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Terrestrial Palaeoecology and Global Change

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Chapter 9. Crises<br />

361<br />

of biological diversity. Mass extinction is a rough regulation of biological diversity via<br />

population strategies adjusted to long-lasting environmental disturbances.<br />

IX.5. Replacement of ecological dominants<br />

The idea of biotic turnovers over the critical boundaries has originated from the replacements<br />

of charismatic groups, in particular, of dinosaurs by mammals. This latter<br />

replacement signifies the end of the Mesozoic <strong>and</strong> the beginning of the new era. Opponents<br />

of the idea have argued that, since dinosaurs constituted but a minor part of biological<br />

diversity, their elimination (typically considered in this line of thought as the result of<br />

a competitive replacement by mammals) does not signify a global biotic turnover of<br />

whatever, if any, scientific meaning. If we instead focus on the replacement of gymnosperms<br />

by angiosperms, then the boundary of geological eras (the Mesophytic/Cenophytic)<br />

must be placed in the middle of the Cretaceous. A choice of one or another<br />

option for the boundary is therefore a matter of convention.<br />

Such considerations have led many evolutionists to believe that stratigraphic levels of<br />

accelerated turnover rates can be objectively defined for particular lineages rather than<br />

for biota as a whole. The eras of life need not be taken literally as distinct stages of<br />

organic evolution. Some lineages do show high turnover rates over the selected boundaries,<br />

others do not. Plants have evolved under a different beat than animals while the<br />

dinosaur extinctions on l<strong>and</strong> at approximately the same time level as the ammonite extinctions<br />

in the sea is a mere coincidence.<br />

However, with the rise of biospheric ecology, this logic no longer seems faultless.<br />

Ecological niches of the larger dinosaur herbivores seem to have remained vacant for at<br />

least 5 m.y. after their extinction, hence competitive elimination is unlikely. Some 150<br />

m.y. long coexistence of dinosaurs <strong>and</strong> mammals only means that the dinosaurian communities<br />

<strong>and</strong> their contemporaneous mammalian communities were fairly well-equilibrated,<br />

with minimal niche overlaps, while the replacement means that this long-st<strong>and</strong>ing<br />

ecological equilibrium was eventually disrupted. Adaptive radiation of modern mammals<br />

postdates, rather than predates, the decline of dinosaurs. Archaic mammals perished<br />

simultaneously with dinosaurs, although over a somewhat longer time interval. The buildup<br />

of mammalian diversity is due to new types of vegetation, such as grassl<strong>and</strong>, as well<br />

as new trophic niches, such as frugivory.<br />

The gymnosperm/angiosperm story is fairly consistent with the above version of<br />

the dinosaur/mammal story (Krassilov, 1997a <strong>and</strong> elsewhere). Both only make sense<br />

in the context of biospheric evolution. Angiosperms first appeared in the mid-Neocomian,<br />

concomitantly with the therian mammals (Aegialodon dawsoni <strong>and</strong> subsequent<br />

records of the metatherian–eutherian grade: Lillegraven, 1969; Lillegraven et al., 1979;<br />

Fox, 1980) <strong>and</strong> the true birds, jointly signifying a major innovation of terrestrial plant<br />

communities.

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