Table Of ContentLecture Notes ni
Earth Sciences
Edited by Somdev Bhattacharji, Gerald M. Friedman,
Horst .J Neugebauer and Adolf Seilacher
30
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Berlin Heidelberg NewYork London ParisTokyo Hong Kong
Editors
Prof Dr Dr. h.c. Erie G. Kauffrnan
University of Colorado
Department of Geological Sciences
Campus Box 250
Boulder, CO 80309, USA
Prof Dr. Otto H. Walliser
University of Gottlngen
Institute and Museum for
Geology and Palaeontology
Goldschmidt-Str. 3, D-3400 Gothngen, FRG
Layout' Gabrlela Meyer, G~)ttlngen
ISBN 3-540-52605-6 Sprmger-Verlag Berhn Heidelberg New York
ISBN 0-38?-52605-6 Sprlnger-Verlag NewYork Berlin Heidelberg
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Loafw September 9, 1965, n~ version its of 24, June 1985, a and copyright fee always must be
paid fall Violahons under the prosecution act of the German Copyright Law
© Berhn Springer-Verlag Heidelberg 1990
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PREFACE
This volume presents results from members of the Project 612 "Global Biological Events in Earth History"
of the International Geological Correlation Programme (IGCP). The project, initiated by the elder editor
(O.H.W.) within the framework of the International Palaeontological Associatio(nI PA) in the late 70s, was
officially establishedi n 1984. Subsequently, it led to the first three conferences on Global Bio-Events, and
their respective symposia volumes: 1) In G6ttingen, West Germany in 1986 (WaUiser, O. H., Ed., 1986,
Global Bio-Events, Springer-Verlag); in Bilbao, Spain in 1987 (Lamolda, M. A., Kauffrnan, E. G., and
Walliser, O. H., Eds., 1988, Paleontology and Evolution: Extinction Events; Rev. Espafiola de Paleont., n.
extraord.); and in Boulder, Colorado, U.S.A. in 1988 (this volume). The next meeting, on Innovations and
Revolutions in the Biosphere, is planned in Oxford, England in 1990, to be hosted by Martin Brasier.
During the history of this project, the focus of our research has shifted significantly. Initial focus was on
specific global mass extinctions (e.g. the Precambrian/Cambrian, Frasnian/FammeniaCnr,e taceous/Tertiary,
and Eocene/Oligocene events) to a broader treatment of Phanerozoic mass extinctions, their differences or
unifying factors, and their causal mechanisms. Subsequent meetings have attempted to focus attention on a
fuller spectrum of global bio-events in Earth history. The Boulder Conference, and this volume, although
still strongly influenced by the excitement of mass extinction research, expresses these new trends in bio-
event studies.
The Boulder conference, held on May 16-23, 1988, focused on a broad spectrum of Abrupt Changes in
the Global Biota. Over 001 participants from 31 nations attended this meeting, representing diverse
disciplines of palaeobiology, palaeoclimatology, palaeoceanography, sedimentology, geochemistry, and a
broad spectrum of the stratigraphic and geological sciences. Four days of talks were supplemented by field
trips to the continental Cretaceous/Tertiary boundary in the Raton Basin, New Mexico, and to the
Cenomanian/Turonian mass extinction interval exposed near Pueblo, Colorado.
