Causes of massive biotic extinctions and Yvonne Herman

Department of Geology
explosive evolutionary diversification Washington State University
Pullman, Washington 9 9 1 6 4
throughout Phanerozoic time

ABSTRACT
Massive extinctions and explosive evolutionary radiations are caused
principally by rates of change and secondarily by the magnitude, duration,
and spatial extent of phenomena; the phenomena themselves may vary
from extinction to extinction. Sudden changes over large areas result in
massive biotic extinctions; gradual changes permit organisms to adapt
behaviorally by migration and biochemically or physiologically by
alterations in their cellular or molecular makeup and result in adaptive
evolutionary diversification.

INTRODUCTION others, 1980; Smit and H e r t o g e n , 1980); and (17) e n o r m o u s

Massive extinctions and bursts in e v o l u t i o n a r y diversifications stellar explosions ( T u c k e r , 1977).
have interrupted the otherwise g r a d u a l and c o n t i n u o u s course of S o m e of these processes recurred t h r o u g h o u t geologic time,
evolution and extinction of o r g a n i s m s t h r o u g h o u t P h a n e r o z o i c but only i n f r e q u e n t l y can they be linked with massive biotic
time ( S i m p s o n , 1944) [or " p u n c t u a t e d " processes (Gould a n d extinctions. For e x a m p l e , a l t h o u g h oceanic t e m p e r a t u r e s were
Eldredge, 1977, and references therein)]. T h e most sweeping lower d u r i n g M i o c e n e and Pliocene time than d u r i n g Oligocene
extinction, m a r k i n g the Permian-Tria.ssic b o u n d a r y - 2 3 0 m . y . time (Savin, 1977, a n d references cited therein), b o t h the
ago, resulted in the elimination of m a j o r g r o u p s of m a r i n e and
land o r g a n i s m s (Newell, 1967; Valentine, 1969). O t h e r massive
extinctions a f f e c t i n g large n u m b e r s of unrelated terrestrial and
o p e n - o c e a n taxa were recorded at the C r e t a c e o u s - P a l e o c e n e
b o u n d a r y , followed by the drastic Eocene-Oligocene reduction in
diversity (Newell, 1967; Valentine, 1969; Fig. 1). Similar although
less severe biological crises are k n o w n to have occurred in Late
C a m b r i a n , Late O r d o v i c i a n , Late D e v o n i a n , a n d Late Triassic
time ( f o r e x a m p l e , Newell, 1967; Valentine, 1969; J o h n s o n and
B o u c o t , 1973). Various hypotheses have been a d v a n c e d to
a c c o u n t f o r these d r a m a t i c episodic biological events. Extinctions
have been a t t r i b u t e d to (1) drastic cooling (for example, H a y ,
1960; Axelrod and Bailey, 1968); (2) cooling followed by a rise
of calcium c a r b o n a t e c o m p e n s a t i o n d e p t h ( C C D ) (Worsley,
1971); (3) unusually high t e m p e r a t u r e s (Urey, 1973; W a t e r h o u s e ,
1973; Emiliani, 1980); (4) hyposaline oceans (Fischer, 1964;
Stevens, 1977); (5) hypersaline oceans (Bowen, 1968, 1970;
W a u g h , 1973); (6) m a r i n e regressions (Newell, 1967; J o h n s o n ,
1974); (7) collapse of the f o o d web by reduction of m a r i n e
p h y t o p l a n k t o n ( T a p p a n , 1968); (8) decreased nutrient supply
f r o m land (Bramlette, 1965; B u d y k o , 1977); (9) injection of
Arctic brackish water into the world ocean ( G a r t n e r and Keany,
1978; Thierstein a n d Berger, 1978); (10) c o n t i n e n t a l suturing
Figure 1. Sudden changes in environment result in massive biotic
(Valentine and M o o r e s , 1970); (11) increase in tropic resource
