Max-Planck-Institut für Physik komplexer
Systeme
International Workshop on
Biological Evolution and Statistical
Physics
May 10-14, 2000
Restructuring the Hystory of Life: The Role of
Mass Extinction, Recoveries and Radiation
Douglas
H. Erwin
Department
of Paleobiology,
National Museum of Natural
History
Washington, DC 20560
erwin.doug@nmnh.si.edu
Patterns in the history of life over tens to hundreds of millions
of years reflect a combination of long-term adaptive trends, filtered by
the perturbations imposed by mass extinctions. How are ecosystems restructured
following extinctions? Biological intuition, reflected in many models
and simulations, suggests that diversification occurs through re-occupation
and diversification into niches. A variety of lines of evidence suggest
this
view is too simplistic. Rather, niches are created during recoveries
(and some other biotic radiations), driving further diversification.
How these niches are created has important implications for the long-term
effect of mass extinctions and subsequent recoveries.
Some mass extinctions appear
to have had little immediate ecological effect, or long-term evolutionary
effect (end-Ordovician, end-Triassic). Other mass extinctions have
dramatically shifted the course of evolution (end-Permian, for marine invertebrates,
some tetrapods, and perhaps insects; end-Cretaceous, for tetrapods).
Detailed studies of post-extinction recoveries have revealed that many
of these changes reflect the pattern of
the recovery at least as much, and perhaps more than the pattern of
extinction. Thus, mass extinctions may remove entire clades and even
ecological guilds, depending on the pattern of selectivity. The pattern
of rebuilding ecosystems during post-extinction recoveries appears to be
driven by a different dynamic, however. The immediate post-extinction
survival interval is often dominated by opportunists, but these clades
do not necessarily dominate ecosystems when diversification increases during
the recovery interval. During this interval ecological relationships
may be paramount in determining the long-term success of clades. The major
features of gastropod evolution during the Paleozoic were driven by the
need to accommodate, water currents through the gill. During the
course of the Paleozoic, gastropods appear to have repeatedly discovered
the same six solutions to this adaptive problem. The appearance of
shell-crushing predators in the aftermath of the end-Permian mass extinction
changed this system. No longer was adaptation to water currents the
primary adaptive problem; this had been replaced by resistance to
predation. Consequently, as these new ecosystems developed in the
Middle Triassic, several groups
which suffered relatively little during the mass extinction disappeared
during the recovery because the trajectory taken during the recovery transformed
the ecosystem. Detailed studies make it clear that this was a product
of the pattern of recovery, not simply a result of the extinction. Recent
work by Kircher and Weil reveals an approximately 10 myr lag
following biotic crises at a broad range of scales, suggesting the
creation of ecological roles is an ongoing component of recoveries. Models
of recovery and radiations need to include a positive feedback component
which dynamically changes the dimensions of the ecospace.
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