The Conference itself was characterized by a great diversity of approaches to bio-event research, and the
phenomenon of mass extinction. In particular, interactive causes involving both extraterrestrial and
earthbound (tectonic, oceanographic, climatic) forces were discussed, and each major Phanerozoic mass
extinction was treated by specialists in the field. In addition, many presentations focused on the causal
mechanism and patterns of bio-eventd evelopment that were not restricted to mass extinction intervals, but
which could cause regional to globbailo tic response at any time in Earth historTyh.u s, both the conference,
and this volume, focus attention on climatic and oceanic perturbations from anoxia, advection, rapid
thermal change, toxic chemical enrichment, and energy shock from impacts and giant tsunamis as forcing
mechanism for regional to global bio-events. The delicate balance of perched ocean/ctimate~fe systems
under typical warm equable non-glacial Phanerozoic conditions, and their susceptibility to shock from even
small perturbations, was a philosophical theme that ran throughout the meeting. The case for
extraterrestrial forcing of tectonic, volcanic, and biological events was greatly strengthened by new data
presented at this conference, with special concern for the effects of small comet/meteorite impacts in the
oceans, and their chemical/physical/biological signature which might be used, in the absence of shocked
minerals, microspheres or trace metals, to identify extraterrestrial events associated with global and regional
bio-events. The conference benefitted from the introduction of much new data at high levels of resolution,
especially from poorly studied mass extinction intervals. Interactive discussions, and many new ideas
characterized the meeting. The new scientific results of this meeting are exciting; they are reviewed in the
Conference Report published in Episodes (1988, .v ,11 n. 4, p. 289-292). Most of the key papers presented
at the Boulder meeting appear in this volume.
VI
What lies ahead in bio-event research? Clearly, a great deal of excitement and an age of discovery. We
have only touched the surface of this new and dynamic field. We are starting to comprehend the dynamics
of global mass extinctions, integrating detailed geochemical, physical and biological data into scenarios of
cause and effect. But in the years ahead lies the job of understanding the whole spectrum of regional bio-
events preserved in the ancient record, and especially the application of this research to solutions of the
critical problems inherent in global change and the modern biotic crisis. Future directions for research at
this conference include the investigation and modeling of abrupt chemical and thermal shifts in the ocean,
the effects of impacts at deep ocean sites, the documentation of successful survival strategies and
repopulation patterns following biotic crises, the deep ocean record of bio-events, and focus on alternative
forces other than impacting to account for mass extinction events. This volume introduces some of these
new pathways in bio-evernets earch.
ACKNOWLEDGEMENT
We are extremely grateful for the encouragement and support of our colleagues, the International
Palaeontological Association, the International Union of Geosciences (IUGS) and the UNESCO. We
further thank the University of Colorado and its Department of Geological Sciences for hosting the 3rd
International Conference on Global Bio-events in Boulder. We received valuable help from Mrs. Dipl.-
Geol.L . Alberti and from Mrs. C. Kaubisch. Special acknowledgement is due to GabrielMae yer who made
any effort to type and to design this volume.
Erie G. Kauffman Otto. H. WaUiser
Host of the Leader of
3rd International Conference IGCP Project 216
of IGCP Project 612
CONTENTS
PREFACE ........................................................................................................................................................... III
GENERAL ASPECTS
,RESILLAW O. H.: WOH to define "global bio-events". ................................................................................. 1
BOUCOT, A. J.: Phanerozoic extinctions: How similar are they to each other? ....................................... 5
,LLEHCTIK J. A.: Biological selectivity of extinction .................................................................................. 31
LEARY, P. N. & RAMPINO, M. R.: A multi-causal model of mass extinctions: increase in trace
metals in the oceans ................................................................................................................................. 45
LEARY, P. N. & HART, M. B.: Important considerations in the investigation of global bioevents ...... 57
RICE, A.: Shock pressures in igneous processes: Implications for K/T events ....................................... 59