extinctions, whereas gradual changes result in rapid evolutionary
f l u c t u a t i o n s (Valentine, 1973); (12) chemical poisoning ( C l o u d , diversification. Solid line shows change in sea level due to change in
1959; H s u , 1980); (13) increased cosmic radiation ( H a t f i e l d and ridge volume for period from 8 5 to 15 m.y. B.P. It is assumed that by
C a m p , 1970); (14) variations in c o n c e n t r a t i o n of a t m o s p h e r i c 15 m.y. B.P. glacial build-up in Antarctica may have been sufficient that
oxygen (McAlester, 1970); (15) reversals in E a r t h ' s magnetic fluctuations in glacial mass became dominant mechanisms causing sea-
field ( U f f e n , 1963; H a r r i s o n , 1968; H a y s , 1971); (16) i m p a c t of level changes. Dashed line gives position of shoreline relative to hinge
line as a function of rate of sea-level change (after Pitman, 1 9 7 8 ) . Dotted
large meteorites, c o m e t s , or tektite showers (De L a u b e n f e l s ,
vertical line with arrow denotes extinctions; solid vertical line with
1956; M c L a r e n , 1970; U r e y , 1973; O ' K e e f e , 1980; Alvarez a n d arrow denotes rapid evolutionary diversification.

104 G E O L O G Y , v. 9 . p. 1 0 4 - 1 0 8 , M A R C H 1981
Miocene and the Pliocene were periods of rapid evolutionary
radiation of m a r i n e organisms, whereas during the Oligocene
m a j o r reductions in species diversity occurred ( T a p p a n and
Loeblich, 1973). While continents f r a g m e n t e d f r o m Jurassic time
on (Valentine and M o o r e s , 1972), massive biotic d e c i m a t i o n s
occurred episodically—for example, at the C r e t a c e o u s - P a l e o c e n e
and the Eocene-Oligocene b o u n d a r i e s ( T a p p a n and Loeblich,
1973, and references therein). Regressions a f f e c t i n g continental
margins have been n u m e r o u s ( D a m o n , 1971), b u t only a few
have coincided with massive extinctions ( S c h o p f , 1974; J o h n s o n ,
1974).
Explosive diversifications c o m m o n l y a p p e a r to follow
massive extinctions (Newell, 1952). T h e most impressive are
those of the Devonian and Jurassic-Cretaceous for the
shelf biota (Newell, 1952, 1967) and those of the Devonian-
C a r b o n i f e r o u s , C r e t a c e o u s , Eocene, and Neogene for m a n y
p l a n k t o n i c o r g a n i s m s (Lipps, 1970; T a p p a n and Loeblich,
1973; H e r m a n , 1979). E v o l u t i o n a r y diversification has been
attributed to t h e d e v e l o p m e n t of diverse h a b i t a t s by continental
f r a g m e n t a t i o n (Valentine and M o o r e s , 1972), changes in land
g e o g r a p h y and in oceanic circulation, increased climatic z o n a t i o n ,
and decreased t e m p e r a t u r e s (Valentine, 1968; Lipps, 1970), as
well as to higher and m o r e u n i f o r m oceanic t e m p e r a t u r e s
(Fischer and A r t h u r , 1977), m o r e stable physical c o n d i t i o n s
( T a p p a n and Loeblich, 1973), and reversals in E a r t h ' s magnetic
field ( S i m p s o n , 1966). T h e geologic record provides evidence
that e v o l u t i o n a r y diversification occurred when some of the
p h e n o m e n a listed a b o v e took place (Valentine, 1968; Lipps,
1970; T a p p a n and Loeblich, 1973; Valentine and M o o r e s , 1972;
Whyte, 1977), but not in all instances.
MASSIVE E X T I N C T I O N S
The most severe a n d / o r the best d o c u m e n t e d biotic
extinctions are the Permian-Triassic, C r e t a c e o u s - P a l e o c e n e ,
and Eocene-Oligocene.
