Leaves

Leaves today are, in almost all instances, an adaptation to increase the amount of sunlight that can be captured for photosynthesis. Leaves certainly evolved more than once, and probably originated as spiny outgrowths to protect early plants from herbivory. Leaves are the primary photosynthetic organs of a plant. Based on their structure, they are classified into two types: microphylls, which lack complex venation patterns, and megaphylls, which are large and have complex venation. It has been proposed that these structures arose independently.[36] Megaphylls, according to the Telome hypothesis, have evolved from plants that showed a three dimensional branching architecture, through three transformations—plantation, which involved formation of a planar architecture, webbing, or formation of the outgrowths between the planar branches and fusion, where these webbed outgrowths fused to form a proper leaf lamina. All three steps happened multiple times in the evolution of today's leaves.[37] It has been proposed that before the evolution of leaves, plants had the photosynthetic apparatus on the stems. Today's megaphyll leaves probably became commonplace some 360mya, about 40my after the simple leafless plants had colonized the land in the early Devonian period. This spread has been linked to the fall in the atmospheric carbon dioxide concentrations in the Late Paleozoic era associated with a rise in density of stomata on leaf surface. This must have allowed for better transpiration rates and gas exchange. Large leaves with less stomata would have gotten heated up in the sun's heat, but an increased stomata

density allowed for a better-cooled leaf, thus making its spread feasible.[38][39] The rhyniophytes of the Rhynie chert comprised nothing more than slender, unornamented axes. The early to middle Devonian trimerophytes, therefore, are the first evidence of anything that could be considered leafy. This group of vascular plants are recognisable by their masses of terminal sporangia, which adorn the ends of axes which may bifurcate or trifurcate.[32] Some organisms, such as Psilophyton, bore ‹See Tfd›enations. These are small, spiny outgrowths of the stem, lacking their own vascular supply. Around the same time, the zosterophyllophytes were becoming important. This group is recognisable by their kidney-shaped sporangia, which grew on short lateral branches close to the main axes. They sometimes branched in a distinctive H-shape.[32] The majority of this group bore pronounced spines on their axes. However, none of these had a vascular trace, and the first evidence of vascularised enations occurs in the Rhynie genus Asteroxylon. The spines of Asteroxylon had a primitive vasuclar supply – at the very least, leaf traces could be seen departing from the central protostele towards each individual "leaf". A fossil known as Baragwanathia appears in the fossil record slightly earlier, in the late Silurian.[40] In this organism, these leaf traces continue into the leaf to form their mid-vein.[41] One theory, the "enation theory", holds that the leaves developed by outgrowths of the protostele connecting with existing enations, but it is also possible that microphylls evolved by a branching axis forming "webbing".