WILDE r P., QUINBY-HUNT, M. S. & BERRY, W.B.N.: Vertical advection from oxic or anoxic
water from the main pycnocline as a cause of rapid extinction or rapid radiations ........................ 85
YANG, Z.: An astronomical explanation of anomalous concentrations of iridium element
during catastrophic extinctions ............................................................................................................... 99
PALAEOZOIC EVENTS
BIERNAT, G. & ,KYZCRANDEB W.: Evolutionary crisis within the Ordovician
acrotretid inarticulate brachiopods of Poland .................................................................................... 105
BERRY, W. B. N., WILDE, P. & OUINBY-HUNT, M. S. : Late Ordovician graptolite mass
mortality and subsequent Early Silurian re-radiation ........................................................................ 115
BOUCOT, A. J.: Silurian and pre-Upper Devonian bio-events ................................................................ 125
TRUYOLS-MASSONI, M., ,SONISETNOM R., GARCIA-ALCALDE, J. L. & LEYVA, F.: The Kacak-
otomari event and its characterization in the Palentine domain (Cantabrian Zone, NW Spain) 331
LOTrMANN, J.: The Middle Givetianpurnilio-Events - A tool for high time resolution
and event-stratigraphical correlation ................................................................................................... 541
,RELDNIHCS E.: The late Frasnian (Upper Devonian) Kellwasser Crisis ............................................. 151
MC GHEE, G. R. Jr.: The Frasnian-Famennian mass extinction record in the eastern ....................... 161
United States
RACKI, G.: Frasnian/Famennian event in the Holy Cross Mts, Central Poland:
Stratigraphic and ecologic aspects ....................................................................................................... 169
KALVODA, J.: Late Devonian - Early Carboniferous paleobiogeography of benthic
Foraminifera and climatic oscillations ................................................................................................. 183
,VOKAMIS K. V.: Major evolutionary events among the spiriferids at the Devonian-
Carboniferous boundary ........................................................................................................................ 189
PALAEOZOIC/MESOZOIC EVENTS
,TREHCIET C.: The Permian-Triassic boundary revisited ........................................................................ 199
BENTON, M. J.: Mass extinctions in the fossil record of late Palaeozoic and Mesozoic tetrapods ...239
IV
MESOZOIC EVENTS
,YELTAHW R.: The relationship between extrinsic and intrinsic events in the evolution of
Mesozoic non-marine Ostracoda ......................................................................................................... 253
,MOLLOC C. J.: The taxonomic analysis of mass extinction intervals: an approach to
problems of resolution as shown by Cretaceous ammonite genera (global) and
species (Western Interior of the United States) ................................................................................ 265
HAR~ES, P. J. & KAUFFMAN, E. G.: Patterns of survival and recovery following the
Cenomanian-Turonian (Late Cretaceous) mass extinction in the Western Interior
Basin, United States ............................................................................................................................... 277
RoY, J. M., MCMENAMIN, M. A. S. S. & ALDERMAN, E.: Trophic differences, originations
and extinctions during the Cenomanian and Maastrichtian stages of the Cretaceous 299 ..................
JOHNSON, C. C, & KAUFFMAN, E, G.: Originations, radiations and extinctions of Cretaceous
rudistid bivalve species in the Caribbean Province ............................................................................ 305
MESOZOIC/CENOZOIC EVENTS
,TRAH M. B. & ,YRAEL P. N.: Periodic bioevents in the evolution of the planktonic foraminifera..325
CALDEIRA, K., RAMPINO, M. R., VOLK, T. & ZACHOS, J. C.: Biogeochemical modeling
at mass extinction boundaries: atmospheric carbon dioxide and ocean alkalinity
at the K/T boundary ............................................................................................................................... 333
FLEMING, R. F. & NICHOLS, D. J.: The fern-spore abundance anomaly at the Cretaceous-
Tertiary boundary: a regional bioevent in western North America ................................................. 347
NICHOLS, D. J., FLEMING, R. F. & ,NESKIREDERF N. O.: Palynological evidence of
effects of the terminal Cretaceous event on terrestrial floras in western North America 351 ............
KAUFFMAN, E. O., UPCHURCH, G. R. & NICHOLS, D. J.: The Cretaceous - Tertiary
boundary interval at south table mountain, near Golden, Colorado 365 ............ .................................. :
LANOLDA, M. A.: The Cretaceous-Tertiary boundary crisis at Zumaya (Northern Spain).
