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Evolution__3rd_Edition

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538 PART 5 / Macroevolution<br />

Plants evolved adaptations for land<br />

life...<br />

Origin of<br />

land plants<br />

Origin of vascular plants<br />

(1) vascularization<br />

(2) roots<br />

(3) leaves<br />

Origin of seeds<br />

change, or internal biological innovations, or some mix of the two. But there is no<br />

consensus yet.<br />

18.5 <strong>Evolution</strong> of land plants<br />

The land was first colonized by microbes. Prokaryotic fossil cells have been found from<br />

terrestrial environments over 1,000 million years ago. Terrestrial life existed only in<br />

microbial form until the land was colonized by plants and fungi. Little is known about<br />

early terrestrial fungi, but for plants we have widely accepted evidence of fossil spores<br />

from 475 million years ago, and further evidence of fossil spores from maybe 550 million<br />

years ago. Land plants are most closely related to a group of green algae called<br />

charophyceans (Figure 18.7). Early fossil spores have a “tetrad” structure that is found<br />

in some modern bryophytes, and most early plant fossils seem to be related to the<br />

phylogenetic branches leading to modern bryophytes and pteridophytes.<br />

The main events in the early evolution of land plants were the evolution of a resistant<br />

spore stage, then the evolution of vascular tissue, followed by roots and then leaves.<br />

Bryophyta such as moss have spores but lack vascular tissue. Pteridophyta such as ferns<br />

have vascular tissue. Vascular tissue, and particularly vascular tissue that is built of<br />

tracheid cells with lignin-containing cell walls, enables a plant to support itself on land.<br />

The earliest fossils of whole plants, as distinct from spores, date back to around<br />

430 million years ago. Clarksonia is one of the commonest fossils of this time. These<br />

early land plants lacked roots and leaves, and simply had branching stems. We can infer<br />

that they photosynthesized through their stems, because stomata are visible in the fossil<br />

stems. Fossils with leaves appear 390–350 million years ago.<br />

The evolution of leaves coincides with a dramatic fall of about 90% in atmospheric<br />

carbon dioxide concentration. One hypothesis connects these events. The initial evolution<br />

of land plants may have removed carbon dioxide from the atmosphere, not only by<br />

their relatively minor photosynthetic activity through the stem but, more importantly,<br />

Charophyceans<br />

Bryophytes (e.g., moss)<br />

Pteridophytes (e.g., ferns)<br />

Gymnosperms (conifers)<br />

Angiosperms (flowering plants)<br />

Figure 18.7<br />

Phylogeny of the main groups of plants. Land plants are most<br />

closely related to a group of algae called charophyceans. The main<br />

events in plant evolution are the evolution of land adaptations,<br />

then seeds, and then flowers.<br />

..

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