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Terrestrial Palaeoecology and Global Change

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

Gigantopterids <strong>and</strong> emplectopterids, both prominent in the Cathaysian-type assemblages<br />

of eastern Asia <strong>and</strong> western North America, typically have composite leaves in<br />

which the entire or simply pinnate leaf blades were formed by marginal fusion of ultimate<br />

<strong>and</strong> penultimate segments, with the original segmentation betrayed by the conservative<br />

venation pattern (Asama, 1976; Krassilov, 1995c, VIII.6). The hierarchical areolate<br />

venation of such leaves owes its origin to meristematic fusion zones acting as a<br />

plate meristem (Krassilov, 1995c). This leaf morphology is evidence of a prolonged meristematic<br />

activity characteristic of megaphyllous life forms in equable climates. An advanced<br />

Gigantopteris morphotype, some variants of which show clearly defined driptips<br />

(Yao, 1986), occurred in southern Cathaysia. A survival of Carboniferous forms,<br />

such as a tree fern Psaronius <strong>and</strong> arboreal lepidophyte Lepidodendron, also indicates<br />

a tropical climate.<br />

Noto-Permian vegetation. Classical Gondwana flora came from the Narmada–<br />

Son–Mahanadi graben system that transects peninsular India. Boulder beds occur at the<br />

base of the basinal sedimentary fill followed by coal measures that contain the bulk of<br />

the flora. The Himalayan occurrences are scattered from Kashmir to Assam, with a<br />

few outliers in Tibet, Yunnan <strong>and</strong> the “mixed” floras of Indonesia <strong>and</strong> New Guinea. A<br />

number of well-defined species are shared by Indian <strong>and</strong> Australian floras, but the other<br />

parts of “Gondwanal<strong>and</strong>” are floristically less similar (Archangelsky, 1990, 1996; Maheshwari,<br />

1992).<br />

A primitive glossopterid leaf morphotype Gangamopteris appeared already in the<br />

pre-tillite deposits. Early Gangamopteris plants are commonly envisaged as growing in<br />

sparse vegetation patches of a glacial l<strong>and</strong>scape. However, fossil plant assemblages<br />

found immediately below <strong>and</strong> above the Dwykka Tillite in South Africa, the Bacchus<br />

Marsh Tillite in Australia <strong>and</strong> the Talchir boulder beds in Peninsular India suggest a less<br />

dismal picture, with a number of Carboniferous survivors (Ch<strong>and</strong>ra et al., 1992; Pant,<br />

1996). Leaf morphology gives no support to either a “Gangamopteris tundra” or a<br />

“Gangamopteris taiga”. Morphologically, Gangamopteris is linked, via Palaeovittaria<br />

<strong>and</strong> Noegerrathiopsis, to the Zamiopteris–Cordaites group of the Arcto-Permian temperate<br />

zone.<br />

The emblematic Glossopteris morphotype came forth <strong>and</strong> diversified in the period of<br />

a widespread coal accumulation. Some variants of its highly polymorphiic leaves show<br />

distinct abscission scars, others had drip-tips characteristic of rainforest foliage. Such a<br />

morphological diversity obviously suggests a wide range of habitats (Pant, 1996), which<br />

is confirmed by taphonomic evidence (Retallack, 1980). Glossopterids are sometimes<br />

preserved as leaf mats containing spur-shoots shed with clusters of intact leaves, as is<br />

typical of deciduous riparian trees. A constant association of Glossopteris with Vertebraria,<br />

an aerenchymatous underground organ, suggests a peatl<strong>and</strong> habitat. The coriaceous<br />

lingulate leaves, as in the modern mangroves, <strong>and</strong> the uniquely preserved seedlings<br />

(Pant & Nautiyal, 1987) are also consistent with a helophyte life form in a majority<br />

of Indo-Australian glossopterids. The greatest morphological diversity of Glossopteris

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