Micropaleontological data ..................................................................................................................... 393
,NESNAN T. A. & UPSHAW, B.: Aftermath of the Cretaceous-Tertiary extinction: Rate and
nature of the early Paleocene molluscan rebound ............................................................................. 401
SWAIN, F. M.: Species duration and extinction patterns in Cenozoic non-marine
Ostracoda, Western United States ....................................................................................................... 411
,ITSUGA J. & ,ALOS-AYOM S.: Mammal extinctions in the Vallesian (Upper Miocene) ................... 425
U ~ E NO~ A
HOW TO DEFINE "GLOBAL BIO-EVENTS"
contribution
to Proiect
GLOBAL
JI BIO -
,RESILL~.W Otto H. *) EVENTS
Abstract : Regarding the manifold pattern of global bio-events, the corresponding terms should not be
defined too restrictive.
I N T R O D U C T I O N : Science needs a common and vocabulary definitions of the used terms. But a
definition must not be necessarily extremely restrictive. Sometimes a relatively wide definition is more
adequate or even necessary, e.g. if the term has to describe a large variety of parameters or patterns or, as
another example, if the knowledge of the investigated subject is still on a low level. On account of these
reasons, the need of a wider definition also applies for those happenings which we call biological or
geologicale vents. Therefore I plead for a non-restrictive usage of the term 'global bio-event', as explained in
the following.
o G b 1 a I : Extinctions are serf-evidently global, even if the last representative of the extinct taxon lived in
a res~icted area for a longer time. Therefore, the problem is to prove whether the disappearance of a
certain taxon, recognized in a certain area, is due to local or regional mortality, to migration or to real
extinction. The disappearance of certain taxa in the Devonian often was thought to be only of local nature,
but later turned out to be a real extinction, i.e. global. Another, but even more important question in
connection with global bio-events is whether a globally distributed taxon disappears contemporaneously
elsewhere. Up to now, normally we only can confine the corresponding time interval of extinctions in the
order of a biozone, i.e. in general about 1 - 0,25 million years. In some other cases (e.g. the Kellwasser
Event = KW Event; see Schindler, 1990, this volume) the time interval of stepwise extinctions can be
globally restricted to the order of 100 ky and less. Even in the first mentioned case, we may call it
contemporaneous with regard to the geological time scale.
E v e n t : Of course, each change, disappearance, new occurrence, interruption, origination etc., i.e. each
happening, is an event. In connection with global biological or geological events we should use this term for
exceptional happenings which took place in a relatively short time span; 'relatively' in comparison with the
tong time intervals between the events. But what does exceptional mean? A few examples may help to
elucidate this: significant higher ex~nction rate or origination rate compared with the average rate of
background extinctions or background origination respectively; sudden changes of facies; intercalation of an
anoxic layer in a sequence with well developed communities.
Now I want to take up again the question of the relative shortness of an event. A tsunami-caused layer
(compare Lottmarm, 1990, this volume), or most probably also the main Iridium layer at the K/T boundary,
are products of an extremely short happening of days or evenh ours. In relation to that, the KW Event with
a duration of several 100 Ky is extremely long. However, comparingt he time-span of the KW Event with the
*) Institut und Museum fiir Geologic und Pal/iuntologie, Unlversit~it G~ttingen, Goldschmidt-Str. 3,
D - 3400 GiSttingen, F.R.G.
preceding Frasnian time, in which over several million years the changes have been much less, the time
interval oft he KW Event is relatively very short and justifies the term 'event'.
The above mentioned bio-events are connected with high extinction rates or even with mass extinctions.
As pointed out earlier (e.g. Walliser, 1986), an extinction event often is followed - after a certain time
interval - by radiations. If this phase is short in comparison with the following time of nomismogenesis
(during which a "normal" background rate of origination can be observed), we may call it an event, too, in
this case a r a d i a t i o n e v e n t. The sudden occurrence and spreading-out of a new biological
construction ("Banplan", e.g. the ftrst ceiling of bactritids, leading to goniatites) is an event, too, in this case
abiological innovation event.
