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— CH. 1 · INTRODUCTION —

Pteridophyte

7 min listen · Ch. 1 of 8
8 sections
  • A pteridophyte produces neither flower nor seed. Its way of reproducing is hidden, which is why these plants earned the old label cryptogams, a word that means exactly that. Instead of seeds, they scatter spores. They carry xylem and phloem, the plumbing of a true vascular plant, and they are the ancestors of the green world we know today. Ferns belong here. So do horsetails, often counted among the ferns, and the lycophytes, which include clubmosses, spikemosses, and quillworts. Yet here is the puzzle this documentary will pursue. These plants were grouped together for centuries, but they do not actually form one natural family. Ferns turn out to be closer kin to seed plants than to lycophytes. So what holds the term together, why has it survived, and how do scientists draw the lines between these spore-bearing survivors?

  • Pteridophyta is no longer a widely accepted taxon, yet the word pteridophyte lives on in everyday speech. The reason is that the plants it once described do not share a single common ancestor to the exclusion of others. Ferns and horsetails are more closely related to seed plants than they are to lycophytes. That makes the old grouping a paraphyletic grade rather than a clade. Modern studies of land plants agree that seed plants emerged from pteridophytes, closer to the ferns than to the lycophytes. The convenience of the term keeps it alive in working vocabulary. The science of these plants is still called pteridology, and its practitioner a pteridologist. Two bodies carry the legacy forward in their names: the International Association of Pteridologists and the Pteridophyte Phylogeny Group, the latter the source of the 2016 classification known as PPG I.

  • Alternation of generations governs the life of every pteridophyte, the same rhythm found in bryophytes and in seed plants. A diploid generation called the sporophyte produces spores. It is followed by a haploid generation, the gametophyte or prothallus, which produces gametes. What sets pteridophytes apart is independence. Unlike in seed plants, both generations live freely on their own, neither tethered to the other. Compared with bryophytes, the sporophyte here is branched and far larger and more conspicuous. The sexuality of these gametophytes splits into named patterns. A dioicous gametophyte is either male, making antheridia and sperm, or female, making archegonia and egg cells. A monoicous one makes both and can act as either. Timing matters too. Protandrous gametophytes mature their antheridia first, male before female; protogynous ones reverse the order. These terms should not be confused with monoecious and dioecious, which describe a seed plant's sporophyte and whether it bears both pollen and seeds or only one.

  • Roots, stem, and leaves divide the body of a pteridophyte sporophyte cleanly. The root system is always adventitious, meaning it arises from places other than a primary root. The stem may run underground or rise into the air. Leaves come in two forms, the small microphylls or the larger megaphylls. These plants carry the apomorphies of vascular plants, the vascular tissue that defines the group, alongside older land plant traits like spore dispersal and the absence of seeds. In Smith's molecular study, the ferns earned a distinctive profile: lateral root origin in the endodermis, usually mesarch protoxylem in shoots, a pseudoendospore, a plasmodial tapetum, and sperm cells bearing 30-1000 flagella. That last figure marks how far these swimmers travel from the single tail of a seed plant's pollen.

  • Ferns account for nearly 90% of the living diversity among pteridophytes. Smith and colleagues, in 2006, published the first higher-level classification of the molecular phylogenetic era. They treated ferns as monilophytes within a scheme that placed lycophytes at less than 1% of extant vascular plants and seed plants at roughly 260,000 species. The monilophytes themselves number about 9,000 species and gather horsetails, whisk ferns, and all the eusporangiate and leptosporangiate ferns. The word moniliform means bead-shaped. Kenrick and Crane introduced it in 1997 as a scientific replacement for fern, including the horsetails, and Pryer and colleagues established it by 2004. Not everyone approved. Christenhusz and Chase, reviewing classification schemes in 2014, called the usage irrational and discouraged it, noting that the name Filicopsida already existed. By contrast, lycopod or lycophyte means wolf-plant, and the older catch-all fern ally was simply a convenient label for non-fern spore-bearers, never a natural grouping.

