosphaticminerals from previouslydescribedgroupsbut perhaps
they identified were the sameas those in fossil rnost closelyrelatedto the hagfishes(Myxinoand recent bones and teeth. However, Gross idea). A similar conclusionis expressed,
but
(1954),a studentoffossil fishes,madethe im- just in passing,by
Jeferies(1986),who notes
portant determinationthat conodontelements that the hypotheticalanimal ..s" proposes
he
as
are not composedof dentineand lack features the first "crown vertebrate,,resembles
the Scotthat could be interpretedas either a purpa or tish Carboniferousconodontanimal
described
dentine channels. Furthermore, Gross pre- by Briggset al. (1983).However,in
the nearly
paredthin sectionsofthe Pa elemenlsof Ozar- three pagesJetreriesdevotes
to listing the charkodina murchisonl (Pander)and determined actersof this hypotheticalcreature,
I find onlv
frorn the arrangementoftheir lamellaethat the six that might be more or less
objectivelyrepelementsmust havegrownby accretionofnew resentedin (or by) the Scottishspecimens(mamaterial to their outer surfaces,not by addition rine habitat, eel-shapedbody, head-trunk-tail,
ofnew lamellaeon the inner surfaceofthe ele- rasprngtee1h,?lensless
pairedeyes,?somites).I
ment, as Pander(1856)had concludedwasthe suggestthat, as Jefferiesintimates,
the resemcase.Gross vr'asthus able to assertwith some blance betweenhypotheticalanimal ..s,'and
confidencethat conodont elementswere not the Scottishconodontsmay be so striking
bethe teeth or derrnal scalesof vertebrates.nor cause the latter were used as a general
model
werethey componentsof the vertebrateendo- for the former.
skeleton,which should show evidenceof ossiIn reachingtheir conclusionson conodontaffication about a core of spongy, cartilagenous finities, Aldridge er al. (1986)
and Jefferies
tissue.Gross concludedfrom his studiesthat (1986) clearly identiry with Pander,
Huxley,
conodonts were primarily soft-bodied chor_ Newberry,Macfarlane,and Ulrich
and Bassler,
datesassignable
to a distinct branchofthe Ae- althoughtheir reasoningis somewhatdifferent.
natha. lt is ironic. I suppose,that his detail;
Becausethe Scottishspecirnenslack any evistudiesled Grossto conclusionsvery similar to denceofjaws or a bony skeleton,it is
the opinthose advancedby Huxley, Macfarlane,and ion of Aldridgeand his coauthon
that
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THE PHYLUM CONODONTA
179
the chordate feature to which the conodont appamtus
shows the greatest similarity is the lingual structure of
hagfishes. . . in which the rows ofteeth are not opposed
when at resl, but are evefted by protractor muscles in
such a way thal the action is bilateral rather than
vertical.
basal plate, which supports the floor of the
mouth and is the rigid, immovable site of at_
tachmentofthe jaw muscles.
Contraction of prolractor muscles attached,
to its anterior end causesthe dental plate to
slide forward in the grooved upper surface of
Sincethis is alsothe chordatefeatureto which the basalplate.
As it doesso, its sidesflatten
Huxley, Newberry, Macfarlane, and Ulrich and and the rows
of lingual teeth above it are
Basslerassignedconodontelements,and a fea- raised, spread
laterally, and eyerted from the
ture of Jeferies' (1986) hypothetical animal mouth (Fig.
8.3, upper left). Dawson (1963)
"s," the first cro\ln vertebrate,a briefreyiew of likens this sequence
ofevents to the openingof
its charactersis in order.
a book. Contractionofmusclesattachedto the
posteriorend of the dental plate efects retraction of the plate, closingof the book, and ap8.4.8 The lingual apparatusofVyxine
proximation of the teeth in a graspingmove_
Figure8.3includesventral viewsofthe headof ment. Repeatedprotraction and retraction
of
Myxine glutinosa, a typical hagfish, drawn to the lingual apparatus altemately spreads
and
show the toothed lingual structure everted(left, approximatesthe rows of teethand is
effective
above) and retracted(right, above).The sagittal in tearing food and conducting it into
the
and transversesectionsbelow these two views mouth. During theseripping motions,the
lone
provide schematicinformation on the anange- palatal tooth may
serve as a gaff by impaling
ment of supporting plates and principal rnus- chunksof foodstuffand therebykeeping
them
cles.Accordingto Dawson(1963),the lingual from slippingout ofthe mouth during prorracslructule of Myxine has four primary compo- tive moyementsofthe dentalplate.
nents. Above the mouth there is a single,backIndividual lingual teeth of Myxine glutinosa
wardly directedpalatal tooth,which is median (Fig. 8.4A) are complexstructuresmade
up of
in position and firmly attachedto an overlying an outer horn cap separatedby
an epithelial
plate of palatal cartilage. The laterally com- layer from an inner
conethat includesdistincpressed,slitlike mouth is flanked by flaps of tive, goblet-shaped
pokal cells.In the centerof
mucousmembrane,which arecontinuationsof each tooth is a mesodermalpulp
cavity. The
the cephalicepidermisand fold into and line base of the outer horny cap is embedded
in a
the oral cavity.Embeddedin the mucousmem- groove in the mucous membrane
of the oral
brane on either side of the oral cavity are two cavlty, exceptlaterally,whereit joins adjacent
longitudinal rows of sevento nine yellowish- teeth. Dawson (1963) notes that
severalaubrown, horny teeth,which are conicalin shape thors believe that growth of the horn
cap takes
and joined at their basesto form a tooth comb- place by cell division and keratinization
in this
When the lingual apparatus is retracted (Fig. basalgroove.Cells in the epithelial
layer be8.3, upper right), the tooth combslie flat, and tweenthe apexofthe horn cap and that
ofthe
apicesof their componentdenticlespoint in- inner pokal-cell cone appearpl/.led.apart,
as if
ward and posteriorly.