Another difficulty with the term event may be demonstrated by the otomari Event (compare Truyols-
Massoni et al., 1990, this volume), which also has been called Kacak Event by House (1985). The
cephalopod limestone sequences of the Rheiulsche Schiefergebirge (Germany) show intercalated dark to
black sediments (shales and sometimes limestones), called Odershausen Formation. At the very base of that
formation occur Nowakia otornari (sensu lato; it is an early form) and Cabrieroceras sp., formerly called .C
rouvillei. The base of the Odershausen Fro. has been taken as the base of the upper Middle Devonian
("Givetian") int he pelagic realm since the beginning of this century.
On account of the observation, that the occurrence of "anoxic" sediments can be recognized world-wide
at this time level, I called it otornari or rouvillei Event. In the meantime I recognized the real C. rouvillei (v.
Koenen, 1886) (from the Montagne Noire, France) is a younger synonym to C. crispiforme (Kayser, 1879),
but not identical with the Cabrieroceras from the Odershansen Fro. C. crispiforrne occurs alreadyw ithin the
late australis Zone and disappears at the end of the kockelianus Zone. T k. kockelianus and a early form of
the P. ensensis (or P. xylus group, respectively) overlap in one limestone layer int he sections Bou Tchrafme
(SE-Morocco) and in the Eifel Mountains (Germany; Weddlge, 1988). On account of holostratigraphic
considerations this layer represents a time-spani nt he order of 10 to 20 Ky. In both mentioned regions, also
the end of the "anoxic" intercalation is time-equivalent: P. hemiansatus occurs very short after the first
limestone layer above, the black shales.
What is the otornari Event? Is ita t position a of Fig. 1 (corrsponding to Truyols-Massoni et al., 1990, this
volume), or at b or c, or is it the time-span between, e.g., b and c? The original intention was to designate
the beginning of the black shales (b in Fig. 1) as 'otomari Event', most probably being connected with a
transgressive pulse. Of course, also the end of the black shale (c in Fig. I) is not only a lithological, but also
a biological event, which could be named separately. Current investigations on the changes between
australis and late ensensis Zones are promising in respect to a better knowledge of the taxonomy and ranges
of the involved taxa. Therefore, closer definitions will be made after completion of these investigations.
Until then the terra 'otomari Event' should be used in the mentioned original, not restricted sense.
C r i s i s : If the extinction rate is higher than the rate of originations over a longer period of time, we
shouldn't call this process an event but rather a crisis. Again we shouldn't make too restrictive definitions,
because there are all possibilities and transitions between a relatively short crisis and a relatively long event.
A good example again is the KW Event (Schindler, 1990, this volume): It is preceded by a crisis, lasting
several million years; the KW Event itself has a stepwise pattern, i.e. it can be subdivided in several special
events, of which the last, culminating one, may have been extremely short. The sequence 'crisis - stepwise
event - final culmination' seems to occur often ine arth history such as at the end oft he Cretaceous.
Category/Order: The fact that globaelv ents
of quite different impact on biosphere or sedimentary
regime appeared in earth history, allows us to distin-
guish different categories or orders of events. Boucot
(1990, this volume) lists such different orders, fortu-
nately without giving strict def'mitions. Fortunately,
because as soon as we would base the def'mitious on
~h
exact percentages of extinctions or originations per
time-unit, we would no longer be able to rank the
0
events, unless having reached an utopian high
standard of knowledge.
0 0 ~ 0 ~
0 ~ 0
" ':<C
Figure 1 : Range of fossils near the
iramoto Event (for explanation see text).
REFERENCES
Boucot, A. J. (1990): Phanerozoic extinctions: How similar are they to each other? - This volume.
House, M. R. (1985): Correlation of mid-Palaeozoic ammonoid evolutionary events with global sedimentary
perturbations. - Nature, 313, 17-22.
Lottmann, J. (1990): The Middle Givetian pumilio-Events - A tool for high time resolution and event-
stratigraphical correlation. - This vohune.
Schindler, E. (1990): The late Frasnian (Upper Devonian) Kellwasser Crisis. - This volume.
Truyols-Massoni, M., Montesinos, R., Garcia-Alcalde, J. L. & Leyva, F. (1990): The Kacak-Otomari event
and its characterization in the Palentine Domain (Cantabrian Zone, NW Spain). - This volume.
