those birds unknown, that left only footprints

Big toes! Huge bird!!

Along the rim of Bull Canyon, on the north slope of the La Sal Mountains in southeast Utah, my field assistant and I followed footprints in sand. Three-toed two-footed creatures had passed this way when the sand was wet, before it turned to rock. They were common then, traveling in packs.

Similar 3-pronged impressions occur 2000 miles to the east, in the Connecticut River Valley in Massachusetts. Some are famous—among the earliest to be studied and published. They were found in 1835 by Dexter Marsh, who was laying a flagstone sidewalk. He showed the slabs to the owner of the property who gave them to a physician who then gave them to state geologist Edward Hitchcock.

"They consist of two slabs, about forty inches square, originally united face to face; but on separation, presenting four most distinct depressions on one of them, with four correspondent projections on the other; precisely resembling the impressions of the feet of a large bird in mud." (Hitchcock 1836, italics mine)

Sandstone slabs, each 36.5" x 34"; depressions (molds) on left, projections (casts) on right (source, click on "Fossil Slabs Found by Dexter Marsh").

Hitchcock was understandably excited. Very few bird fossils had been found anywhere, and geologists had decided that because most birds were lightweight creatures of the air, they were unlikely to be submerged and preserved on the bottom of lakes, oceans and such. "Even when they chance to perish in the water, they float so long upon the surface, as to be most certainly discovered, and devoured by rapacious animals."

As it turned out, such marks were fairly common in the area. Hitchcock studied slabs from five quarries, concluding that the impressions must have been made by birds:

1. These impressions are evidently the tracks of a biped animal. For I have not been able to find an instance, where more than a single row of impressions exists.

2. They could not have been made by any other known biped, except birds. On this point, I am happy to have the opinion of more than one distinguished zoologist.

3. They correspond very well with the tracks of birds.

Some of Hitchcock's drawings of modern-day bird tracks, from his 1836 publication (source).

However, many of the tracks Hitchcock studied were too large to have been made by the birds we know—to 18 inches long and 13 inches wide, and separated by 6-foot strides. Therefore these tracks must have been made by large birds now extinct. Eminent geologists of the day agreed with Hitchcock (2). But by the end of the century they had been "proven" wrong. These tracks were not avian, they were reptilian (Dean 1969).

Science marches on of course, and we now know that in a sense, Hitchcock and his colleagues were correct. The creatures that left footprints in the Connecticut River Valley and southeast Utah were indeed birds. But they also were dinosaurs, specifically theropods (study the images below before discussing this at cocktail parties).

Dinosaur classification (3). While all birds are dinosaurs, not all dinosaurs are birds; similarly, birds are a subset of theropods (from Zureks).

Evolution of birds from a dinosaur ancestor; Manti-La Sal National Forest, Bull Canyon Tracksite.

You might be wondering why dinosaurs traveled the rim of Bull Canyon. Well ... actually they didn't. There was no Bull Canyon 157 million years ago. Instead this was a broad coastal plain, where large bipeds could cruise along at 2.5–3.5 mph (Hunt-Foster 2016).

The Bull Canyon Tracksite includes at least 50 well-preserved large theropod tracks, to 18 x 14 inches in size. But aside from footprints little is known about these creatures, for no bones have been found. So rather than naming a species, paleontologists named their tracks: Megalosauripus. These are ichnofossils—"a fossil record of biological activity by lifeforms but not the preserved remains of the organism itself." They're also called trace fossils, the term I learned.

 Theropods passed this way, in a pack perhaps.

They were big! (40-pound dog for scale).

Megalosauripus is a theropod track, not the theropod itself.

The wet sand where theropods once walked is now sandstone, part of the Moab Member of the Curtis Formation, dating from 157 million years ago (Late Jurassic). The setting was dynamic—changing sea level, oscillating shoreline, occasional sand dunes—making classification and dating of rock units difficult (Mathis 2021). But no matter. Whatever geologists decide to call the rock, its theropod tracks go on and on and on. They occur across Arches National Park, east to the Bull Canyon area and the Colorado–Utah state line, and perhaps as far south as Blanding. This is the Moab Megatracksite, also known as the Dinosaur Freeway. A conservative estimate of its size is 700 square miles; as of 2016, c. 3000 tracks had been reported from 30 sites (Hunt-Foster 2016).

Dinosaur Stomping Grounds, aka Jurassic Dancefloor, with at least 2000 theropod tracks (Sierra Club).