they had been stretchedbetweena horn cap
The lingual teeth of Myxine are firmly at- that gIew more rapidly than the
underlying
tachedto a cartilaginols dental plate (FiE. 8.3), pokal-cellcone.The latter,composedofwhite,
which is a troughlike structure of V-shaped opaque,resilientectodermalcells,is peculiar
to
crosssectionthat lies just below the mucous the Myxinoidea and surroundsa small pulp,
membraneof the oral cavity. Its longitudinally which contains blood vesselsand nerve
fibers.
keeledundersidemoves forward and backward Various authors have speculatedthat
the
in the grooved upper surfaceof a cartilaginous pokal-cellcone might be a replacement
tooth,
Fig.8.3..
.Myxine glutrroJa. Top two views show venrral side of head,with lingual apparatusextended(left) and
retracted (dght). central and bottom figures are tralsverse and sagittal sections-of t}re rieao
to strow erem'enti a.J
'
musculatureofthe lingual apparatus.Rledrawnfrom oa*son fts65i_
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VJNOCONOJ IIHJ
THE PHYLUM CONODONTA
lconiform element
rhich are the
donts collected
rtilaginous pala are required
. in Myxine. I
brtable just to
eiltrer. That is,
tEerve rmpresI havebeeninnld have been
American Silu:served of the
Ens preserves
n might be inal;' three likely
natus designed
imately ingest
herd grasping
F,ir€(d lalam.I join Dawson
trses of any of
cuetoped inde-
l8l
pendentlymany times. If so, similarity in me- the repositoryfor the Conodonta
or its sister
chanical action, even if real, may not imply group.
very much about biologicafiniries.
Without in any way minimizing the possible
signilicance of the bonelike features reported
by Barskov, Moskalenko, and Starostina
8.4.9 Summaryoffeaturcsindicatingchordate
(1982)
from the basalplate of severalMiddle
qlfinities
Ordovicianelernentsfrom Siberia,it shouldbe
In summary. evidencetaken by previous au- notedthat no suchfeatureswerenotedin cornthors to suggestchordate (or vertebrate) affiri- parableparts of any of the conodontelemeqts
ties for conodontsconsistsof (l) similarity in studiedby owig (195t), Gross(1954),Miiller
chemical composition of conodont elements and Nogami(1971),or in the closelysimilareleand the mineralizedtissuesof yertebrates;(2) ments of ArchaeognathusrccEntly analyzed in
the shapesofconodontelements,which suggest detail by Klapper and Bergstrtim(1984).The
that they functionedas teeth or jaws; (3) the affinities of Coleodus, to which the Siberian
presencewithin the basalplatesof severalSi- specimenshave been assigned,
are unknown
berian Ordovician specimens of structures amongconodonts;and the Siberianfossils
are
suchasthosethat characterize
vertebratebone; diferent enough from the tlpe of Coleodusto
(4) tlte chevron-shaped
impressionsin the pos- raiseat leastmodestdoubt about their identiterior half of seyeral Scottish Carboniferous lication.In short,the featuresnotedby Barskov
specimens,which have beeninterpretedto be and his coauthorsmay indicatethat the basal
myotomes comparableto those of Branchio- plates of the specimensin which they found
stoma(:Amphiorus); and (5) imptied similar- them are bone.But it is not entirelyclear that
ity of function betweenlhe cephalic apparatus the specimensare conodont elements!
Nor
of conodonts and the lingual apparatusof have those featuresbeen seen in the basal
Myxine and.other hagfishes.
platesof elementsrepresenting
other species.
With respectto compositionalsimilarity, it
Orvig (1951)notedthat someofthe conodont
has been noted repeatedly that phosphate elementsin his collectionfrom the Silurian
of
mineralizationis widespreadwithin the animal Estoniaand the middle DevonianofOhio have
kingdom and henceis not, ofitsell evidenceof "a baseconsistingofa substancewhich differs
anything more than an enzymatic system that . . . frorn. . . materialof the tooth-likecusps.. .
hasnot beencompletelymodifiedto permit ac- but in this substance
oneis not concernedwith
cumulationofcarbonateminerals.Note,in this any kind of bone tissue."He also pointed
out
regard,the recent determination by Tillier and that the Middle OrdovicianHardingSandstone
Cuif(1986) that the buccalelementsof certain of Colorado, which yields elementsof Coleodus
aplacophoranand scaphopodanmollusks are and a greatvarietyofother typesofconodonts,
phosphatic.
also yieldsspecimensassignedto the ostmcodWith respectto shape,I needonly point out erms Astraspis and. Eriptychius that exhibit
that teeth,graspingspines,copulatoryhooklets, practicallyall the typesofhard tissueknown
in
and the like, have similar shapesin whatever vertebrates.To my mind, this statementis
siganimal body they are formed becausethoseare nificant becauseit suggests
that conodontelethe shapesthat do the job, not becauseall or- ments with superficiallybonelikematerials
a1
ganismswith teeth,graspingspines,and copu- the baseand specimenscomposed
of real bone
latory hooklets are closely related. In addition occur in collectionsfrom the samebeds
and
to those formed in various ways by diferent that tbey may be readily distinguishedby
one
groupsofchordates,structurcsthat are similar with a trained eye.
to chordate teeth in form and function are
Finally, anyone familiar with the elegant ilfound in distantly relatedgroupssuchas mol- lustration of Myxine glutinosa in Miiller's
lusks, arthropods,trematodes,nemertineans, (1836) famous monograph on the hagfishes
gnathostomulids,and chaetognaths.
