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Terrestrial Palaeoecology and Global Change

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Chapter 9. Crises<br />

367<br />

lobes, making its second entry in the latest Permian (Krassilov, 1999b; Fig. 128). A<br />

Cretaceous example is Czekanowskia, a widespread pioneer of Mesozoic peatl<strong>and</strong>s,<br />

exceedingly rare over the Late Cretaceous in general, but prominent in the Maastrichtian<br />

dinosaur beds of Amur Basin (Bugdaeva, 2001). Encephalartites, a semiextinct<br />

Mesozoic cycadophyte, is likewise common in the contemporaneous deposits of Koryak<br />

Highl<strong>and</strong>s (Krassilov et al., 1988) while the relatively poor KTB assemblage of Augustovka,<br />

western Sakhalin, contains two such cycadophytic genera, Pterophyllum <strong>and</strong><br />

Cycadites, giving it an archaic mid-Cretaceous aspect (Krassilov 1979; Fig. 146).<br />

An association of macromutational newcomers with Lazarus forms is a distinctive<br />

feature of crisis communities that combine a low diversity with a high morphological<br />

disparity <strong>and</strong> morphological polymorphism of dominant species (IX.9.3).<br />

IX.8. Recovery<br />

In plant communities, recoveries after local crises tend to restore the pre-existing<br />

structure. The models of recovery based on recent examples (Carr-Kitchell, 1980; Barry<br />

et al., 1991; Rosenzweig & McCord, 1991; Frederiksen, 1994) emphasize the significance<br />

of an initial diversity <strong>and</strong> of speciation rates that are fostered by vacant ecological<br />

niches as well as by immigration.<br />

The climax cut-off model (Krassilov, 1992a; IX.6) maintains that, in a stressful environment,<br />

ecological succession is halted at an early stage never reaching the potential<br />

climax. In effect, the recovery of climax species is hampered <strong>and</strong>, in the long sustained<br />

disclimax situations, might never occur. Major ecological crises truncate ecological successions<br />

over a wide range of terrestrial <strong>and</strong> marine habitats, thus driving the climax species<br />

to extinction. This explains the great extinctions that are recorded as nearly simultaneous<br />

disappearances of dominant forms over a wide range of habitats.<br />

The post-crisis recoveries eventually bring forth new dominants that are recruited<br />

from among pioneer species of pre-existing, or recently arising, taxa. Thus, dinosaurs<br />

recovered, with new dominant forms, after their near-extinction over the mid-Cretaceous<br />

crisis but were replaced by mammals over the KTB crisis (IX.5).<br />

The recovery or non-recovery of a formerly dominant group basically depends on its<br />

ecological status, with severity of disturbance <strong>and</strong> competition as additional factors. Recovery<br />

usually involves pioneer species or, in a broader sense, taxonomic groups having<br />

their representatives in the pioneer/early seral stages. It is problematic for the taxa represented<br />

by climax forms only. In biotic communities, the pioneer <strong>and</strong> seral species may<br />

belong to the same higher taxon (evolutionary grade) as the climax species. For instance,<br />

in broadleaved forests, the climax (lime, oak) <strong>and</strong> seral (poplar, maple) dominants are of<br />

the same evolutionary grade. The recovery of the lime–oak climax is impeded in heavily<br />

polluted environments but the broadleaved forest may recover through its persistent<br />

seral stages.

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