  • Christenhusz and Chase, in 2014, split the old assemblage into two separate, unrelated taxa. They counted the lycopods as one subclass, three orders each with a single family, five genera, and about 1,300 species. The ferns they sized at four subclasses, 11 orders, 21 families, about 212 genera, and roughly 10,535 species. The four fern subclasses ran Equisetidae, Ophioglossidae, Marattiidae, and Polypodiidae, matching Smith's four classes, with Ophioglossidae corresponding to Psilotopsida. Formal ranking proved awkward. Both Infradivision and Moniliformopses are invalid names under the International Code of Botanical Nomenclature. Ferns, though a monophyletic clade, are formally treated as only four classes, 11 orders, and 37 families, with no higher rank assigned. Inside Polypodiopsida, the largest grouping, botanists recognised a string of informal clades, including leptosporangiates, core leptosporangiates, polypods, and the eupolypods split into Eupolypods I and Eupolypods II.

  • PPG I, the 2016 scheme from the Pteridophyte Phylogeny Group, sorts the living plants into two classes. Lycopodiopsida holds three extant orders: Lycopodiales for clubmosses, Isoetales for quillworts, and Selaginellales for spikemosses, each with one extant family. Polypodiopsida, the ferns, spreads across 11 extant orders. Within it, Equisetidae carries the horsetails in a single order, family, and genus, Equisetum. Ophioglossidae gathers whisk ferns and grape ferns, Marattiidae the marattioid ferns, and Polypodiidae the leptosporangiate ferns across seven orders. The largest of those, Polypodiales, alone spans 26 extant families. Beyond the living lie the vanished. Several pteridophyte groups survive only as fossils: the Rhyniopsida, Zosterophyllopsida, Trimerophytopsida, the Lepidodendrales, and the Progymnospermopsida. These extinct lineages mark the deep roots of a grade from which the seed plants themselves would eventually grow.

  • Around 1880, English speakers coined the name Pteridophyte as a Neo-Latin compound. The prefix pterido- means fern, a Latin borrowing of the Greek pterís, which traces back to pterón, meaning feather. The suffix -phyte means plant, from the ancient Greek phyton. So buried inside the formal term is an image of a feather, the very shape a fern frond suggests to the eye. It is a fitting end for a word that has outlasted the taxon it was built to name, surviving on the page and in the lab long after the family it described was shown to be no family at all.

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Common questions

What is a pteridophyte?

A pteridophyte is a vascular plant with xylem and phloem that reproduces by spores rather than flowers or seeds. Ferns, horsetails, and lycophytes such as clubmosses, spikemosses, and quillworts are all pteridophytes.

Why are pteridophytes called cryptogams?

Pteridophytes are sometimes called cryptogams because the word means that their means of reproduction is hidden. They produce neither flowers nor seeds, reproducing instead by spores.

Are pteridophytes a monophyletic group?

No, pteridophytes do not form a monophyletic group and instead constitute a paraphyletic grade. Ferns and horsetails are more closely related to seed plants than to lycophytes, so Pteridophyta is no longer a widely accepted taxon.

How does the pteridophyte life cycle work?

The pteridophyte life cycle involves alternation of generations between a diploid sporophyte that produces spores and a haploid gametophyte, or prothallus, that produces gametes. Both generations are independent and free-living, unlike in seed plants.

How many species of ferns and lycophytes are there?

Christenhusz and Chase in 2014 counted about 10,535 fern species across about 212 genera and roughly 1,300 lycopod species across 5 genera. Ferns account for nearly 90% of living pteridophyte diversity.

Where does the name pteridophyte come from?

The name pteridophyte is a Neo-Latin compound created by English speakers around 1880. It joins the prefix pterido-, meaning fern and derived from the Greek pterón for feather, with the suffix -phyte, meaning plant from the Greek phyton.

What is the PPG I classification of pteridophytes?

PPG I, put forward by the Pteridophyte Phylogeny Group in 2016, divides living pteridophytes into the class Lycopodiopsida with three extant orders and the class Polypodiopsida, the ferns, with 11 extant orders. Its largest fern order, Polypodiales, includes 26 extant families.

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