It is thus (Fig. 8.2)is boundto be struckby the many feano surpnsethat someone,sometime,has con- tures that seemto be sharedwith the best
Dresideredone or anotherofthose groupsaseither servedof lhe ScottishCarboniferoussoecimens
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YJ,NO(IONO] 3HJ
z8r
THE PHYLUM CONODONTA
.rc be€n trying
h Chordataor
crtebratephyla,
rsrated convic'organizational
E a aroselate in
c Paleozoicand
since
Han.
However,
-tracted)
lin the late Prely of metazoan
le such organihve survived,
hce" between
ruch less than
rvrvors.
rtte idea(albeit
t-aflhropod" is
I independently
te gradesof ord rhat at least
confiisingcomthn characters,
fled to survive
lested recently
ne members of
lave been aslbn to several
:nlatlves, may
organizational
rtitrgdom!)that
mmotiids and
rps. And it has
tt eventhe l/enD mor€ than a
hates and verhtives may reI of the "chortral sides into
€-, the Echinor phyla, or even
r-Yof trying to
: bas done little
ipeness. Their
ftatures of the
le1- have comr linle hard in-
183
formation we have on soft-part anatomy Branson,E. B., and Mehl, M. G. (1933).Conodont StudiesNo. l. Univ. Missouri Studiest,
speaksto us in severaltongues.Thus, I am
5-72.
comfortablein consideringthe Conodontaan
Briggs,D. E. G., Clarkson,E. N. K., and Aldridge,
extinctphylum of small.solitarymarineaniR. J. (1983). The conodont aqimaL Lethaia
mals,with compressed
or depressed
vermifom
16,t-14.
bodies that were soft except for a variously dif- Ctark, D. L., Sweet, W. C., Bergstriim, S. M.,
Klapper, G., Austin, R. L., Rhodes, F. H. T.,
ferentiated assemblageof phosphatizedepithelial elementsin the cephaliclobe.The elements Mtiller, K. J., Zieg)er, W., Lindstriim, M.,
Miller, J. F., and Harris, A. G. (1981). Conby which conodontsare represented
in the fosodonta. Pt. W, Suppl. 2 il Trcatiseon Invertesil record may have functioned as grasping brate Paleontology(ed. R. A. Robison), Geol.
spinesand buccal or pharyngealteeth. And the
Soc.America and Univ. Kansas,202 pp.
cosmopolitanand broadly facies-independentConway-Morris, S. (1976). A new Cambrian lophophorate from the BurgessShale of British
distributionofmany speciessuggests
they were
Columbia. Palaeontology 19, 199-222.
pelagicanimals, probably nektonic, although
Dawson, J. A. (1963). The oral cavity, the'jaws'
others,of more limited distribution and facies
and the horny teeth of Myxine glutinosa. Pp.
dependence,may have been nektobenthicor
231-255 il The Biology of Myxlze (ed. A. Brodal and R. Fange).Universitetsforlaget,Oslo.
even benthic. Conodonts were probably a
minor phylum.neverparlicularlyconspicuousDubois, E. P. (1943). Evidence on th€ nature of
conodonts. Pal. 17. 155-159.
componentsof Paleozoicor Triassic marine Ellison, S. P., "L
Jr. (1944).The composition ofconfaunas.Their ultimate extinctionat the end of
odonts. J. Pal. lE, 133-140.
the Triassicwasprobablythe cumulativeresult Gross, W. (1954).Zur Conodonten-Frage.,SerrckenbergianaLethaea 35, 73-85.
of a long decline through the Carboniferous
Harley,
J. (1861).On the Ludlow bone-bedand its
and Permianand a gradualreductionin genocrustaceanremains.Q. L Geol,Soc.London 11,
mic diversity. In spite of their probablerarity
542-5s2.
in Paleozoicand Triassic seas,conodontsare Hass, W. H., and Lindberg, M. L. (1946).Orientation of crystal units ofconodonts. J. Pal.20,
abundantand commonasfossilsright up to the
501-504.
top ofthe Triassic,probablybecausethey were
Hinde, G. J. (1879). On conodonts from the
blessedwith heala, chemicallyresistantskeleChazy and Cincinnati group of the Cambro-Sital elementsthat providedthem a placein poslurian, and from the Hamilton and Geneseeterity denied to other less suitably endowed shaledivisions ofthe Deyonian in Canadaand
groups.
the United States. 0. "/. Geol. Soc. London 35,
351-369.
Hubrecht, A. A. W. (1883).On the ancestralform
of the chordate. Q. J. Microsc. Soc. 23, 349References
368.