Terminal Permian Environments
Vast areas of the P a n g e a n s u p e r c o n t i n e n t , which lay
near the South Pole during m u c h of Paleozoic time, glided into
the w a r m mid-latitudes (Crowell and Frakes, 1970; Frakes, 1979).
T h e drift of the f r a g m e n t e d supercontinents was followed by
their reassembly, with a t t e n d a n t regression f r o m most conti-
nental p l a t f o r m s (Valentine and M o o r e s , 1970; J o h n s o n , 1974;
B o u c o t , 1975), owing mainly to cessation or decreased rates of
sea-floor spreading and collapse of ridges (Valentine and M o o r e s ,
1970, 1972; S c h o p f , 1974). The sudden global w a r m i n g (King,
1961; Menzies and others, 1973; Frakes, 1979) and substantially
lowered surface-water salinities (Stevens, 1977) in Late P e r m i a n
time m a y have been responsible for the a b r u p t elimination of
vast h a b i t a t s and consequently of their biota, although chemical
poisoning should have had similar effects ( C l o u d , 1959). This
m a j o r biotic crisis, which a f f e c t e d mainly shallow-water
o r g a n i s m s (Newell, 1952; Valentine and M o o r e s , 1972; J o h n s o n ,
1974) but also land animals and plants (McAlester, 1973),
occurred at a time when global sea levels stood very low
( S c h o p f , 1974; J o h n s o n , 1974).
Late Cretaceous Environments
T h e replacement of latitudinal t h e r m a l gradients by u n i f o r m
warm global t e m p e r a t u r e s (King, 1961; Menzies and others,
1973; M c L e a n , 1978) and a sudden w a r m i n g pulse at the
M a e s t r i c h t i a n - D a n i a n b o u n d a r y are indicated in both terrestrial
(Colbert and o t h e r s , 1946; C o l b e r t , 1965; H u g h e s , 1976;
M c L e a n , 1978) and m a r i n e studies. Oxygen-isotope analyses of
GEOLOGY
benthonic and p l a n k t o n i c f o r a m i n i f e r a and calcareous n a n n o -
fossils (Kroopnick and others, 1977) and belemnites (Stevens
and C l a y t o n , 1971), as well as the distribution of larger w a r m -
water f o r a m i n i f e r s , coral reefs, and rudists (Voight, 1964), and
chemical evidence ( J o r g e n s e n , 1975) indicate a late Maestrichtian
warming, although this interpretation has not been u n a n i m o u s
(Russell, 1970). Regression of continental-shelf water coincided
with the terminal C r e t a c e o u s w a r m pulse. This water with-
drawal, like the Permian-Triassic event, is a t t r i b u t e d to mid-
ocean ridge c o n t r a c t i o n (Valentine and M o o r e s , 1970, 1972;
S c h o p f , 1974; P i t m a n , 1978). Decreased nutrient supply f r o m
land to the ocean, as a consequence of lowered continental relief
and aridity, has been advocated by several a u t h o r s (for example,
Bramlette, 1965; B u d y k o , 1977). It has been postulated that
density stratification in the world ocean was enhanced by the
u n i f o r m global t e m p e r a t u r e s (Bramlette, 1965). C o n s e q u e n t l y ,
nutrient recycling to the surface-water layer, which today occurs
mainly by upwelling and d i f f u s i o n , was p r o b a b l y minimal at that
time (Bramlette, 1965).
