Bestia longipes (Sullivant & Lesquereux) Brotherus

Bestia longipes is a large (to 12 cm), pleurocarpous moss, with soft, regularly pinnate branches, bright green when moist, olive green to brown when dry, which grows in large loose mats on rock walls and boulders in canyons along the immediate coast.
Scroll down for more photos (click to enlarge) and descriptions of habitat, male and female plants, leaves, sporophytes, distribution map, name origin, and similar species.
Bestia longipes
Bestia longipes CC BY-NC Amanda Heinrich
Habitat:  Bestia longipes is found on rock walls, large boulders, and occasionally the bases of trees, along streams in canyons, primarily on the immediate coast of California.
Bestia longipes
Bestia longipes moist, on a rock wall in a canyon on the immediate coast. CC BY-NC Amanda Heinrich
Bestia longipes
Bestia longipes dry, on rock wall in a canyon on the immediate coast. CC BY-NC Amanda Heinrich
Habit:  Bestia longipes plants dangle softly downward in loose mats, often extensive. When moist, the plants’ regularly pinnate, bright green branches look like tiny ferns. When dry, branches curl gently and plants often turn to the side, becoming more olive green-brown.
Bestia longipes
Bestia longipes moist, dangling out and down like soft miniature ferns. CC BY-NC Amanda Heinrich
Bestia longipes
Bestia longipes dry, branches gently curled, many plants turned to the side. CC BY-NC Amanda Heinrich
Plants:  Bestia longipes plants are quite large, up to 12 cm, and have fairly regular, short, pinnate branches.
Bestia longipes
Bestia longipes plants moist, branches once or twice pinnate (CAH #931). CC BY-NC Amanda Heinrich
Bestia longipes
Bestia longipes plants dry, gently curled (CAH #931). CC BY-NC Amanda Heinrich
Leaves:  Bestia longipes leaves are held away from the stem when moist, and are loosely appressed to the stem when dry. Leaves are triangular-ovate, slightly concave, 1.5-2 mm long, with those on the stem larger than those on the branches. The margins are recurved from the base of the leaf to about 3/4 up, and are toothed near the apex of the leaf. The costa is also toothed near the apex.
Bestia longipes
Bestia longipes leaves spread from stem when moist (CAH #931). CC BY-NC Amanda Heinrich
Bestia longipes
Bestia longipes leaves are loosely appressed to the stem when dry, their slight concave shape resulting in a bit of a poofy look to the branches and stem (CAH #931). CC BY-NC Amanda Heinrich
Bestia longipes stem leaf
Bestia longipes stem leaf, triangular-ovate and slightly concave, margins recurved to ~ 3/4, viewed from bottom (CAH #931, 40x). CC BY-NC Amanda Heinrich
Bestia longipes leaf margin
Bestia longipes recurved leaf margin (CAH #931, 400x). CC BY-NC Amanda Heinrich
Bestia longipes leaf tip
Bestia longipes toothy leaf tip (CAH #931, 400x). CC BY-NC Amanda Heinrich
Bestia longipes costa spine
Bestia longipes spiny costa near leaf apex (CAH #931, 400x). CC BY-NC Amanda Heinrich
Bestia longipes branch leaf
Bestia longipes branch leaf, also triangular-ovate with recurved margins, but more petite than stem leaf, viewed from above (CAH #931, 100x). CC BY-NC Amanda Heinrich
Perichaetia:  Bestia longipes appears to be dioicous, with separate male and female plants. The female sex organs, archegonia, are housed within budlike perichaetia situated along the main stem. Each perichaetium houses many narrow, flask-shaped archegonia, each of which houses an individual egg. Upon fertilization by a sperm, the now diploid zygote develops into the sporophyte.
Bestia longipes female
Bestia longipes bud-like perichaetia along stem (CAH #1021). CC BY-NC Amanda Heinrich
Bestia longipes archegonia
Bestia longipes dissected perichaetium housing reddish, flask-shaped archegonia interspersed with colorless finger-like paraphyses (CAH #1021, 100x). CC BY-NC Amanda Heinrich
Perigonia:  Bestia longipes male plants have tiny onion-like perigonia along the primary branches. Each perigonium houses several sausage-shaped antheridia, which in turn contain thousands of sperm.
Bestia longipes male
Bestia longipes male plant, with tiny onion-like perigonia along branch stems (CAH #931). CC BY-NC Amanda Heinrich
Bestia longipes male branch
Bestia longipes brown onion-shaped perigonia house male sex organs (CAH #931). CC BY-NC Amanda Heinrich
Bestia longipes perigonium
Bestia longipes perigonium, paritally dissected to reveal sausage-shaped antheridia (CAH #931, 100x). CC BY-NC Amanda Heinrich
Bestia longipes antheridium
Bestia longipes individual antheridium with associated transparent, finger-like paraphyses (CAH #931, 100x). CC BY-NC Amanda Heinrich
Sporophytes: When an egg in an archegonium within a perichaetium is fertilized by a sperm from a perigonium, it develops into a sporophyte, the part of the plant that produces spores. Accordingly, sporophytes are found along the stem of female plants. Sporophytes are not often seen on Bestia longipes plants, but 2023 was a bumper crop year here in Santa Barbara County.
Bestia longipes sporophyte
Bestia longipes sporophytes. CC BY-NC Amanda Heinrich
Bestia longipes perichaetia
Bestia longipes perichaetia in the field. CC BY-NC Amanda Heinrich
Distribution:  Bestia longipes is endemic to California, possibly reaching into southern Oregon (one collection from 1935). It is not exactly plentiful in Santa Barbara County, but can be found within each stream canyon along the coast, sometimes in extensive populations.
Name: The genus name, Bestia, honors George Newton Best, an American bryologist, who lived from 1846-1926. “longus” is Latin for long, and “pes” is Latin for foot or stalk, though it is unclear to which stalk this refers – stem or seta.
Similar Species in Santa Barbara County
Beginners may confuse Bestia longipes with Dendroalsia abietina, another large moss with regularly pinnate branches.  Dendroalsia abietina is typically larger, and more robust, with much stiffer and wirier stems and branches. When dry, its leaves overlap neatly and tightly, and plants curl up in a highly characteristic manner. Additionally, Dendroalsia abietina is less restricted in its habitat, and is found on rocks and trees away from streams and canyons.
Dendroalsia abietina on the left, with two Bestia longipes plants on the right, moist. CC BY-NC Amanda Heinrich
Dendroalsia abietina with Bestia longipes
Dendroalsia abietina on the left, with two Bestia longipes plants on the right, dry. CC BY-NC Amanda Heinrich
References and Further Reading
Bestia longipes, James R. Shevock, Flora of North America Vol. 28, www.efloras.org
California Mosses, Bill and Nancy Malcolm, Jim Shevock and Dan Norris. Available for purchase from the California Native Plant Society here.
Contributions toward a Bryoflora of California II. A Key to the Mosses, Daniel H. Norris and James R. Shevock, Madrono 51(2): 133-269. 2004. Available for purchase  here.