Colonization of land

Land plants evolved from chlorophyte algae, perhaps as early as 510 million years ago;[4] some molecular estimates place their origin even earlier, as much as 630 million years ago.[5] Their closest living relatives are the charophytes, specifically Charales; assuming that the Charales' habit has changed little since the divergence of lineages, this means that the land plants evolved from a branched, filamentous alga dwelling in shallow fresh water,[6] perhaps at the edge of seasonally desiccating pools.[4] The alga would have had a haplontic life cycle: it would only very briefly have had paired chromosomes (the diploid condition) when the egg and sperm first fused to form a zygote; this would have immediately divided by meiosis to produce cells with half the number of unpaired chromosomes (the haploid condition). Co-operative interactions with fungi may have helped early plants adapt to the stresses of the terrestrial realm.[7] Plants were not the first photosynthesisers on land; weathering rates suggest that organisms were already living on the land 1,200 million years ago,[4] and microbial fossils have been found in freshwater lake deposits from 1,000 million years ago,[8] but the carbon isotope record suggests that they were too scarce to impact the atmospheric composition until around 850 million years ago.[9] These organisms, although phylogenetically diverse,[10] were probably small and simple, forming little more than an "algal scum".[4] The first evidence of plants on land comes from spores of Mid-Ordovician age (early Llanvirn, ~470 million years ago).[11][12][13] These spores, known as cryptospores, were produced either singly (monads), in pairs (diads) or groups of four (tetrads), and their microstructure resembles that of modern liverwort spores, suggesting they share an equivalent grade of organisation.[14] They are composed of sporopollenin – further evidence of an embryophytic affinity.[15] It could be that atmospheric 'poisoning' prevented eukaryotes from colonising the land prior to this,[16] or it could simply have taken a great time for th

necessary complexity to evolve.[17] Trilete spores similar to those of vascular plants appear soon afterwards, in Upper Ordovician rocks.[18] Depending exactly when the tetrad splits, each of the four spores may bear a "trilete mark", a Y-shape, reflecting the points at which each cell squashed up against its neighbours.[11] However, this requires that the spore walls be sturdy and resistant at an early stage. This resistance is closely associated with having a desiccation-resistant outer wall—a trait only of use when spores must survive out of water. Indeed, even those embryophytes that have returned to the water lack a resistant wall, thus don't bear trilete marks.[11] A close examination of algal spores shows that none have trilete spores, either because their walls are not resistant enough, or in those rare cases where it is, the spores disperse before they are squashed enough to develop the mark, or don't fit into a tetrahedral tetrad.[11] The earliest megafossils of land plants were thalloid organisms, which dwelt in fluvial wetlands and are found to have covered most of an early Silurian flood plain. They could only survive when the land was waterlogged.[19] There were also microbial mats.[20] Once plants had reached the land, there were two approaches to dealing with desiccation. The bryophytes avoid it or give in to it, restricting their ranges to moist settings, or drying out and putting their metabolism "on hold" until more water arrives. Tracheophytes resist desiccation: They all bear a waterproof outer cuticle layer wherever they are exposed to air (as do some bryophytes), to reduce water loss, but—since a total covering would cut them off from CO2 in the atmosphere—they rapidly evolved stomata, small openings to allow gas exchange. Tracheophytes also developed vascular tissue to aid in the movement of water within the organisms (see below), and moved away from a gametophyte dominated life cycle (see below). Vascular tissue also facilitated upright growth without the support of water and paved the way for the evolution of larger plants on land.