Several a u t h o r s have suggested that the terminal C r e t a c e o u s
extinctions could have been triggered by a spillover of cold,
brackish water f r o m the Arctic into the world ocean via the
Atlantic ( G a r t n e r and Keany, 1978; Thierstein and Berger, 1978)
when G r e e n l a n d was separated f r o m E u r o p e by sea-floor
spreading. A l t h o u g h this idea is interesting, it is based on several
basic a s s u m p t i o n s that need f u r t h e r e v a l u a t i o n — n a m e l y , that
the Arctic was filled with brackish or fresh water and that this
low-salinity water gushed into the Atlantic, whence it spread
into the other oceans, killing p h y t o p l a n k t o n and z o o p l a n k t o n ,
and that sea-floor spreading was relatively rapid. H o w e v e r ,
according to Sclater and others (1977), spreading rates prior to
- 5 4 m . y . B . P . were very slow. Recent discovery of open-ocean
calcareous n a n n o f o s s i l s (Worsley and H e r m a n , 1980; H e r m a n
and B a c k m a n , in p r e p . ) and of silicoflagellates (Ling and others,
1973) of Late C r e t a c e o u s (Maestrichtian) age in Central Arctic
basin sediments suggests that salinities were at least near
" n o r m a l " sea-water values. F u r t h e r m o r e , i m p o r t a n t deductions
concerning C r e t a c e o u s global surface-water paleocirculation
patterns during nonglacial conditions were m a d e by Luyendyk
and others (1972) on the basis of l a b o r a t o r y experiments. T h e
trans-Arctic surface-water flow was shown to be f r o m the
Atlantic via the pole to the Pacific (Luyendyk and others, 1972;
H e r m a n , 1979; Fig. 2). No flow of Pacific water via the Arctic
into the Atlantic Ocean was observed (Luyendyk and others,
1972). For the reasons outlined, an Arctic fresh-water spillover
into the Atlantic does not seem likely, although this possibility
need not be ruled o u t .
A n o t h e r hypothesis a b o u t massive extinctions involves the
impact of a meteorite or c o m e t . This idea (De L a u b e n f e l s ,
1956) has been revived by M c L a r e n (1970), Urey (1973), Alvarez
and others (1980), Smit and H e r t o g e n (1980), Hsu (1980), and
Emiliani (1980). T h e effect of such collision would have been
catastrophic to the biota of the entire planet. Chemical analysis
of m a r i n e sediments has shown a n o m a l o u s l y high levels of
iridium and o s m i u m , considered to indicate influx of extra-
terrestrial material at the Cretaceous-Tertiary b o u n d a r y in New
Z e a l a n d , D e n m a r k , several sections near G u b b i o , Italy (Alvarez
and others, 1980), and C a r a v a c a , Spain (Smit and H e r t o g e n ,
1980). This trace-element enrichment was believed by these
a u t h o r s to s u p p o r t the idea of a meteorite collision. Finn Surlyk
geologists (1980) q u e s t i o n e d the validity of this hypothesis:
" S i n c e the relevant analytical techniques are quite c u m b e r s o m e ,
relatively few d e t e r m i n a t i o n s of Ir and related trace-elements are
to be f o u n d in the literature . . . the question t h u s arises: how
105

Figure 2. Reconstruction of continents and oceans in Cretaceous
time and inferred surface-water paleocirculation patterns (after
Herman, 1 9 7 9 ) .
m a n y analyses have to be u n d e r t a k e n in order to assess with
certainty that an o c c u r r e n c e is a n o m a l o u s ? " F u r t h e r , Wezel
(1979) published the results of detailed investigation of the
G u b b i o sequence, in the key section at Bottaccione, which is
interpreted to represent an essentially c o n t i n u o u s sequence of
C r e t a c e o u s - P a l e o c e n e calcareous pelagic sediments by Alvarez
a n d o t h e r s (1977). A c c o r d i n g to Wezel (1979), these p r e s u m e d
pelagic sediments are t u r b i d i t y - c u r r e n t deposits with very few
indigenous fossils. It is i m p o r t a n t to d e t e r m i n e whether other
sections believed to represent c o n t i n u o u s C r e t a c e o u s - P a l e o c e n e
sequences are indeed u n i n t e r r u p t e d .
D r o u g h t and s u d d e n increase in t e m p e r a t u r e on land are,
I believe, the principal cause of d i n o s a u r extinction ( M c L e a n ,
1978; Bramlette, 1965; B u d y k o , 1977; H s u , 1980).
