Aquarium and Aquatic Plants, Part 3 Codexery

Ceratophyllum

Cosmopolitan aquatic genus lacking roots and stomata.

Ceratophyllum

Ceratophyllum, better known as coontails, is a group of flowering plants found worldwide in ponds, marshes, and slow-moving streams, thriving in both tropical and temperate climates. It is the sole living genus in the family Ceratophyllaceae, which itself is the only living family in the order Ceratophyllales. Though these plants are sometimes called "hornworts," that name also refers to Anthocerotophytes, which produce a distinct horn-shaped sporophyte. Since coontails have nothing resembling a horn, using "hornworts" for them is strongly advised against.

These plants grow entirely underwater, often floating at the surface. Their stems can reach 1–3 meters long. At intervals along the stem nodes, they produce rings of bright green leaves that are narrow and frequently forked. Depending on the species, the forked leaves may be brittle and stiff or softer to the touch. Coontails lack roots entirely, even as embryos, though they sometimes develop modified leaves that look like roots to anchor the plant to the bottom. They have no stomata on any part. The flowers are tiny and unremarkable, with both male and female flowers occurring on the same plant. In autumn, the plant forms thick buds called turions that sink to the bottom, making it seem as if frost has killed it; come spring, these buds grow back into long stems, gradually filling the pond.

Carl Linnaeus first described coontails in 1753, with *Ceratophyllum demersum* as the type species. The genus is distinct enough to have its own family, Ceratophyllaceae, though it was once grouped with Nymphaeaceae in the order Nymphaeales under the Cronquist system. Recent research shows it is not closely related to Nymphaeaceae or any other living plant family. Early molecular studies suggested it might be the sister group to all other flowering plants, but newer work indicates it is the sister group to the eudicots. This was supported by sequencing the *C. demersum* genome in 2020. The APG IV system places the family in its own order, Ceratophyllales, and provides a cladogram.

More than 30 species of coontail have been described, but how many are valid remains unresolved. A narrow interpretation rejects over 23 of these as variants, accepting only 7 species. This lumping was so extreme that even subspecies were not recognized. Defined this broadly, the genus includes: *Ceratophyllum australe*, *C.

Quick Facts

Taxon
Ceratophyllum
Type Species
Ceratophyllum demersum L.

Facts from the source article.

Lore & Background

Ceratophyllum grows completely submerged, usually floating on the surface, with stems reaching 1–3 m in length. At intervals along nodes of the stem, it produces rings of bright green leaves that are narrow and often much-branched; the forked leaves are brittle and stiff to the touch in some species, softer in others. Roots are completely absent, even in the embryonic stage, but sometimes modified leaves with a rootlike appearance anchor the plant to the bottom. Stomata are not present on any species. The flowers are small and inconspicuous, with male and female flowers on the same plant. In ponds, it forms thick buds (turions) in autumn, which sink to the bottom and give the impression that the plant has been killed by frost; come spring, these grow back into long stems, slowly filling up the pond.

Coontails were first described by Carl Linnaeus in 1753 with Ceratophyllum demersum as the type species. The genus is considered distinctive enough to warrant its own family, Ceratophyllaceae, but was once considered a relative of Nymphaeaceae and included in Nymphaeales in the Cronquist system. Recent research has shown that the plant is not closely related to Nymphaeaceae or any other extant plant family. Some early molecular phylogenies suggested it was the sister group to all other angiosperms, but more recent research suggests it is the sister group to the eudicots, a notion supported upon sequencing of the C. demersum genome in 2020. The APG IV system placed the family in its own order, the Ceratophyllales.

More than 30 species of coontail have been described, but the specifics of how many are valid is yet to be settled. A narrow interpretation rejected over 23 of these taxa as variants, accepting only 7 species, and even creating subspecies was rejected. A 2018 phylogenetic analysis found support for 4 species being monophyletic, while 3 others are paraphyletic. It found C. australe to be a divergent lineage of C. tanaiticum, rendering C. tanaiticum paraphyletic, but still regards C. australe as a separate species due to significant morphological differences and geographic isolation. C. demersum and C. platyacanthum were recovered as paraphyletic with respect to each other, and could possibly be considered one species, although their morphologies and ecologies are distinct.

Reader's Guide

Ceratophyllum is notable as the only extant genus in its family and order, making it a unique lineage among flowering plants. Its phylogenetic position has been debated: early molecular phylogenies suggested it was the sister group to all other angiosperms, but more recent research, supported by the 2020 sequencing of the C. demersum genome, indicates it is the sister group to the eudicots. The APG IV system places it in its own order, Ceratophyllales. The genus is also distinctive for its complete lack of roots and stomata, and for producing turions that overwinter on pond bottoms. Taxonomically, the number of valid species remains unsettled, with a narrow interpretation accepting only 7 species out of more than 30 described. A 2018 phylogenetic analysis revealed paraphyly among several species, suggesting that some may be conspecific, though morphological and ecological differences have kept them separate. For aquarium and aquatic plant enthusiasts, coontails are a hardy, rootless, submerged plant that can fill ponds with long stems, and their ability to form overwintering buds makes them resilient in temperate climates.

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