The Monthly Fern??—Prairie Spikemoss

Selaginella densa—moss, fern, fern ally, or none of the above? Coin is 19 mm across.

This month the South Dakota fern series features another oddity—a spikemoss, genus Selaginella. It's even more unusual than last month's Water Clover, for while water clovers are ferns, spikemosses are not, at least not anymore. So where in the greater scheme of plant classification do they belong?

The Prairie or Dense Spikemoss, Selaginella densa, is the more common of South Dakota's two spikemosses. It occurs in the Black Hills and scattered across the west half of the state. If you live in or have wandered across the Great Plains or Rocky Mountains, you may have seen it, for it grows on a wide range of sites—prairies, alpine meadows, dry rocky slopes, rock crevices, sandstone, quartzite or granite rock, and dry gravelly, clayey or sandy soil (Flora North America). Or maybe you overlooked it, as I used to do. After all, it looks very much like a moss (1).

Spikemosses were first classified—given a name and assigned to a plant group—by the great Swedish botanist Carl Linnaeus, founder of today's system of naming organisms. In his Species plantarum (1754) he placed them in the CRYPTOGAMIA MUSCI section—the mosses. That was a big mistake, but at the time it was a reasonable decision. Like mosses, Selaginella produces spores (2). But unlike mosses, it has vascular tissue—plumbing for transporting water and nutrients.

In the late 1890s another Swedish botanist was studying spikemosses, while preparing a Catalogue of the Flora of Montana and the Yellowstone National Park. Per Axel Rydberg had emigrated to the United States in 1882, hoping for a career as a mining engineer. But after a serious injury in an iron mine in Michigan, he moved to eastern Nebraska to teach mathematics. He also studied botany at the University of Nebraska—the beginning of a "lifelong devotion to plant studies" in the Great Plains and Rocky Mountains (source).

In 1895 and 1896, Rydberg was sent to Montana by the US Department of Agriculture to collect grasses and forage plants. The next summer he returned, with the first field expedition of the New York Botanical Garden. He made about 1800 collections representing 800 species—20,000 specimens in all (replicates were collected for exchange or sale to other institutions).

Rydberg's Catalogue included a large foldout map showing localities mentioned in the text. He noted that the eastern half of the state was "practically unexplored botanically." (BHL)

Going through his collections that winter, Rydberg saw that the flora of Montana was poorly known, even with his additions. "It was therefore considered advisable to extend the work and study all the material from the state that was accessible." He examined specimens from 16 institutions and private individuals, ranging from the Lewis & Clark collection (1803–1804) to the Montana Ladies' [Columbian] World's Fair Set (1893). By the time the Catalogue was published in 1900, Rydberg had added 776 species to the flora of the Rocky Mountain region, including 163 novelties—species new to science (Rydberg would become known as a notorious splitter).

Among the novelties was a low densely-tufted plant with very short stems covered in bristle-tipped leaves 3–5 mm long. Fertile stems were taller, to c. 4 cm, with spore-bearing leaves (sporophylls) neatly arranged in four ranks, forming terminal strobili (aka cones).

Prairie Spikemoss forms dense mats in this soil crust. Matt Lavin photo.

Selaginella densa's 4-angled strobili rise above very short sterile stems that look like clusters of bristle-tipped leaves. cinthyadasilva photo.

Rydberg knew the plant was a spikemoss, but the dense "moss-like" form was not something he had seen before. After careful study of seven specimens, he concluded it was a new species, calling it Selaginella densa. The holotype (basis for formal description) was a specimen collected in 1889 by Valery Havard, a French-born American military physician, explorer and botanist.

Havard identified his specimen (NYBG) as S. rupestris, which is widespread in the east half of the US.

In his Catalogue Rydberg followed the accepted classification of the day. He included Selaginella densa in the Pteridophytes—spore-bearing vascular plants, mainly ferns. He put it near the end of the section, with horsetails, clubmosses and other oddballs. These were the Fern Allies. Like ferns they bore spores, had vascular tissue, and reproduced via two distinct independent life stages. But otherwise they were decidedly unfernlike, and very different from each other. Just look below!