T h e c a r b o n - i s o t o p e record of surface-dwelling p l a n k t o n i c
f o r a m i n i f e r a indicates t h a t they were depleted in 13 C d u r i n g
Pleistocene c o l d - t e m p e r a t u r e m i n i m a ( S h a c k l e t o n , 1977), times
of extreme land aridity. If Shackletori's a s s u m p t i o n that n C
variations in t h e calcific shells of f o r a m i n i f e r a reflect changes in
land vegetation is correct, then the strong 13 C depletion at the
C r e t a c e o u s - P a l e o c e n e b o u n d a r y a n d d u r i n g the early Paleocene
(Savin, 1977; Thierstein and Berger, 1978) may be indicative of
extreme aridity on c o n t i n e n t s . W h a t e v e r the initial causes, the
effect was mass d e c i m a t i o n of o p e n - o c e a n a n d , to a lesser
extent, shelf biota, as well as of d i n o s a u r s , when the critical
threshold of their tolerance to t h e e n v i r o n m e n t was crossed.
C u r r e n t estimates of t h e d u r a t i o n of the events that b r o u g h t
a b o u t the massive extinctions vary f r o m essentially i n s t a n t a n e o u s
106
to 104 to 10 5 yr (Kent, 1977; Van Valen and S l o a n , 1977;
Russell, 1979, a n d review d a t a therein; Smit and H e r t o g e n , 1980).
Terminal Eocene Environments
T h e drastic r e d u c t i o n in species diversity of shell-bearing
p l a n k t o n ( T a p p a n and Loeblich, 1973, and references therein)
and of early land m a m m a l s (Stokes, 1966) at the Eocene-
Oligocene b o u n d a r y (Fig. 1) h a p p e n e d at a time of sudden
regression ( H a l l a m , 1963; D a m o n , 1971; Vail and others, 1977).
T h e sea-water retreat f r o m continental shelves was t h o u g h t to
coincide with t h e collapse of spreading ridges (Valentine and
M o o r e s , 1970, 1972) a n d with the expansion of the A n t a r c t i c
ice sheet. T h e cooling was alternatively a t t r i b u t e d to tektite
showers ( O ' K e e f e , 1980). Aridity on land (Stokes, 1966; F r a k e s ,
1979) beginning in middle E o c e n e time, coupled with the s u d d e n
t e m p e r a t u r e decline, was p r o b a b l y responsible for the
elimination of early m a m m a l s ( M a t t h e w s , 1962).
EVOLUTIONARY DIVERSIFICATION
T h e geologic record provides evidence that bursts in
evolutionary r a d i a t i o n occurred when e n v i r o n m e n t a l c h a n g e s
took place gradually over extended periods such as those of the
C r e t a c e o u s or the N e o g e n e ( T a p p a n and Loeblich, 1973), when
continental f r a g m e n t a t i o n was proceeding at a slow and even
rate, latitudinal t e m p e r a t u r e z o n a t i o n was increasing (King,
1961; Menzies and others, 1973; F r a k e s , 1979), a n d sea level
was c h a n g i n g gradually (Vail a n d others, 1977), c o n d i t i o n s that
allowed o r g a n i s m s to keep pace with and be able to a d a p t to the
changing e n v i r o n m e n t s .
E v o l u t i o n a r y diversification took place while sea level rose
gradually d u r i n g the Jurassic P e r i o d , and it also occurred while
sea level was d r o p p i n g d u r i n g the Miocene E p o c h (King, 1961;
Menzies and others, 1973; Vail and others, 1977; F r a k e s , 1979).
T e m p e r a t u r e s were a p p a r e n t l y increasing d u r i n g Early and
Middle Jurassic time, as well as d u r i n g the Early C r e t a c e o u s , but
were declining t h r o u g h o u t the N e o g e n e (Menzies and others,
1973; Fischer and A r t h u r , 1977; P e a r s o n , 1978; F r a k e s , 1979).