Our visit to the Bull Canyon Tracksite last fall was but a brief introduction. Many more opportunities to commune with large extinct birds await. Fortunately many of the sites are on public land, and there's a handy guide available (Hunt-Foster 2016). We shall return!

Dreaming of giant birds after a day in the field.

Notes

(1) The title of this post comes from a poem by Henry Wadsworth Longfellow—To the Driving Cloud. He referred to fossil bird tracks in several poems (Dean 1969).

(2) Edward Hitchcock was a cleric and amateur geologist. In his time, so-called amateurs made major contributions, his study of fossil birds being a good example. Charles Lyell confirmed Hitchcock's findings, and included them in lectures and later editions of Principles of Geology. Louis Agassiz and others also spread the word (more in Dean 1969).

(3) Are you wondering, as I did, why birds are NOT included in the seemingly eponymous Ornithischia (lower right in diagram)? That group includes dinosaurs with hips that superficially resemble those of birds. Maybe the name predates the realization that birds evolved from a theropod.

Sources (in addition to links in post)

Dean, DR. 1969. Hitchcock's Dinosaur Tracks. American Quarterly 21:639–644. https://www.jstor.org/stable/2711940

Hitchcock, E. 1836. Ornithichnology—description of the footmarks of birds, (Ornithichnites) on New Red Sandstone in Massachusetts. American Journal of Science and Arts, XXIX:307-40. Internet Archive.

Hunt-Foster, RK, et al. 2016. Tracking Dinosaurs in BLM Canyon Country, Utah. Utah Geological Association, Geology of the Intermountain West, Vol. 3. PDF

Mathis, A. 2021. Moab, Goblin Valley, and the Curtis Formation. Moab Happenings Archive.

The Monthly Fern: Water Clover & its odd spores

"It is worth clarifying that these plants are not clovers." (Photo by Bill Dodd).

For July, South Dakota's fern-of-the-month is the Hairy Water Clover, Marsilea vestita. Among our ferns it's quite the oddball—in habitat, behavior, leaves, and especially the spores. They suggest that one of its ancestors was the evolutionary start of seed plants!

Water Clovers grow nearly worldwide. On the order of 45 to 65 species are recognized (experts disagree on number), of which 5 are native to North America. M. vestita, the Hairy Water Clover, is the only species in South Dakota (so far). It's known from many sites across the state, in shallow water and on mud. Plants are rooted (not free-floating) and tolerate seasonably dry conditions, for example persisting after a pond has dried up. In fact, wet followed by dry can aid in reproduction and dispersal (Montana Field Guide).

Marsilea vestita is rhizomatous and colonial; in wet habitat, leaflets usually are horizontal (MWI).

M. vestita in a field; in drier habitat, leaflets often are ascending (Mary Ellen (Mel) Harte photo).

Hairy Water Clover can grow as tall as 20 cm on moist soil, or to 40 cm in water (in order to reach the surface). Leaves are dimorphic—sterile and fertile—but neither is fern-like. Sterile leaves have blades with four rounded triangular leaflets, and look a lot like four-leaf clovers! No other plant can be confused with Marsilea (1). 

Sterile leaves of Hairy Water Clover have divided blades on long slender stalks (MWI).

The sterile leaves also are unusual in behavior. Water Clovers are the only ferns known to be nyctinastic—moving with the onset of darkness. During the day leaf segments are nearly horizontal. Then as the sun sets they bend upward, forming a packet of sorts (Montana Field Guide has an account of this and other interesting features of Water Clover).

Fertile "leaves" are located near the base of sterile leaf stalks. In shape and size they resemble beans or peppercorns (source of another common name, Pepperwort). Being unusual they of course have a special name—sporocarp (= spore body); oldtimers like Linnaeus called them capsules.

Marsilea vestita. The hairy sporocarps contain 2 kinds of spores (lower right). Britton & Brown 1913.

Young sporocarps are greenish, hairy and slightly soft. With maturity they dry out, darken and become very hard. In this state they can survive for many years; the record is said to be 100. The Marsilea sporocarp is an effective unit of dispersal, often by way of waterfowl digestive tracts.

Like almost all ferns, Water Clovers have sori—clusters of sporangia which contain the spores. But the arrangement is quite different. In a typical fern, sori are located on the underside of leaves. In Marsilea, they're neatly arranged inside the sporocarp.

A typical fern; sori are clusters of sporangia, which contain dust-sized spores (2). USDA Forest Service.