Aldridge,R. J., Briggs,D. E. c., Clarkson,E. N. (1887). The relation of the Nemertea to
K. and Smith, M. P. (1986).The amnitiesof
the Vertebrata.Q. J. Microsc. Sci.21,605-644.
conodonts-new evidencefrom the Carbonif- Hyman, L. H. (1951). The Invertebrqtes:Plqtyerousof Edinburgh,ScotLand.
Lethaia19,| -l 4.
helminthes and Rhynchocoela-The acoeloAldridge,R. J., Srnith,M. P., Norby,R. D., and
mate Bilateria.y.lL Mc Graw-Hill. New York.
Briggs,D. E. G. (1987).The architectureand
550 pp.
polygnathacean
functionof Carboniferous
(1959). The Invertebrates: Smaller Coelocon- odontapparatuses.
Pp. 63-75in Palaeobiology mate Groups. V. 5. Mc Graw-Hill, Ne\ryYork,
of Conodonts(ed. R. J. Aldridge). Ellis Hor783 pp.
wood,Chichester,
180pp.
James,U. P. (1884).On conodontsand fossil anBarskov,I. S.,Moskalenko,
T. A., andStarostina, nelidjaws. Cincinnati Soc.NaL Hist. J.7. 143prinadlezh- 149.
L. P. (1982).NorTe dokazatel'stva
nosti konodontoforid k pozvonochnym.Pal Jefferies,R. P. S. (1986). The Ancestryofthe VerZhur. 19E2,80-86. [English translation in Patebrates. Carnbidge Univ. Press, Cambridge,
leontologicallournal 1982,82-90.1
376 pp.
Bengtson,
S. (1976).The structureof someMid- Kirk, S. R. (1929).Conodontsassociatedwith rhe
dle Carnbrianconodonts,and the earlyevolu- Ordovician fish fauna of Colorado-A prelimtion of conodont structure and function. Iernary roe. Am. J. Sci.Ser.5, 18(108),493-496.
thaia7, 185-206.
Klapper, G., and Bergstriim, S. M. (1984). The
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6e lower Sile apparatus
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&n* (ed. R.
rst€r, 180pp.
l rhe Harding
lcolorado. "/.
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APPENDIX A. A SUMMARY CLASSIFICATIONOF THE
CONODONTA
Phylum ConodontaPander,1856
ClassCar.idonti,new
Order Proconodontida. new
Family ProconodontidaeLindstr6m,
t970
E oconodont us Miller, 1980
Miller, 1969
Proconodontus
Family Cordylodontidae Lindstriim,
r970
CambrooistodusMiller, I 980
CordylodusPzndet 1856
I apetognathus Ianding, | 982
Family FryxellodontidaeMiller, l98l
Fryxel lodontus Miller, I 969
Family PygodontidaeBergstriim,198I
N eri codus Li'j.ds1'Jom,| 955
PolonodusDzik, 1976
PygodusI-amont and Lindstriim,
1957
Order Belodellida, new
Family BelodellidaeKhodaleyich and
Tschernich,1973
B eIodeIIa Ethi'lgtoll, 1959
Coelocerodontus Ethington, 1959
Dvorakia Klapper and Barrick, 1983
Stoladus Lindstrdrn, 1955
Il aIIiserodus Serpagh,| 967
Family Ansellidae Fihraeus and
Hunter, 1985
AnsellaFirhtaeu;s
andHunter, 1985
Yiira,
1975
Hamarodus
2
n€w
Dapsilodontidae,
Family
BesselodusAJdridge,1982
Dapsi lodus Cooper, | 9-l6
ClassConodontiBranson,1938
Order Protopanderodontida,new
Family Protopanderodontidae
Lindsrrtim, 1970
Glypt oconus KenJc,dy, 1980
M onocostodus M.rller, 1980
OneotodusLindstriim, I 955
Landing, 1982
Parutahconus
ProtopanderodusLlndstriim, I 97I
Drygant, 1974
2Pseudooneotodus
ScabbardeIIa Orchard, I 980
SemiacontiodusMiller, I 969
Stauferella Sweet,Thompson, and
Satterfield,1975
2Strachanognat
hus Rhodes,1955
Teridontus Miller, 1980
TropodusKennedy, 1980
UtahconusMiller. 1980
VaiabiloconusLanding,Bames,and
Stevens.1986
FamilyClavohamulidae
Lindstrdm,
t970
Clavohamulus Fumish, | 938
H i rsutodont us Mjlle\ | 969
Seftat ognathus I-fJe,| 970
?Paraserratognat
hzs Yang, 1986
Family AcanthodontidaeLindstrtim,
t970
Acanthodus Fu;mish, | 938
Cor nuodus F ahrircrs, | 966
D repanodusPander, 1856
Parapaltodus Stov$e, 1984
ScalpellodusDzik, 1976
Ulric hodina Furnish, | 938
