Xu
Cui
Whether
periodicity exists in extinction process has long been debated since 1984 when
Raup and Sepkoski first raised it. The debate has focused on the extinction
database, the definition of mass extinction and the statistical methods. Much
more data are needed before the debate is resolved. At the same time several
extinction mechanisms, mostly extraterrestrial ones, had been put forward. But
none seems to well explain the dynamics of extinction.
In
the history of life a vast majority of species that have ever lived have been
extinct1,2, and the pattern and process of extinction is still in
controversy3. One of the suggested patterns is that extinction
happens periodically, first raised by Fisher and his student Arthur of Princeton
University in 1977 in a virtually ignored paper4. They postulated a
32-million-year periodicity in extinction basing on data culled from the fossil
record of the past 250 million years. However, their study was not supported by
a rigorous statistical analysis and
the database itself was limited2,4,5, leaving their postulation
dubious. They also guessed that cycles of convection within the Earth, though
still little understood, drove the extinction process4.
The
suggestion was reexamined seven years later. In 1984 Raup and Sepkoski of
University of Chicago advanced a hypothesis of a 26-million-year periodicity in
extinction (published in PNAS2) which arrived in the heat of the
debates on whether mass extinction was caused by extraterrestrial impacts
triggered by Alvarez, et al 6 in 1980. The study of Raup and
Sepkoski was based on a pioneering statistical analysis of the record of 3,500
families of marine animals (vertebrate, invertebrate, and protozoan) and thus
arrested the attention of scientists, and triggered new research in broad
fields, particularly in astronomy and astrophysics7-12. More
importantly, they also triggered a heated debate that does not end now:
periodicity exists in extinction?
Periodic
extinction?
First
of all, why the Raup and Sepkoski 1984 paper2 is so controversial? In
the paper, Raup and Sepkoski investigated the temporal distribution of the 12
major extinctions, defined as the peaks (local maximum) in extinction intensity
– time diagram, over the past 250 million years statistically using various
forms of time series analysis: Fourier analysis, standard autocorrelation
analysis and nonparametric testing. The database came from Sepkoski’s
compilation of the temporal ranges of about 3,500 families of marine animals,
mainly based on Harland time scales. The 12 events showed statistically
significant periodicity and so they concluded, “It seems inescapable that the
post-late Permian extinction record contains a 26-million-year periodicity,
assuming that the Harland time scale is a reasonable approximation of
reality.”2 The paper left 4 open questions: 1. What if the definition of
“major extinction” changes? 2. The statistic method is reliable? 3.
The Harland time scale is reasonable? 4. The database is reliable? What if the
database is expanded to continental organisms and longer geological time? These
unsolved questions led to many objections to their periodicity conclusion.13,14.
Just
two months later Hallam13 gave some reasons why open-mindedness on
the problem of periodic extinction was necessary. First he questioned that the
accuracy of Harland time, and then he regarded only 5 of the 12 major
extinctions defined by Raup and Sepkoski were unquestionable. The other 7 were
either dubious themselves or minor extinctions. Thus he concluded that the
evidence given by Raup and Sepkoski hardly seemed warrant the conclusion of
periodicity13.
Then
in November of the same year Kitchell and Pena questioned the periodicity using
deterministic and stochastic methods14. They showed that in the past
250 million years extinction displayed a pseudoperiodic behavior with a cycle
length of 31 million years. And more, the periodicity weakens when the analysis
was extended to the entire Phanerozoic. They used a quite different analyzing
method from Raup and Sepkoski’s and first extended the analysis beyond 250
million years and thus the paper attracted much interest from other scientists.
The
database Raup and Sepkoski used was also challenged. Patterson and Smith39,40,
after checking the echinoderm and fish family extinctions which made up about
20% of the data, found that only 25% of their sample was signal and the 75%
noise component included pseudoextinctions of non-monophyletic groups. Sepkoski 41admitted
that data from echinoderm and fish failed to exhibit periodicity. But he argued
the data were only 25% of the entire marine extinction data and he did not treat
taxonomic families as real evolutionary entities in the analyses of extinction
patterns. Thus the periodicity conclusion did not fail.
