Monthly Fern: polypodies & fern sex (or do they?)

Left, Polypodium saximontanum, leaves to 25 cm long (Matt Berger); right, P. virginianum, leaves to 40 cm long.

For May, the South Dakota "Monthly Fern" series features polypodies—Polypodium saximontanum and P. virginianum. The genus name comes from the Greek "poly" meaning many and "podion" meaning little foot, referring to the bumps (old leaf bases) on the creeping stems. Polypodies occur worldwide, but are more common in the Northern Hemisphere. About 100 species are recognized, with 11 in North America. They grow mostly on rock (source).

Creeping stem (rhizome) of Common Polypody; bumps on right are old leaf bases. © 2007 Robbin Moran.

South Dakota's two polypodies have a notable distribution—one on each side of the state. P. saximontanum, Rocky Mountain Polypody, grows on granite outcrops in the Black Hills, in the far west. P. virginianum, Common Polypody, is restricted to a small area of Sioux quartzite, in the far east very close to Minnesota (where it's common). It's a good thing they live 350 miles far apart. They're very similar and would be difficult to distinguish if their ranges overlapped.

South Dakota polypodies: Rocky Mountain Polypody in Black Hills (green dot); Common Polypody in Minnehaha County (pink dot); SEINet search May 2025.

Rocky Mountain Polypody on granite, Black Hills, SD (JD McCoy).

Common Polypody on Sioux quartzite, Palisades State Park, SD.

Both species have evergreen leathery deeply-lobed leaves with straw-colored stems. Common Polypody leaves tend to be longer and wider than those of Rocky Mountain Polypody (see first photo). No other clear differences in vegetative characters were found in published descriptions.

As for reproductive structures—which are critical for fern id as we've been told repeatedly—the South Dakota polypodies again are very similar. Both have round sori (spore clusters) arranged in two rows on the underside of leaf lobes; indusia (covers) are absent. The spores are yellow, so much so that even though they're housed in brownish sporangia, they give the sori a yellowish cast.

Common Polypody, P. virginianum (MWI).

Rocky Mountain Polypody, P. saximontanum (Kelly Fuerstenberg).

It is possible to distinguish Common and Rocky Mountain polypodies based on their sori, but it isn't easy. Both have sporangiasters—tiny transparent jelly-like blobs separating the sporangia (1). In P. virginianum, most sporangiasters have gland-tipped hairs, while in P. saximontana, gland-tipped hairs are few or absent. Be forewarned—sporangiasters are said to be so small that one needs macro photos or a good hand lens to examine them.

Polypodium virginianum sori, with sporangiasters with gland-tipped hairs. © 2007 Robbin Moran (arrows added).

Sporangiasters with and without glandular hairs, Polypodium amorphum; very helpful photo by James Thomas.

While we're on the subject of "reproductive" structures, let's address a common misconception (fern buffs excepted). Strictly speaking sori, sporangia and spores are not reproductive structures, for ferns cannot reproduce themselves directly. Instead, their spores give rise to plants quite unlike the parent fern.

This leads us to the fern life cycle, the so-called bugbear of beginning botany students. But we can dispense with complicated details and off-putting terms and still understand and appreciate the curious life of ferns. They and their relatives the lycophytes (formerly "fern allies") are the only land plants that exist as two different free-living beings—kinda like a butterfly and its caterpillar (2).

I find it helpful to first think about flowering plants (angiosperms) with their familiar sex organs. Flowers have eggs in ovaries and sperm in pollen. When a cell of each joins in fertilization, the result is a seed. If conditions are right, the seed germinates and grows into a plant like its parent.

Angiosperm life cycle; note the single free-living being—the plant (modified from source).

But ferns are different. Instead of seeds, they produce millions of tiny asexual spores. If conditions are right, a spore germinates and grows into a minute green plant very different from its parent, even though they have the same DNA. I find this so cool to think about! Unfortunately there seems to be no user-friendly term for these little beings, only "gametophyte" or "prothallus".

Gametophyte of Polypodium vulgare (light microscope at x4 magnification); Viséan.

John Lindsay, a British surgeon working in Jamaica, was the first to describe fern gametophytes (1794), though he didn't call them that and didn't fully understand what they were. Hoping to figure out how ferns reproduce, he had sprinkled "dust" from a fern leaf (today's spores) on dirt in a flowerpot. "I placed the pot in a window of my room, watered it daily, and every day or two examined a small portion of the [dirt] by the microscope ... but observed no alteration till about about the 12th day after sowing." At that point the soil began to turn green "as if it were covered with some small moss".

Lindsay made a nice drawing showing the stages of fern development he observed (full illustration here).

Excerpt from Lindsay's drawing: top, germination; lower right, tiny scales; lower left, first fern leaves.

With the microscope Lindsay could see particles of dust germinating—"pushing out their little germ, like a small protuberance, the rudiment of the new fern" (8–11 above). After a few more weeks the "moss" had grown enough to be visible to the naked eye, looking like small scales (13). These grew to be roundish and bilobate, similar to liverworts (14). Finally a tiny leaf emerged from the scale (15), followed by larger ones (16) until there was a fern like the one that produced the dust. Understandably, Lindsay concluded the dust was fern seed.