Family DrepanoistodontidaeFihraeus
and Nowlan, 1978
D repanoistodusLindstrdm, I 97I
Paltodus Pande\ 1856
ParoistodusLindstrttm, I 97I
Order Panderodontida,new
Family Panderodontidae
Lindslriim,
r9'10
BeIodi na EthilJigton,| 959
CulumbodinaMoskalenko,1973
N eopanderodus Ziegler and
Lindstrtim, 1971
PanderodusEthinglon, 1959
Parabelodina Sweet,| 9'l 9
ParapanderodusStouge,I 984
PlegagnathusEthington and Fumish,
1959
Pseudobelodina Sweet, 1979
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A SUMMARY CLASSIFICATION OF THE CONODONTA
llae Bergstrilm,
r6m, 1983
r. 1966
ftr. 1935
he Harris,
mson and
L 1 938
rb and Harris,
ton and Clark,
ad Harris,
-,genusfor
Ete P
t5
B5
33
r llass, 1959
t935
rd Mehl,
: Lindstriim,
a|IOmand
rrp.elj, 1974
E- Mehl, and
dstrdm, 1970
rtu. l97l
I
Etrson and
nd Mehl,
ud Mehl,
I Mehl, 1933
sler. 1925
Dd Mehl,
tL 1982
Hibbardella Bassler,1925
I dioprioniodusGlnnell, 1933
Kl adognathus Rexroad, I 958
OulodusBrarson and Mehl, 1933
Prio ni odina Bassler,1925
PristognathusStone and Furnish,
1959
?Ligo nodina Bassler,1925
?Lonchodina Bassler,1925
Family BaclrognalhidaeLindstriim,
t9'70
Bactrognathus
Bransonand Mehl,
194l
DoliognathusBransonand Mehl,
t94l
Dollymae Hass, 1959
Eotaphrus Pierce and Langenheim,
t97 4
E mbaysgnathus Metcalfe,I 98I
ScaliognalhusBransonand Mehl,
t94l
Staurognathus Branson and Mehl,
194l
Family EllisoniidaeClark, 1972
EllisoniaMij.ller,1956
Furnishius Clark, 1959
GladigondoIelIa Miiller, I 962
Hadrodontina Staesche,I 964
Metillina Kozur and Mock, 1974
Pachycladina Staesche,I 964
SweelinaWardlawand Collinson,
1986
Family GondolellidaeLindstrdm,
l9'10
CarinellaBt trov,1973
CypridodeIla Moshey 1968
EpigondoIella Mosher I 968
Gondolella Statffer and Plummer,
1932
Misikella Kozur and Mock, 1974
Mosherella Kozur, 1972
NeogondolellaBender and Stoppel,
r965
Neospathodus Mosher, I 968
Platyvillosus Clark, Sincavage,and
Stone1964
Pseudolurnishiusvan den Boogaard,
1966
Xaniognathus Sweet,1970
Order Ozarkodinida Dzik, 1976
t 87
Family Spathognathodontidae
Hass,
l9s9
Amydrotaxis Klapper and Murphy,
1980
AncyrodellaUlich and Bassler,1926
A ncyrodeIloides Bishoff and
Sannemann,
1958
AphelognathusBranson,Mehl, and
Branson,l95l
BispathodusM.ilrlle4 | 962
E ognathodusPhllip, | 965
Lochriea Scott, 1942
Mehlina Y oungquist, 1945
OzarkodinaBransonand Mehl, 1933
PandorinelIi na M;jller and Miiller,
t95'l
?"Plectodina" (new genusfor specres
without pastinateP elements)
PolygnathoidesBranson and Mehl,
t933
Pseudopolygnathus Branson and
Mehl, 1934
Rhachistognat
hus Dunn, I 966
Scaphignathus
Helms, 1959
Torlodus Weddige, 1977
VogelgnathusNorby and Rexroad,
1985
Yaoxianognathus An, | 985
Family KockelellidaeKlapper, l98l
A ncoradelIa W zlliser, 1964
KockeIella W alliser, 195'1
Family Pterospathodontidae
Cooper,
1977
Apsidognalhus
W alliser,1964
Astropentagnat
husMostler, 1967
AulacognathusMostler, 1967
Carniodw Wallisel 1964
Johnognathus Mashkova,I 977
PterospathodusW alliser, I 964
Family PolygnathidaeBassler,1925
Ancyrognalhus Bransonand Mehl,
1934
?A ncyroIepis Ziegle4 1959
PolygnathusHindq | 879
PolylophodontaBranson and Mehl,
t934
Family Palmatolepidae,new
Klapperina l-ane, Miiller, and
Ziegler,1979
MesotaxisKlaEEerand Philip, 1972
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; 19'72
APPENDIX B. STRATIGRAPHICRANGE CHARTS
in and
rnd
I Mirchell,
tin and
lin and
d Mehl,
h and
Alekseev,
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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Ranges of selected Triassic conodont species.