Raup
and Sepkoski continued extending their analysis down to the more detailed
taxonomic level of genera, analyzing 11,000 of them17,18,19. As
Sepkoski20 said in 1990, the 26-million-year cycle, evident as peaks
in the extinction rate, appeared even more strongly than it did in the original
family data. What was remarkable was that almost at the same time, a new
extinction peak, recognized in the same year in the Middle Jurassic Period, fell
at a cycle time predicted by the periodicity hypothesis5.
However,
when Benton considered both marine and continental organisms, support for
periodicity of mass extinctions was not found42. But Benton did not
use vigorous statistical methods and thus the question was still left open.
Hoffman
of the University of Warsaw seriously disputed the data culling by Raup and
Sepkoski, the definition of mass extinction, dating of extinction events and
statistical methodology15. As Hoffman argued, because Raup and
Sepkoski used “peaks” as the definition of mass extinctions, it was
necessary that the number of families extinguished should be greater than in the
preceding and succeeding stages. This effect was to exclude the possibility that
consecutive stages could be associated with mass extinction, which was bound to
prejudice the analysis towards a periodic interpretation. If the probabilities
of extinction in successive stages were entirely independent, simply looking for
peaks in the record of extinction implied a one-in-four chance that any stage
would be recognized as a time of mass extinction and, given the exclusion of the
possibility that consecutive stages may be thus described, the analysis was
certain to yield the conclusion that, on the average, extinction peaks occur
every four stages. This, said Hoffman, was the origin of Raup and Sepkoski’s
periodicity15,16.
At
the same time Maddox 33 suggested, “Defining a mass extinction is a
more urgent need.” Hoffman argued that some objective criterion was needed and
he would prefer a criterion based on the rate or probability of extinction per
unit time (say a million years) rather than one based on counting the number of
family extinctions in a stage, which he said was biased towards periodicity15.
Again in October 1985 when replying Lewin’s criticisms, he said, “it (mass
extinction defined by Raup and Sepkoski) accepts all peaks of extinction
intensity, and not only truly major events as mass extinctions. Why should we
wish to accept this particular-counterintuitive-definition rather than any
other?” 34
Raup
and Sepkoski argued17 that most of the extinction events defined in
the way they did2 had long been recognized as times of high
biological turnover and had been used since the mid-19th century to
define the major units of geologic time.
Stigler
and Wagner36 and Lutz 35 questioned the statistical method
used by Raup and Sepkoski. Stigler and Wagner36 held that regardless
of whether or not the data were periodic in origin (even regardless of the
actual period if they were in fact periodic), the nonparametric test used by
Raup and Sepkoski were substantially biased toward their periodicity conclusion.
This was because the test was sensitive to measurement error due to the
“Signor-Lipps effect”, early recording of extinctions due to missing fossil
specimens. Also because of the unequal spacing in time, models used by Raup and
Sepkoski might be expected to produce statistically significant but artifactual
periods. (But according to Benton et al37, the past 540
million years of the fossil record provided uniformly good documentation of the
life of the past if scaled to the stratigraphic level of the stage and the
taxonomic level of the family.) On the other hand, Lutz35 argued that
if episodic models were allowed as alternatives to the Poisson model, then no
geologic time series were sufficiently regular over a long enough time interval
to be convincingly periodic. At the same time he admitted that it was also not
possible to positively exclude periodicity based on statistics alone.
Raup
and Sepkoski 18 rebut that Stigler and Wagner ignored the geology of
the situation and ignored their treatment. The qualitative model used by Stigler
and Wagner was inappropriately applied over the entire time series and the
parameter value was extreme and thus unreasonable. Stigler and Wagner’s
response 38 was that they remained convinced that their conclusion
was correct and stated their evidence again.