It wasn't until the 1840s that botanists finally got rid of fern seed. It had become obvious that despite their alluring beauty, ferns are not sexual creatures. That honor belongs to their gametophytes.

A typical gametophyte, with antheridia and archegonia (source).

On the underside of a gametophyte are little bumps that come alive in the presence of water. Some release wriggling spiral filaments that swim away. Others open to receive a spiral filament if one happens by. These are sex organs: male antheridia release wriggling sperm, and female archegonia each contain an egg. If a sperm wriggles down the neck of an archegonium, it arrives at a large cell—an egg. Fertilization produces a zygote, which develops into a baby fern growing out of the gametophyte (15 in Lindsay's drawing above). If conditions are right, it will become a full-sized fern, thereby completing a life cycle.

Here's the life cycle of a fern, emphasizing the two independent free-living beings that make it so cool! (3)

Fern life cycle—green fern (aka sporophyte) and brown gametophyte (Sigel et al. 2018, much modified).

Once again I'm ending a post without addressing a promised topic. With ferns, it's too easy to go down a rabbit hole! So I will do another Monthly Fern for June—about "the burning question" of how many fern spores fit in a Coke can. What's your guess? Here's a hint: a typical soda can holds 0.355 liters (1.5 cups). And here's an average-sized spore:

Polypodium virginianum spore. Copyright © 2007 by Robbin Moran.

Notes

(1) Sporangiasters may help keep sporangia from drying out prematurely (source). Moran (2017) notes that "In immature sori they form a continuous, protective covering over the young sporangia, thus acting like an indusium."

(2) In thinking about ferns and gametophytes, butterflies and their caterpillars came to mind. In both cases, two forms are produced from the same DNA by using different genes. This is dramatic in butterflies and caterpillars, but not so much in ferns and their gametophytes. In fact, Sigel et al. (2018) found a nearly 90% overlap in genes expressed in Polypodium amorphum ferns and gametophytes. And there's an even bigger difference. Butterflies and caterpillars are both diploid (two sets of chromosomes); there is no independent haploid form that produces gametes—no equivalent of the fern gametophyte. So my comparison of butterflies and ferns was a stretch.

(3) Strictly speaking all land plants alternate between sporophyte and gametophyte life stages (spore- and gamete-producing). But only in ferns and lycophytes are both stages free‐living beings. In seed plants only the sporophyte is free-living; in mosses and liverworts, only the gametophyte is free-living. More here.

Sources in addition to links in post

Lindsay,  John. 1794. Account of the Germination and Raising of Ferns from the Seed. Trans. Linn. Soc, London 2:93–100. BHL.

Moran, RC. 2004. The Natural History of Ferns. Timber Press.

Moran, RC. 2017. Division Polypodiopsida, Ferns in New Manual of Vascular Plants of Northeastern United States and Adjacent Canada. NYBG Press Digital Content (not available as of June 2025, pers. comm.)

Rothfels, C. 2022. Fiddleheads: Fern life cycles and identification. Online workshop for Jepson Herbarium (videos).

Sigel, EM, et al. 2018. Overlapping patterns of gene expression between gametophyte and sporophyte phases in the fern Polypodium amorphum. Front. Plant Sci. 9:1450. FREE

USDA Forest Service. Fern Reproduction.

The Monthly Fern: Bracken—dreadful or delightful?

"Shelter" by Colin.

Come my sweet and let us lie in

Some idyllic wooded glade

And let us stay til merry-made

Amid the Bracken.

For April, the South Dakota fern-a-month series features the world's most widespread fern—Pteridium aquilinum, Bracken (aka Pasture Brake, Eagle Fern, Helecho Macho, and more). Thought to be native to the Northern Hemisphere, it's now widely naturalized and known from all continents except Antarctica. In South Dakota, it grows in the Black Hills in the western part of the state (1).

Bracken is said to thrive in a variety of habitats—woodlands, fields, old pastures, thickets, disturbed soils, burned areas, and marshes. But sources vary on this. For example, some say it's intolerant of wet soil and shade; others say it can grow well in all but very alkaline soils. In any case, Bracken forms extensive colonies of robust plants to 1.5 m or more tall, from deep rhizomes to 20 feet long. The large triangular leaves (fronds) are twice or thrice pinnately compound (2- or 3-times divided into leaflets). When fully grown, the blades often bend to horizontal, shading much of the ground (source).

Leaf division is an important character in fern identification, but can be hard to understand and explain. However I will try. Below is a thrice pinnately compound Bracken leaf. It's divided into 9 large segments (one terminal), which are divided into many narrow segments, which are divided (or nearly so) into small ultimate segments. "Pinnately compound" means segments line up on each side of an axis (rachis). More here.

Bracken frond by Olegivvit (labels added).

