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Terrestrial Palaeoecology and Global Change

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84 Valentin A. Krassilov. <strong>Terrestrial</strong> <strong>Palaeoecology</strong><br />

Other plant communities for which light poses a major problem thrive in the highlatitude<br />

areas of winter darkness. It is commonly taken for granted that a high latitude<br />

fossil plant of rainforest affinities must be of a tropical derivation, but it might have been<br />

the other way round: polar life forms, preadapted to shading due to their tolerance of<br />

prolonged darkness, might find their way under the canopies of primordial rainforests.<br />

IV.2.2. Temperate deciduous vegetation<br />

Seasonal deciduousness might have appeared in leafless plants that shed green<br />

branchlets, as extant Ephedra species do. The Devonian archaeopterids were shedding<br />

leaf-like syntelomic systems that sometimes formed mat-type accumulations (Fig.37)<br />

indicating seasonal deciduousness, as in the Late Devonian Tanaitis, a leafless heterosporangial<br />

archaeopterid genus akin to Svalbardia, the northernmost (present-day<br />

latitudes) representative of the group. The branch-mats occur on bedding planes of a<br />

levee-type detrital siltstone containing also abundant fossil wood fragments (Krassilov et<br />

al., 1988).<br />

The modern-type deciduousness appeared with the Permian Angarian <strong>and</strong> Gondwana<br />

floras of the broadleaved vojnovskyan “cordaites” <strong>and</strong> glossopterids respectively,<br />

forming mats of individually shed leaves <strong>and</strong> leafy spur-shoots. In the Late Permian,<br />

a dominant morphotype of ribbon-shaped leaves with a callous abscission scar was<br />

joined by the newly appearing flabellate Ginkgo-like (Rhipidopsis) <strong>and</strong> aciculate coniferoid<br />

morphotypes that became more prominent in the Mesozoic. The mid-Triassic<br />

to Early Cretaceous taphofloras of northern Asia were dominated by Czekanowskia,<br />

Phoenicopsis, Pseudotorellia, Ginkgoites, Pityophyllum <strong>and</strong> Nilssonia forming<br />

extensive leaf-mats. All these plants were heteroblastic, shedding spur-shoots with<br />

leaves (Fig. 38).<br />

Shoot dropping was inherited by deciduous conifers, such as Parataxodium, dominant<br />

over northern Asia in the Late Cretaceous. They formed mixed leaf mats with<br />

broadleaved platanoids <strong>and</strong> hamamelids (Trochodendroides) that grew as understorey<br />

<strong>and</strong> seral communities of the Platano-Parataxodietum (Krassilov, 1979). With a decline<br />

of the latter over the Cretaceous/Tertiary boundary, new broadleaved deciduous<br />

forest types were formed on the basis of the seral Platano-Trochodendroidetum, incorporating<br />

the deciduous forms taxodiaceous conifers derived from Parataxodium<br />

(Metasequoia, Taxodium, Glyptostrobus). They constituted the core of the Arcto-<br />

Tertiary biome mixed forest types, with biserrate betuloid components first recorded as<br />

dominant elements from intermontane depressions of circum-Pacific coastal ranges<br />

(Krassilov, 1989c), supposedly representing an upl<strong>and</strong> differentiation of the early hamamelid<br />

complex.<br />

Summer-dry deciduousness is poorly recorded because leaves are shed in a season<br />

unfavorable for preservation scarcely forming leaf mats. It might have first appeared in

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