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
Apparatuses,
38
of, 142
€laboration
reductionof, 143
Apsidognathus,
99, 187,Fig.5.43
173,I82
Archaeocyathida,
Archeognathus,
123,l8l, 188,Fig.5.59
| 73
Aschelminthes,
AshlockFormation,155
Aslraspis,l8l
Astropentaghathus,
99, 187,Fig.5.43
40
Altachmentsurfac.e,
Aulacognathus,
99, 187
Au lobodui, 186
Bactrognathidae,
83, 145,156,187
Bactrognathus,
83,84, 136,159,lE1,Fig.5,32
anchorarius,84
excavatut,84
hamatus,84
Balognathidae,
63,64,1l,15, 144,186
Balognalhus,
63
Baltoniodus,63,
64,65,186,Fig.5.15
Eovince, 167
Baltoscandic
Bars,15,16
Basalcavily,13,15,40
Basalfilling,12
Basalpit, 13,22
Basalplate,179
Base,13,15
Belodella,
45,49,185,Fig.5.3
42,45,49, 133,185
B€lodellida,
42,49, 185
Belodellidae,
Belodia,18,55,57,58,141,185,Fig.5.10
57
calciprcminens,
comprcsso,
58
monilorcnsis,
58
Berys!rcemoghathus,
75,76,186,Fig.5.26
extensus,l6
Besselodus,50,
51, 185,Fig.5.3
Bimembrateskeletalappalatus,24
iamiformelement,18,24
Bip€nnate
Bnksleldia,65
pectiniformelement,22
Bisegminiscaphat€
Bispathodui,
89,90,94,96,97,136,l4l, 159,187,
Fi gs.5.39,5.41
aculealus,96
biswthodus,96
s,96
spinulicostat
109,I12,Fie.5.50
stabilis,96,97,
utahensis,
96, 159
Blades,15,20
205
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INDEX
Distacodidae,36
Distacodontacea,41, 42, 5l
Distacodontidae,36, 5l
Distacodus,51,54
Distomodontidae,67, 68,11, 134, l4l, 144, 167, 186
Distomo&rs, 66, 61, 68,1 l, 98, 186,Fie. 5.19
kentuckyensis,61, 68
Diversity cycles,130
firsr-order, 130, 132, 133
long-term, 130, 132, 133
second-order,13l, 133, l 40, 146
Dolabrate coniform elemcnt, l8
Dolabrateramiform elemenl, 18, 24
Doliognathus,83, 84, 136, 159, 187,Fig. 5.32
Do ytnae, 83, 136, 187
Drakes Formation, 154
Drcpanodus,54, 185,Fig. 5.?
slrkttus,56
Drepanoistodontidae,54, 185
Drepanoistodus,54, 55, 59, 134, | 57, 185,Fig. 5.8
sp. aff. suberectus,23
Dvorakia, 45, 149, 185,Fig. 5.3
]7-Fi& 5.35
Il. 137,139,
Echinodermata,173, 182
EinfacheZahne, 15, 35
Element, I I
Elictognathidae,89, 96, 106, 136, 188
E lictoghathus, 106, 107
Elltsonia,83,85, 86, 89, 137,138,139,161,163,187,
Fig. 5. 33
teichefti, Il7
ttiassica,86,87
Ellisoniidae,83, 85, 142,166, 187
Embaysgnathus,84, 18?,Fig. 5.32
Enanliognathus,18, 83
Eobelodina,5T
Eoconodontus,46, 47, 185,Fig. 5.2
Eognathodus,89, 93, 94,95, 96,91, 135, l4l,181,
Fies . 2. 11, 5. 39,
5. 41
sulcalus,26,96
Eoneop oniod s,72
EopIacognathus, 1 l, 72, 102, | 43, 186,Fie. 5.21
Eotaphrus,83, 136, 159, 187
Epigondolella,87, l3'1,142,146, 187,Fig. 5.35
,g/rkd, 81, 186,Fig. 5.30
divaricala, 8l
Eriplychius, l8l
Eismod s, 18,79,81,134,186,Frg-5.29
quadridactylus, 79
Erratrcodon,
78, 79, 81, 85, 186,Fig.5.29
palu, 79
Euconodontelement,42
Euconodonts,4l
Evolutionary index. ,SeeIndex ofevolutron
Exoc hognot hus exwnsu-s, 68
Extensiformdigyrateelement, 18, 24
Fairview Formation, 155
Falcodus, lO4, 106, loj
anguhts, lO7
Fibrous elements,?9, 81, 165
207
Fibrous struclure, 14
Flotant marsh, 164
Form taxonomy, 5, 35
Freeblade,I13, ll7
Fryxellodontidae,48, 185
Fryxellodontus, 48, 185, Fig. 5.2
Fumishinacea,42
Furnishiut,85,86, 138,187,Fig. 5.33
Fus€dcluster,24, 3?, 41, l7l
GamachighalhrLt,64, 65, 67, 186,Fig. 5.15
Geniculateconiform element, 15, 16, 25
Germanic Province, 168
Gladigondolella,87, 138, l8?, Fig. 5.34
meeki, 81
tJtY?toconus,
)J, l6)! frg. ).)
quadruplicatus,53
Gnathodontidae,
89, 95,96, lO9, ll2,I13, 135,l8E
Gnathodus,
109, Ill,ll2, 136,14l, 159,t88, Fig. 5.50
austi L lll
bilineatus, IIO, I I t, 112,Fig. 5.50
anneifurmis,109,Fig. 5.50
delicarus,109,t1l, Fig.5.50
girtyi, l l l,137, Fig. 5.50
praebilineatu,s,lll
p nctalus, lO9,I10, Fig. 5.50
semiglaber,111,Fig. 5.50
texaws,lll,Fit
5,50
typicus, 109, I10, Fig. 5.50
Gnathostomulids,173
Gondolella,81,88, 89, 137, 138, 142, 16I, 162, 163,
165,166,187,Fig. 5.35
navicula,1'l
Gondolellacea,41, 42, 78
Gondolellidae,78, 83, 87, 88, 89, 138,142,146, 166,
187
Gondwanaglaciation, 134
Grant Irke Formation, 155
Grier Member (of Lexington Limestone),155
Habit (ofconodon$), 148, 150
benthic,152
nektobenthic,l5O, 152, 164
pelagic,150,152,164
Hadrcdoatina,85,86, 138,187,Fig.5.33
Hadrcgnalhus,68, l4l
sta rcgnathoides,68
Hagfrsh. See Myxine
Hamarcdus,49,50,185,Fig. 5.3
Harding Sandstone,l?7
Hemichordata, 173
Heterochrony,145
H ibbardella, 11, 8 | , 82, 84, 187, Frg. 5.29
angulata,8l
Hibbardellacea,42, 7E
Hindeodus,l15,l16, l17, l18, 120,137,139,I40, 159,
1 6 0 ,1 6 l , 1 6 4 ,1 8 8
uassidenlatus,I 15, 116,Fig. 5.54
crlstulus, | 16, I l'1, Fig. 5.54
jufensis, I 17
minulus, lI7
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INDEX
Neogondolella,
87,88, 137,138,142,146,187,Fig.