H o w e v e r , b o t h the Mesozoic a n d Neogene represent times of
rapid increase in n u m b e r s of t a x o n o m i c units. O r o g e n i c move-
m e n t s were u n d e r w a y in t h e Early C a m b r i a n , Silurian,
D e v o n i a n , P e r m i a n , C r e t a c e o u s , a n d Neogene, and quiescent
periods prevailed d u r i n g most of the C a m b r i a n , O r d o v i c i a n , and
Triassic ( D a m o n , 1971; J o h n s o n , 1971), all times of e v o l u t i o n a r y
diversification ( S i m p s o n , 1966; Newell, 1967).
DISCUSSION A N D CONCLUSIONS
T h r o u g h o u t geologic time, o r g a n i s m s have developed
d i f f e r e n t strategies to cope with and a d a p t to the various
e n v i r o n m e n t a l stresses. A d a p t a t i o n can be achieved behaviorally,
m o r p h o l o g i c a l l y , physiologically, or biochemically ( S o m e r o a n d
H o c h a c h k a , 1976). T h e simplest m o d e is b e h a v i o r a l ; it involves
m i g r a t i o n to a region where c o n d i t i o n s are m o r e f a v o r a b l e . T h e
m o r e complex a d a p t i v e strategies c o m p r i s e changes at t h e
cellular a n d / o r molecular level. D u r i n g the last few million
years, E a r t h has been subjected to climatic f l u c t u a t i o n s of
variable a m p l i t u d e s and d u r a t i o n s . These climatic c h a n g e s
characterizing the late C e n o z o i c ice ages were a p p a r e n t l y suf-
ficiently slow to permit n o t only animals but also most plants to
shift their d i s t r i b u t i o n a l ranges to regions with f a v o r a b l e
e n v i r o n m e n t s ( B u d y k o , 1977). T h e causes of late W u r m i a n
( W i s c o n s i n a n ) extinction of herbivores a n d large p r e d a t o r s
outside the tropical regions are still not k n o w n , a l t h o u g h climatic
changes and e x t e r m i n a t i o n s by primitive m a n have been suggested
as possibilities ( B u d y k o , 1977).
M A R C H 1981
Causes of Mass Extinctions and Rapid Evolutionary
Diversification
F r o m t h e vast l i t e r a t u r e t h e f o l l o w i n g c o n c l u s i o n s c a n b e
d r a w n : of t h e several e l e m e n t s t h a t p l a y c r i t i c a l r o l e s in t h e f i n a l
o u t c o m e of e v e n t s , t h e m o s t i m p o r t a n t a p p e a r s t o be t h e rate of
change, f o l l o w e d b y t h e m a g n i t u d e , d u r a t i o n , a n d s p a t i a l e x t e n t
of p h e n o m e n a o r p r o c e s s e s . W h e n t h e r a t e s of e n v i r o n m e n t a l
c h a n g e a r e r a p i d a n d t h e m a g n i t u d e , d u r a t i o n , a n d e x t e n t of
p h e n o m e n a a r e g r e a t e r t h a n t h e a b i l i t y of o r g a n i s m s t o c o p e
with a n d a d a p t to t h e m , extinctions result. Conversely, when
t h e r a t e s of a d a p t a t i o n of o r g a n i s m s c a n k e e p p a c e w i t h o r
exceed t h e r a t e s a t w h i c h c h a n g e s in t h e e n v i r o n m e n t o c c u r ,
adaptive evolutionary radiations ensue.
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ACKNOWLEDGMENTS
Reviewed by R. S. Dietz, C. L. Drake, J. N. T h o m p s o n , J. W.
Valentine, and P. R. Vogt. I t h a n k t h e m and W. A. Berggren, H. D.
Holland, J. W. Mills, N. D. Newell, and P. E. Rosenberg, w h o read an
earlier version of t h e manuscript and provided valuable suggestions.
MANUSCRIPT R E C E I V E D AUGUST 20, 1980
REVISED MANUSCRIPT R E C E I V E D DECEMBER 8, 1980
MANUSCRIPT ACCEPTED DECEMBER 18, 1980
MARCH 1981