The
question of periodicity was placed in a new light in 1987 by Shaw31.
In the pioneering paper, he used non-linear dynamics methods, which were
different from conventional statistical analysis, to detect new symmetrical
patterns and order in several earth processes. With respect to the periodicity
debate, he thought it impossible to demonstrate valid periodicity patterns,
because the data sets are too limited for both conventional and nonlinear
analyses. Things would be better if more statisticians became versed in
nonlinear-dynamics methods, but the “bridge between the two approaches to data
analysis is just beginning to be built”5.
Debate
still goes on. But it is obvious that in 1990s the debate cooled down. The
papers involved in the debate from 1984 to 1989 are twice as those in the 10
years of 1990s, as shown by SCI. Further, a majority of the papers in 1990s did
not take periodicity problem as their main subjects. This might due to the
futility of debate if data is limited.
The mechanisms of periodic extinctions
Mechanisms
of periodic extinctions were raised before the debate was settled down. It is
interesting that even before the objection of periodic extinctions arose,
several mechanisms of period extinctions had been put forward. This is partly
because Raup and Sepkoski’s suggestion in their original paper2.
They suggested that the causes of periodicity might be related to
extraterrestrial forces. Only two months later several scientists suggested
possible causes7-10. Alvarez and Muller indirectly supported the
extraterrestrial causes11. They subjected the records of large impact
craters to time-series analysis and concluded that most occurred on a
28.4-million-year cycle. However, because only 13 craters met their rigorous
criteria, their conclusion must be viewed with caution13. Jetsu et
al 43-45 also suspected the existence of periodicity in impact.
The
causes suggested until now can be summarized as19:
(1)
Orbital dynamics of an unobserved solar companion9,10.
(2)
Vertical oscillation of the solar system about the plane of the Galaxy7,8.
(3)
Transit of the solar system through the spiral arms of the Galaxy.
(4)
Precession of an undetected tenth planet 27.
(5)
Biogeochemical cycles, climate and sea-level changes5.
(1)
Davis et al.10 and Whitmire and Jackson9
independently put forward a model that postulated the existence of an unseen
companion star to the Sun, occupying a highly eccentric orbit. Beginning with
the assumption of an orbital periodicity of 26 to 28 million years, they
calculated a major axis of about 3 light year for the companion’s orbit and a
perihelion distance of 0.3-0.5 light year from the Sun. The estimated mass of
the companion was 10-3-10-1 solar mass. When near the
perihelion it is brought into the dense inner region of a comet cloud and by
perturbing the cometary orbits initiates and intense comet shower, leading to a
series of terrestrial impacts lasting up to a million years13.
This
hypothesis was soon be seriously challenged by the stability analysis of the
hypothesized companion.23-26. The analysis indicated that the orbital
periodicity should be somewhat irregular, varing over 10-20%, and that the
expected half life for the orbital configuration should be on the order of 1000
million years. Thus the companion was unstable. However, Perlmutter et al
30 argued that only the companion hypothesis was consistent with all
the known data. Until 1990, they had rejected about 20% of the 2,691 stars in
their list of candidates in the Northern Hemisphere and kept searching. However,
because until now the companion has not been observed despite considerable
searching by an automated telescope, the hypothesis probably failes1.
(2)
Schwartz and James8 and Rampino and Stothers7 proposed
galactic models13. The former authors speculate that long-term
changes in cosmic radiation flux, soft X-rays and hard UV radiation, due to the
Sun’s oscillation about the galactic plane have provoked sufficient
alterations of the biosphere (disturbing the ionization balance of the upper
atmosphere) to cause mass extinctions. Rampino and Stothers’ reanalysis of
Raup and Sepkoski’s data led them to suggest a extinction cyclicity of about
30 million years, which correlated strongly with a galactic cycle. They argued
that tidal forces from intermediate-sized molecular clouds of 103-104
solar masses, concentrated near the galactic plane, might perturb the Oort Cloud
and inner cometary reservoir as the solar system approached the plane. Such
perturbations could produce comet showers of up to several million years
duration, during which one large or several comets might impact the Earth7.