bisselli,131, 138
clorki, 137
mofiberye sis,138
pfaeD6te t, tt I
Il, l0r
! Fs 5.51
! 154,155
j
I Fre.5.58
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It2
L 1,14,150,
t 5.52
Neomuhioistodus,2L,72,73, 186,FiE 5.22
clypeus,73
Neopanderodus,
55, 57, 185
Neoptioniodus,82,83
Neospathodus,87, 137, 138, 142, lE7, Fig. 5.35
ll7, ll9, 131,138, 188,Fig. 5.55
Neostteptognathodus,
Neoteny, 146
Ne codl,ts,48,185
Neurodontiformes,14, 36, 79, 81, 155
Noixodonlus,64, 186
Nongeniculateconiform elemenl, 15, 16
Nonh American Midcontinent Province, 167
Nonh Atlanric Provinc€, 167
Nolhognathella, 104
Octimembrateapparatus,64
Odontogtiphus, 28, l'l 3
omolius,173
Oelandodus,61,185, Fig. 5.13
Oepikodut,63, 186,Fig. 5.14
Otrshorefauna, 165
Ohio Valley Province, 167
Oistodontidae,42, 50, l4l, 186
Oistodontinae,61, 186
Oistodus,61, 186,Fig. 5-l3
Oneotodontidae,52
OneotodtLt,52, 53, 185, Fig.5.5
Origination-extinctionratio. SeeIndex of evolution
Ostracoderms,176
Oulodw 18,81,154,155,156, l81,Fie. 5.29
se atus,8l
Outsideshale,161,163
Ozarkodina,91, 92,93, 95, 96, 98,99, l2O, 115, 142,
187,Figs.5.38,5.39
abrupta,97
confluens,33
inclinata,92,94
murchisoni, 177
n. sp. of Bergstritm,9l
renscheideksis, 95
sannemana,96
se$,93,94,95,99
semialtemens,93, 106
tortilis, 37
Ozarkodinida,
42, 45, 75,89,90,lJ4,135, 140,l4l,
187
P position (or element),24
Pa position (or element),24, 30
Pachycladina,85, 86, 138, 187,Fig. 5.33
Paedomorphosis,145, 146
Palataltooth, 179
Palmatodella, lO4
Palmatolepidae,
89, 102,106,135,187
Palmatolepis,102, 104, 105, 135, 136, 145, 188,Figs.
5.44,5.4',7
pe obata,105
rugosa, 105
rugosa ampla, 105
rugosa lrachyleru, 105
Itia gula s, lO5
PaLodus,53,54, 185, Fig. 5.8
Pander,Christian Heinrich, 3
PanderSociety,7
Panderodontacea,
4l
Panderodonlida,42, 45, 55,56, l4l, 185
Panderodontidae,56, 167, 185
Panderodus,
55, 57, 58, 59, l4l, 156,167,170,185,
Fig.5.9
berystrcerni,56,58, 141
gr4cuts,zJ, rtE z.t
sulcatus,5l
unicostatus,32, 57, Frg. 3.3
Pahdetulepis, lO5
Pandotinellina, 92,93, 95, 96, 107, 123, 136, 142, | 59,
l8?, Fis. 5.37
insita,93,96,l2l, 123
Parubelodiha,18,
55, 58, l4l, 185,Fig.5.l0
denticulata,56, l4I
Paruchtognathus, 79
Paraconodonta,4l
Paraconodontida,41, 42, 45, 175
Paraconodonts,28, 41, 42
Paragnalhodus,lll
Parupaltodus,54, 185
Parupandetudus,55, 56, 185,Fig. 5.9
asymmetficus, 56, 51
striatus,56
Parapet,109
Paruprioniodus,73, 134, 186,Fie. 5.22
costalus,73, l4l
Paraserrutognathus, 185
Psroistodus,54,55, 185,Fig. 5.8
horridus, 55
Porutahconus, 53, 185
Pastinatep€ctiniform element,2l
Pastiniscaphate pectiniform element, 22
Patrcgnathus,96, 120,122, 136,I88, Fig. 5.57
Pb position (or element),24, 30
Pectinat€t€eth, 19
Pectiniform elem€nt, 15, 20, 24, 36
Peda s, 65, 66,67, 68,7L, 134,135,146,l86,Fi&
5.18
lalialalus,7I
Pelelcysgnathus,
60, 65, 65,69,70,71, 135, 136, l4l,
142, t46, t86,Fie. 5.20
csakyi,lO
inclinatus,70
index, 70, Il, l4l
"Pelekysgnathus," 7o
Peiodon, 18, 35, 16, 77, 156, 186,FLl- 5.27
acuteal4, to, t t
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INDEX
5-47
)
l zl l, 1 45,
t 3 3,l4l,
Lll
45. Fis.5.2
188.Fig.5.50
rt5
Fr. 5.10
13i
!
