Evolution and classification

Fossil forms

The lycophytes represent a wide range of extinct and living plants that have contributed important data on evolutionary trends in primitive vascular plants. The earliest lycophytes included Baragwanathia and Protolepidodendron, dating from the early Devonian Period. Both were small herbaceous plants. During the Carboniferous Period, which followed (beginning 358.9 million years ago), the treelike forms of the Lepidodendrales appeared.

Over the years, fossil parts of lepidodendronic plants have been discovered and assigned by taxonomists to so-called form genera, or organ genera: Lepidophyllum for detached leaf fossils, Lepidostrobus for fossil strobili. These form genera are now recognized as portions of one main fossil genus designated Lepidodendron. Some other lycophytes coexisting with the tree lycophytes were small herbaceous plants that resembled modern Lycopodium and Selaginella species.

Annotated classification

Groups marked with a dagger (†) in the listing below are extinct and known only from fossils.

• Class Lycopodiopsida (lycophytes; club mosses and allies)

  Primitive, seedless vascular plants with true roots, stems, and leaves; sporangia associated with leaf bases, the fertile leaves often aggregated to form cones; distributed worldwide but concentrated in the tropics.

  • †Order Protolepidodendrales

    Extinct herbaceous (rarely woody), homosporous lycophytes; about 8 genera, including Baragwanathia and Protolepidodendron.

  • †Order Lepidodendrales

    Extinct tree lycophytes, therefore capable of secondary growth; heterosporous, with some strobili (cones) forming seedlike structures; about 6 genera, including Lepidodendron and Sigillaria.

  • Order Lycopodiales (club mosses)

    Living and extinct plants with primary growth only; homosporous; 16 living genera, mostly tropical: Huperzia (10–15 species), Lycopodium (9–15 species), Lycopodiella (15 species), and Phylloglossum (1 species), the latter of which is restricted to Australia and New Zealand; includes the extinct Lycopodites. The other extant genera are Austrolycopodium, Dendrolycopodium, Diphasiastrum, Diphasium, Lateristachys, Lycopodiastrum, Palhinhaea, Phlegmariurus, Pseudodiphasium, Pseudolycopodiella, Pseudolycopodium, and Spinulum.

  • Order Selaginellales (spike mosses)

    Living and extinct plants with primary growth only; heterosporous; the sole living genus is Selaginella, with more than 700 species, widely distributed around the world; Selaginellites is an extinct genus.

  • Order Isoetales (quillworts)

    Living and extinct plants with secondary growth; heterosporous, with endosporic gametophytes; Isoetites is an extinct genus; for many years the species of Isoetes were difficult to distinguish, but, since the discovery that frequent hybridization was obscuring the differences between species, they are more clearly understood; Isoetes includes about 250 species in swampy, cooler parts of the world.

  • †Order Pleuromeiales

    Extinct unbranched plants, with subterranean, rootlike rhizophores; heterosporous; a single fossil genus, Pleuromeia.

Critical appraisal

This group is treated as a separate class, Lycopodiopsida, in recognition of its distinctive reproductive structures and long fossil history. The number of genera and species in the group has been contentious, but the PPG I consensus has phylogenetic support. As a result, Lycopodium was reorganized into additional genera based on different chromosome numbers, spore sculpturing, and gametophyte morphology.