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An incredible variety of seedless plants populates the terrestrial landscape. Mosses grow on tree trunks, and horsetails ( [link] ) display their jointed stems and spindly leaves on the forest floor. Yet, seedless plants represent only a small fraction of the plants in our environment. Three hundred million years ago, seedless plants dominated the landscape and grew in the enormous swampy forests of the Carboniferous period. Their decomposing bodies created large deposits of coal that we mine today.
Bryophytes, an informal grouping of the nonvascular plants, are the closest extant relative of early terrestrial plants. The first bryophytes most probably appeared in the Ordovician period, about 490 million years ago. Because of the lack of lignin—the tough polymer in cell walls in the stems of vascular plants—and other resistant structures, the likelihood of bryophytes forming fossils is rather small, though some spores made up of sporopollenin have been discovered that have been attributed to early bryophytes. By the Silurian period (440 million years ago), however, vascular plants had spread throughout the continents. This fact is used as evidence that nonvascular plants must have preceded the Silurian period.
There are about 18,000 species of bryophytes, which thrive mostly in damp habitats, although some grow in deserts. They constitute the major flora of inhospitable environments like the tundra, where their small size and tolerance to desiccation offer distinct advantages. They do not have the specialized cells that conduct fluids found in the vascular plants, and generally lack lignin. In bryophytes, water and nutrients circulate inside specialized conducting cells. Although the name nontracheophyte is more accurate, bryophytes are commonly referred to as nonvascular plants.
In a bryophyte, all the conspicuous vegetative organs belong to the haploid organism, or gametophyte. The diploid sporophyte is barely noticeable. The gametes formed by bryophytes swim using flagella. The sporangium, the multicellular sexual reproductive structure, is present in bryophytes. The embryo also remains attached to the parent plant, which nourishes it. This is a characteristic of land plants.
The bryophytes are divided into three divisions (in plants, the taxonomic level “division” is used instead of phylum): the liverworts, or Marchantiophyta; the hornworts, or Anthocerotophyta; and the mosses, or true Bryophyta.
Liverworts (Marchantiophyta) may be viewed as the plants most closely related to the ancestor that moved to land. Liverworts have colonized many habitats on Earth and diversified to more than 6,000 existing species ( [link] a ). Some gametophytes form lobate green structures, as seen in [link] b . The shape is similar to the lobes of the liver and, hence, provides the origin of the common name given to the division.
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