Walliser, O. H. (1986): Towards a more critical approach to bio-events. - In: Walliser, O. H. (ed.): Global
Bio-Events. A Critical Approach. - Lecture Notes in Earth Sciences, 8, 5-16.
Walliser, O. H. (1988): Proposal for an Eifelian/Givetian Boundary Stratotype. - Document submitted to the
Subcommission on Devonian Stratigraphy (ICS, IUGS), Rennes,A ugust 1988, 4 pp.
Weddige, K. (1988): Eifel Conodonts. - In: Ziegler, W. (ed.): 1st International Senckenberg Conference and
5th European Conodont Symposium (ECOS V), Contributions I. - Courier Forsch.-Inst.
Senckenberg, 102, 103-110; Frankfurt.
? N
PHANEROZOIC EXTINCTIONS: HOW SIMILAR ARE A contribut;on
to Project
THEY TO EACH OTHER?
GLOBAL
II BIO
-
EVENTS
,TOCUOB Arthur J. *)
Abstract: A brief account of the major Phanerozoic extinction events suggests that each one has a flavor all
its own. Major extinction events involving varied combinations of different taxonomic levels, such as many
classes, or many orders, or many families, and the varied impacts on community structure in varied
combinations are what is referred to here by "flavor". Most of the evidence discussed is taken from the
relatively well sampled, marine, level bottom environment. Recent evidence developed by Kauffman and a
few others suggests that extinctions take place over a geologically brief interval of time in which more
stenotopic taxa are eliminated, followed by an abrupt termination of the more abundant, higher biomass
eurytopes, during community collapse. Adaptive radiations too, appear to take place during a geologically
brief interval (up to a few million years) rather than "instantaneouslf' in ecologic time (less than a thousand
years). This suggests that "instantaneous" extra-terrestrial events are not likely to have influenced the
gradual phase of either extinctions or adaptive radiations in any very important manner, unless they are in
need of ecological amplification in order to be recognized. Long term extra-terrestrial events that
potentially resulted in terrestrial climatic gradient changes might have been significant. However, the more
instantaneous community collapse, high biomass effects, might involve other factors, including certain types
of extra-terrestrial events. The decoupling, non-coincidence in time of the extinction events and adaptive
radiations in varied parts of the global ecosystem,s uch as the marine and nonmarine environment, does not
support a concept of "instantaneous" extra..terrestrial influences. It is concluded that we still have little
understanding of the specifics involveidn any major extinction event or adaptive radiation on land or sea.
Many of our paleontologic e u m geologic data have been collected in such an inadequate manner as to
make new sampling mandatory if we are to attain a better understanding of these problems.
"Those who cannot remember the past
are condemned to repeat it."
George Santayana (The Life of Reason)
INTRODUCTION
D'Orbigny (1849-1852, 1850-1852) compiled the original biostratigraphic synthesis that made him the
Father of Biostratigraphy, and provided the foundation for future studies on extinctions and adaptive
radiations.F ollowing Cuvier, d'Orbigny employed a catastrophist interpretation (1849-1852, 2: 252-254) of
the biostratigraphic facts. We no longer subscribe to this view, for which many paleontologists have
substituted a neocatastrophist-evolutioantatriyt ude. However, we are still faced with trying to understand
what are basically the same data.
Most of the major extinction events have, of course, been recognized for a century or more (see Newell,
1956, 1963, 1967, for incisive summaries). The relative ease with which the earlier paleontologists and
biostratigraphers had recognized the major extinctions was largely based on the presence of major biofacies
and biomass (communities of modern parlance) changes across the boundaries combined with significant
taxonomic changes; there was much more to this recognition process than a mere change of taxa at the
boundaries (Newetl, 1967). After most of the Treatise on Invertebrate Paleontology and Osnovy
Paleontologii volumes became available, as well as "The Fossil Record" (Harland et al., 1967), it was
*) Department of Zoology, Oregon State University, Corvallis, OR 97331-2914, U.S.A.