Marsilea sporocarp with sori; below it, a sorus with sporangia, which release spores (no source given).

A cross-section through a Marsilea sporocarp (above) reveals an orderly but complicated interior. Inside the container-like sori are 2 kinds of sporangia. This is where things get exciting. They produce 2 kinds of spores—male and female!

Most ferns, 99% in fact, release a single type of spore—tiny, 1-celled, asexual. But not Water Clovers. They're among the 60 fern species (out of c. 10,500 total) with male microspores and female megaspores. These are heterosporous ferns; interestingly, all are aquatic (more here).

When a Marsilea microspore bursts open, many sperm (aka spermatozoids) swim off in search of an egg to fertilize—not unlike sperm of typical ferns. It's the megaspores that are so unusual. Not only are they 10 times the size of a typical fern spore, they're complex, with specialized parts.

Marsilea megaspore, c. 0.8 mm long, with 2 cells (no source given).

Shortly after leaving its sporangium, the megaspore divides to become two joined cells. The upper cell will produce an egg, which gives off chemicals to attract sperm. If a sperm successfully wriggles through the opening and reaches the egg, fertilization takes place, leading to development of an embryo and then a baby fern.

In comparison, the megaspore's basal cell is huge, and rich in carbohydrates and fats. These will sustain the developing fernling in its first days, before it can photosynthesize. In this way, a megaspore is like a seed, which supplies nutrients for its young seedling. Perhaps an ancestor of Marsilea was the evolutionary beginning of seed plants (more here).

Some readers may be wondering where the gametophytes are—those tiny independent plantlets that are the sexual stage of ferns. Good question! Water Clovers do have gametophytes, but they are minute and NOT independent (another similarity to seed plants). For more about the life cycle of heterosporous ferns, see (3) in Notes.

Simplified?

Now we finally arrive at the long-promised answer to the burning question, "How many spores would fit in a typical [empty] can of soda?"

Fern spores are truly tiny. A handful looks like a pile of dust. To show just how small they are, Robbin Moran (2021) calculated the number of average-sized spores that would fit in a typical can of soda, which has a capacity of 355 milliliters. For spore volume, he used 125,000 µm3, assuming for simplicity that a spore is a cube. What do you think? How about a ballpark estimate?

Hmmm ... 777,000?

Maybe 10 million?

According to Robbin's calculations the answer is 4,440,000,000 (4.4 billion). Yikes, that's a lot! Yes, spores are tiny indeed (4).

Notes

(1) While Water Clovers are easily recognized, distinguishing the Hairy Water Clover from others in the genus is not easy, requiring sporocarps. If you intend to document an occurrence, be sure to collect both types of leaves. See Marsilea in Flora of North America for a species key and descriptions.

(2) It seems sporangia are readily mistaken for spores, as a search for images of "fern spores" suggests. Many of the images actually are sporangia clustered in sori.

(3) In the previous Monthly Fern, I made a big deal out of the 2-stage life cycle of ferns, which involves separate tiny green sexual plantlets—gametophytes—that give birth to baby ferns. Heterosporous ferns have gametophytes, but they are minute and not independent. They develop inside the persistent spore wall, where they give rise to either sperm-producing antheridia or egg-producing archegonia (gametophytes of typical ferns usually have both). For the female gametophyte, developing inside the spore wall provides additional protection for the embryo, but there's no access to sunlight to photosynthesize food for the fernling. That's why the nutrient-rich basal cell is so important.

Life cycle of Marsilea, a heterosporous fern (labels added). See Milne Publishing for details.

(4) I couldn't find Robbin Moran's article online. If you'd like to read more about fern spores, and all of Robbin's calculations and conclusions (e.g. 4.44 billion spores taken together has a surface area nearly equal to 8.5 ping-pong tables), send me an email address and I'll send you a PDF file.

Sources (in addition to links in post)

Hooker, WJ, and Greville, RK. 1831. Figures and descriptions of ferns, principally of such as have been altogether unnoticed by botanists, or as have not yet been correctly figured. Vol. 2. BHL

Milne Publishing. Marsilea. Accessed June 2025.

Montana Field Guide. Montana Natural Heritage Program. Hairy Water Fern—Marsilea vestita.

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

Moran, R. 2021. Fern Spores, Soda Cans, and Ping-Pong Tables. Fiddlehead Forum (May–Dec).

Pinson, J. About Ferns, American Fern Society.

PremaBotany (Prema Iswary). December 2018. Marsilea.