[3,1.187
al- lE7, Fig.
Rabeignathus,117, I19, 188,Fig. 5.56
Radular teeth, 172
Ramiform element, 15, 16,24
Rastrat€element, 15, 18, 55, 57
Recapitulation,145
Recurrentgroup, 39, 40, 4l
Red River Province, 167
Rhachistognathus,97, 136, 137, l4l, 187, Figs. 5.39,
5.41
Rhipidognathidae,62,7 5, 99, 186
Rhrpidog4athu:,
15,16, 155,156,157,l86,Figs.2.ll,
5.26 \
slmmetricttl;,l54
Rhodesognathui564,134, 156, 186,Fig. 5.16
Rhynchocoela,3l! 176, 182
Robtglicoslatusstiock(of Polygnalhus),99,100, l0l
Rossods,53,60,61,133, 186,Fig. 5.12
Rostral ridge, 102
Rostrum, 102
Rotundacodina,
67, 68, 186,Fig.5.19
dubius,68,7l,146
S position (or element),24,25, 30
Sa position (or element),25
Sagiua,41,174, l'l5
Sagittodohtina,64, 67, t86, Fig.5.l5
Sannemannia,65, 66, 186
pesanse s, 66
Sauk sequence,132,133, 134
Sb position (or element),25
Sc position (or element),25
Scabbardella,53, 185,Fig. 5.5
kaliognath s, 33,84, 159, 187
Scalpellodus,54, 185,Fig. 5.7
Scandodus, 54
Scaphaleattachments!fiace, 15,22
Scaphignathus,96, 107, 123, l4l, 187,Fig. 5.41
Schmidtognathus,102, 104, 106, 135, 188,Fig. 5.47
Scolopodontidae,52
Scolopodri, 53
gracilis,56
Sclttula, lO4
Scyphiodtu,73,'14, 143, 186
primrs,74,Fig. 5.25
Sd position (or element),25
Secondaryprocess,21
Segminate pectiniform element, 22
sefitacon oaus,)2, )J, )o, t6), l.lg. ).)
Septimembrate skeletal appamtus, 24
Serratognalhus,53, 185,Fig. 5.6
Seximembrate sk€letal apparatus, 24
Siphonodella,89,96, 99, 106, 107, 108, 136, 188, Figs.
5. 48, 5. 49
duplicala, 109
pruesulcata,107, 108, 109
sulcato, 107, IO9
Siphonognathui, 106
Skeletalappararus,23, 37, 38
Skeletalelemenl, 11
Solenogastres,
172, 173
Somites,29, l7l
2tl
Spathodus,ll5
Spathognathodontidae,
89, 90, 91, 94, 95, 96, 97, 106,
tt2, t34, 116,t4t, 144,167,t87
Spathognalhodtls,ll5
Spatula,I I ?
SlauflercL\a,53,185,Fig. 5.5
Staurcghathla,83, 84, 136, 187,Fig. 5.32
Stellatepectiniform element,21
Steptotaxis,65,66, 186,Fig. 5.18
fumishi grotrp, l4l
Steteocohtls,123, 188, Fig. 5.59
Stolodus,49, 185,Fig. 5.3
Sttuchahoghath s, 185
Strcptognathodl,ts,
ll4, ll5,137,146, 150,l6l, 163,
188,Fig. 5.52,5.53
Subbryantodus,I 15, 116, 188
Survivorship curves, 139
Sweetina,85, 137, | 38, 187,Fig. 5.33
Sw€erognathidae,
90, 92,96, ll2,ll7, 188
Sweetognathinae,I l7
Sweelognathus,89, 117, l19, 137, 138, 188,Fig.
5.55
menilli, l19,131
whitei,IL9,l2O
Symmetry,classesof, 25
Symmetry-tran$ition series,22
Synpioniodina, lO4
TanglewoodMember (of Lexington Limestone),
155
Tangshanodus, 186
Taphrognalhus,l2O, 122, l88, Fle. 5.57
Tenraculata. See Lophophorata
Teridonlidae, 52
Teridontus,45, 52, 5!, 133,142, 157, 185,Fig. 5.5
Tertiopedateramiform element, l7
Tethyan Prcvince, 168
Tippecanoesequence,132, 134
Tooth comb, 179
Tortodus,93,94, 187,Figs. 5.39,5.41
Ttiangulodus, T2
Trichonodello, 11
2expansa,68
T gonodur,12
Trimembrate skeletal apparatus, 24
Tipodellus, IO5
Tripodontinae, 62, 186
Tripodus,60,61,62, 63,72,13,75, 133,14l, 185,Fig.
5.t2
deltotus, '12
Trisegminiscaphate pectiniform element, 22
I ropodus,)5, lu), nrg. ).)
complus,53
Truchercgnalhus, 79
True conodonts,4l
Ulrtchodina,54, 185,Fig. 5.?
Unimembrate skeletal apparatus, 24
Utahfonus,53, 185,Fig. 5.5
"Utica" Shale,155
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