The
problem with these hypotheses is the disparity between the 33-million-year
half-period for solar oscillation and the 26-million-year period for extinction
events19. Rampino and Stothers7 argued that the data of
Raup and sepkoski2 fitted a 30-million-year period and, molecular
clouds were randomly distributed so there should be a fair amount of
stochasticity in extinction events so that the extinction period estimated from
a small number of events might not correspond precisely to the oscillatory
half-period. However, as argued by Sepkoski and Raup19, analysis
based on new data showed that a 30-million-year period was untenable as an
estimate of the extinction periodicity, leaving the disparity unresolved22.
Also, as stated by Bahcall, the apparent periodicity in the mass extinction and
cratering records could not be caused by a population of objects that
contributes a major fraction of the total mass density at the solar vicinity29.
(3)
As the solar system orbits the Milky Way Galaxy, it passes through spiral arms,
or density waves, approximately every 50 million years19.
Interstellar “planetismals” were captured by the sun during these transits
and this capture could increase “asteroid” bombardment of the Earth during
ensuing 20-30 million years intervals, as argued by Napier and Clube21.
The major problem with this hypothesis, as argued by Sepkoski and Raup, was the
periodicity in extinction it predicted of about twice the observed length, and
thus it could be rejected as a plausible ultimate cause of periodic extinction19.
(4)
An undetected tenth planet, often called “planet X”, was put forward to
explain the periodicity of extinction in 1985 by Whitmire and Matese27.
They argued that such a planet might be able to produce periodic comet showers
if its orbit were highly inclined and if the inner edge of the comet cloud
extended as a thin disk almost to the orbit of Neptune. Under such conditions,
the orbit of Planet X, would precess through the planetary plane. If the
precession period were approximately 56 million years, the planet, with an
estimated one to five Earth masses, would sweep comets out of the inner disk as
it passed through the plane every half period. This hypothesis was largely ad
hoc, said by Sepkoski and Raup19, since the precession period was
determined entirely from periodicities in extinction and cratering that were
being explained. Also, whether the planet would have sufficient mass to scatter
a large number of comets was questioned by Kerr28.
This
hypothesis was further questioned by Collander-Brown and Williams46.
They found that the tenth planet had to be placed on an orbit at such a large
heliocentric distance that no evidence for the existence, or non-existence,
could be found.
(5)
Hallam argued, in the same article as above13, that the most obvious
relationship of mass extinctions was with sea-level changes, which had exhibited
a cyclicity on more than one time scale throughout the Phanerozoic. Mass
extinctions of marine groups took place as a result of the drastic reduction in
area and quality of habitat when extensive epicontinental seas regressed because
of a fall in the sea level. But it was noteworthy that the Cenomanian,
Pliensbachian and Devonian events were not preceeded by significant falls in sea
level, as Hallam said in the same article, so many problems and uncertainties
remained.
Later
the searching for mechanisms of extinction was turned to extinction itself.
Sneppen et al 32 and Sole et al33
used the nonlinear methods to study the process evolution and extinction.
Sneppen concluded that evolution was a self-organized critical phenomenon and
thus large bursts of species and mass extinctions might be a simple consequence
of ecosystems dynamics and required no external cause. Sole’s conclusion was
that extinction process was self-similar fluctuation and a nonlinear response of
the biosphere to perturbations provides the main mechanism for the distribution
of extinction events. But as early as in 1990, Sepkoski disputed internal forces
as the cause of extinction because “internal dynamics was not consistent with
data”20.
Until
now none of the above mechanisms seems to well explain the periodicity of
extinction. Indeed the existence of periodicity in extinction is still in
debate. Further data collection and test methods are necessary to solve the
debate.
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