''Ceratopteris'' is the only genus among

homosporous In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...

ferns that is exclusively aquatic. It is pan-tropical and classified in the

Parkerioideae Parkerioideae, Synonym (taxonomy), synonym Ceratopteridoideae, is one of the five subfamilies in the fern family Pteridaceae. It includes only the two genera ''Acrostichum'' and ''Ceratopteris''. The following diagram shows a likely phylogenic re ...

subfamily of the family Pteridaceae.

Description

Erect aquatic or subaquatic ferns of moderate size. Rhizome short, fleshy, horizontal and ascending to erect, loosely rooted in the mud or +/- floating, radial, dictyostelic with numerous meristeles and medullary strands, young parts bearing thin, ovate, +/- cordate, clathrate scales. Fronds stipitate, the stipes fleshy, with numerous longitudinal air canals, abaxially rounded and ribbed, adaxially flattened, vascular bundles in a peripheral ring, one with each rib and several to the adaxial side, and several smaller medullary strands; lamina dimorphic, sterile fronds +/- spreading, 2–3-pinnatifid with broad membranous lobes, venation reticulate without included free veinlets, often with proliferous buds in the axils; fertile fronds erect, longer, narrower and more divided than the sterile, the lobes strongly recurved to completely cover the adiaxial surface, venation longitudinal, branching at the bases of the lobes. Sporangia solititary, scattered along the veins, exindusiate but protected by the continuous reflexed margin of the lamina, large, short-stalked, annulus broad, irregular, of 30–70 thickened cells, or lacking, containing 16 to 32 spores. Spores large, trilete, ribbed with irregular long meshes.

Taxonomy

''Ceratopteris'' was long placed in the monogeneric family Parkeriaceae, thought to be unique because of its aquatic adaptations. However, recent genetic analysis has shown it to be clearly allied with ''

Acrostichum ''Acrostichum'' is a fern genus in the Parkerioideae subfamily of the Pteridaceae. It was one of the original pteridophyte genera delineated by Linnaeus. It was originally drawn very broadly, including all ferns that had sori apparently "ac ...

'' in the subfamily Parkerioideae, within the family Pteridaceae.

Species

At one time, some authorities recognized only one species; now some authorities recognize only four species. However, recent work by Masuyama and Watano '' has suggested that C. thalictroides'' actually consists of four cryptic species (''thalictroides'', ''froesii'', ''gaudichaudii'', ''oblongibloba''). *''

Ceratopteris cornuta ''Ceratopteris'' is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. Description Erect aquatic or subaquatic ferns of moderate size. ...

'' (P. Beauv.) Le Prieur; Ann. Sci. Nat. 19: 103. t. 4A (1830) *''

Ceratopteris froesii ''Ceratopteris'' is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. Description Erect aquatic or subaquatic ferns of moderate size. ...

'' Brade, Arch. Jard. Bot. Rio de Janeiro 18: 31 (1964). *''

Ceratopteris gaudichaudii ''Ceratopteris'' is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. Description Erect aquatic or subaquatic ferns of moderate size. ...

'' Brongn.; Bull. Soc. Philom. 187 (1821) *''

Ceratopteris oblongiloba ''Ceratopteris'' is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. Description Erect aquatic or subaquatic ferns of moderate size. ...

'' Masuyama & Watano—Acta Phytotax. Geobot. 61(2): 84 (-85; figs. 3-4). 2010 ov 2010 *''

Ceratopteris pteridoides ''Ceratopteris'' is the only genus among homosporous ferns that is exclusively aquatic. It is pan-tropical and classified in the Parkerioideae subfamily of the family Pteridaceae. Description Erect aquatic or subaquatic ferns of moderate size. ...

'' (Hook.) Hieron.; Bot. Jahrb. Syst. 34: 561 (1905) *''

Ceratopteris richardii ''Ceratopteris richardii'' is a fern species belonging to the genus ''Ceratopteris'', one of only two genera of the subfamily Parkerioideae of the family Pteridaceae. It is one of several genera of ferns adapted to an aquatic existence. ''C. ri ...

'' Brongn.; Dict. Class. Hist. Nat. 3: 351 (1823) - "C-fern" *''

Ceratopteris thalictroides ''Ceratopteris thalictroides'' is a fern species belonging to the genus '' Ceratopteris'', one of only two genera of the subfamily Parkerioideae of the family Pteridaceae. Common names ''Ceratopteris thalictroides'' is commonly known as wate ...

'' (L.) Brongn.; Bull. Sci. Soc. Philom. Paris, sér. 3, 8: 186 (1821)

Distribution and habitat

This widespread genus of four to six species inhabits the humid tropics. ''Ceratopteris'' is also important in the study of pteridophytes and is a commonly used

model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...

for use in

genomic Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...

studies, due to the ease and rapidity with which it can be grown in laboratories, as well as having well-characterized phenotypes. Patented strains of this plant have been developed.

Outcrossing and self-fertilization

A gametophyte is one of the two alternating multicellular phases of the fern life cycle. The gametophyte is

haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...

. The other stage is the diploid

sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote pr ...

. A gametophyte is a multicellular organism that develops from a haploid spore (that has one set of chromosomes). ''Ceratopteris thalictroides'' has two types of gametophytes with different sexual expression: these types are

hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...

and male. Hermaphroditic gametophytes have one or several

archegonia An archegonium (pl: archegonia), from the ancient Greek ''ἀρχή'' ("beginning") and ''γόνος'' ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female ga ...

and a few antheridia. Archegonia are multicellular structures of the gametophyte that produce and contain the ovum or female gamete. The corresponding male organ is the antheridium, a haploid organ producing and containing

sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...

. The second of the two types of gametophyte are male that produce only antheridia. When ''Ceratopteris richardii'' ferns are grown alone they develop into hermaphrodites (with mainly female archegonia). However, plants that grow near established genetically identical individuals develop into males. The hermaphrodite secretes a pheromone (

antheridiogen Antheridiogens are a class of chemicals secreted by fern gametophytes that have "been shown to influence production of male gametangia and thus mating systems in a large number of terrestrial fern species". Antheridiogens are only observed in homos ...

) that tells others nearby to be male. Thus when plants are near each other outcrossing is promoted. Self-fertilization predominates when there is no nearby partner.

Toxicity

The plants may have carcinogenic properties.

Uses

''Ceratopteris'' is a fairly popular aquarium plant, often sold under the name "water sprite." It may be grown as an emersed but natant (floating) plant, or as an immersed plant rooted in the substrate. Under the right conditions the plants will grow fully emerse erect leaves. Under a bright light the plants can grow fast, and be used to help cycle an aquarium. This plant is often used as a vegetable, particularly in Asia.