As in most ferns, Bracken's spores are borne on the underside of leaves in clusters called sori (remember?). In Bracken, sori are continuous along leaflet margins. In youth, the sori are covered; with maturity, leaflet margins unroll, exposing mature sporangia (spore shooters).

In Bracken, young sori are protected under rolled leaflet margins, sometimes with tiny membranous indusia (flaps, click image to view); Zharkikh photo.

In this frond, leaflet margins have unrolled and lines of brown sori are visible; Zharkikh photo.

Brown "beads" are sporangia; each contains many minuscule spores ready to be "shot"; Zharkikh photo.

My first encounter with Bracken was in the Bear Lodge Mountains in the northwest Black Hills. That was at least 40 years ago, but the memory remains vivid. In a stand of tall quaking aspen, Bracken's horizontal fronds formed a lovely lacy ground cover. It was an idyllic setting, and still comes to mind when I think about Pteridium aquilinum.

That memory prompted me to search for a bit of English poetry about Bracken (it has close ties to moorlands). But it was rarely mentioned, and never in an idyllic setting (2). Perhaps poets know of Bracken's reputation. It's not a particularly nice fern, and there are many reasons to dislike it. The horrors that follow were provided by Robbin Moran, a man who who loves ferns!

Bracken grove with conifers and ferns and little else; Charlesblack photo.

The common objection to Bracken is that it's weedy—in fact a noxious invader. With its vigorous growth and colonial habit, it swamps (with litter) or shades out other species. Eradication is difficult due to its deep extensive rhizomes.

Other dangers lie hidden. Bracken is filled with nasty stuff, ostensibly for defense against insects and other herbivores. It contains at least two kinds of insect hormones, which cause uncontrolled molting and death in any insect that eats it. It also contains thiaminase, an enzyme that breaks down vitamin B1. This makes Bracken hazardous to livestock, which often find it palatable. Overconsumption will cause thiaminase-induced staggers (treatable with vitamin B1, thiamine).

And there's more. Bracken is rich in tannins and therefore bitter-tasting, which is good. For if consumed, for example in the absence of other forage, tannins inhibit enzymes critical to cellular metabolism. Bracken also produces a deadly chemical weapon, hydrogen cyanide, in response to tearing of leaf tissue, thereby deterring or killing the perpetrator.

Gosari, a popular Korean dish of Bracken fiddleheads; Hyeon-Jeong Suk photo.

Bracken fiddleheads (young shoots, also called croziers); Phil Gayton photo.

After reading of Bracken's many hazards, I was surprised to learn that people happily eat its fiddleheads. But this is dangerous too. Though cooking removes tannins and thiaminase, carcinogens remain, and increased rates of stomach and esophageal cancers have been reported where fiddleheads are popular, for example in Japan, Korea, and Britain.

The main carcinogen is the compound ptaquiloside (3). It occurs throughout Bracken plants, but is highly concentrated in young growth, in spring and early summer. Humans take up ptaquiloside mainly by eating fiddleheads, but there are other sources—airborne spores, milk and meat from affected animals, and contaminated ground and surface water where Bracken grows (source).

Now that I know of Bracken's nefarious ways, do I feel foolish about my early love affair? No, for it also offers delights, as the Radnoshire Wildlife Trust in Wales explains: "Bracken can be an important habitat in its own right.  It supports over 40 species of invertebrate, forming an important part of the diet for 27 of these while 11 are found only on bracken. It is an important breeding habitat for moorland birds ... and reptiles and mammals benefit from its shelter." It's also a great candidate for areas in reserves and gardens where nothing else will grow. Just keep an eye on it!

Finally, Bracken is beautiful, in fact so beautiful that its lovely lacy fronds make a woodland irresistible. Even Robbin Moran agrees. "The grove seems so peaceful and idyllic" he writes, though he knows that in the shadows there lurks a femme fatale.

"Afternoon light" by Colin.

PS Last month I promised to include the fern life cycle in April's Monthly Fern post. But being overwhelmed by Bracken's dark side, I've postponed "the bugbear of botany students" until May. You're off the hook for now! But this also means you must wait for an answer to the burning question: "How many average-sized fern spores does it take to fill a can of Coke?" Stay tuned.

Notes

(1) I'm surprised Bracken hasn't been reported from eastern deciduous forests in the far eastern counties of South Dakota. Looks like good habitat to me, and it grows in Minnesota not all that far away.

(2) In the absence of suitable poetry, I wrote my own (I promise I won't do it again). I did find a poet named Bracken, as well as Bracken, a literary magazine: "Bracken is green and lush, coarse and delicate, drinks from the earth, and spreads underground, more root than frond. Bracken is understory, invades, takes over, shades and protects. We seek poetry and art that will root, tender and tough, in us."

(3) At least one of Bracken's insect hormones also is a carcinogen (source).

Sources

Moran, RC. 2004. The Natural History of Ferns. Timber Press.

Royal Horticultural Society. Advice—Bracken.

Stone, J. A successful fern, or a case for control. Radnorshire Wildlife Trust (blog).