Horsetails and scouring rushes, genus Equisteum, have jointed stems with cylindrical sheaths tipped with teeth. These are thought to be highly modified leaves. Spores are born in terminal cones.

Unbranched species of Equisetum are called scouring rushes. Andre Zharkikh photo.

Whisk ferns, genus Psilotum, have linear shoots that fork in the upper half. Minuscule scale-like leaves subtend globose spore containers 2–3 mm across.

Sideways view of a whisk fern. Mary Keim photo.

Quillworts, genus Isoetes, are aquatic, with grass-like clusters of linear leaves. Spores are born in sac-like structures in enlarged leaf bases.

Bolander's Quillworts in a lake in the Wasatch Mountains, Utah. Andrey Zharkikh photo.

Clubmosses, family Lycopodiacee, are a more diverse group, with 7 genera and 27 species in North America. Some are suggestive of spikemosses; in fact spikemosses were put in the genus Lycopodium by Linnaeus.

These clubmosses, all formerly genus Lycopodium, are now 4 separate genera; from Ferns and Evergreens of New England, 1895 (BHL).

The Fern Allies group came into use in the early 1800s, as a catchall for diverse, puzzling, somewhat fernlike plants. But after about a century botanical experts began to object. Some Allies appeared to be more closely related to ferns, others not so much. Then less than a century later, the Allies got caught up in a revolution. Biologists were switching to a phylogenetic approach to classification. In a nutshell (a very tiny one), they now hope to classify organisms based on evolutionary relationships, i.e., so that all members of a group share a common ancestor. The Fern Allies do not, so they were reclassified (3).

The commonly accepted classification splits the Allies into two groups that diverged long ago, early in the evolution of vascular plants. One includes ferns, horsetails, whisk ferns and seed plants. The other group is much smaller, a collection of relatively primitive plants: quillworts, clubmosses and spikemosses. These are lycophytes (answer to question at top of post). For a longer summary, see The Ferns and their Allies at Cliffnotes. For a deep discussion, start with Pteridophyte taxonomy on Wikipedia.

Fern and lycophyte classification from the Pteridophyte Phylogeny Group. Black labels added, not sure how that guy in the corner snuck in.

Notes

(1) Not all spikemosses are as humble and mosslike as ours. Selaginella is a large genus with c. 800 species, mainly of the tropics and subtropics. In hospitable habitat, spikemosses can be quite showy—some are iridescent!

Selaginella uncinata, Blue Spikemoss, is native to moist shady sites in southern Chile and is widely cultivated; leaves are 3–4 mm long (Flora North America). GKA Dickson photo.

(2) Actually Linnaeus couldn't decide whether "fern dust" was pollen or seeds. The concept of spores would come later.

(3) It's been really hard to give up Fern Allies! It's such a handy label for those diverse kinda-fernlike species. Not surprisingly, the name hasn't gone away. Sometimes it appears under an alias, for example "Fern Relatives" in Ferns of Northeastern and Central North America (2005). More often it pops up in casual conversation, or is used by older botanists who haven't bothered to learn the new scheme. After resorting to "Fern Allies" in a message to pteridologist Robbin Moran, I committed to learning it (Robbin is much too kind to disapprove directly, but he did refer to "lycophytes" in his reply).

Sources (in addition to links in post)

Cobb, B, Farnsworth, E, Lowe, C. 2005. Ferns of Northeastern and Central North America. 2nd ed. Peterson Field Guide Series.

Linné, Cv. 1754. Species plantarum v2. BHL

Moran, Robbin. 2004. A Natural History of Ferns. Timber Press.

Rydberg, PA. 1900. Catalogue of the Flora of Montana and the Yellowstone National Park. Memoirs of the New York Botanical Garden. BHL