Marsupella bolanderi Austin (Underw.)

Marsupella bolanderi is a small (<6mm high, 0.1-0.2mm wide) leafy liverwort, with two rows of green to reddish bilobed leaves, that often overlap near the apex of the shoot, but are more remotely spaced further down the stem. The species is dioicous, with separate male and female plants. It is typically found growing on sandy soil or on sandstone itself, either in sun, where plants turn reddish, or in shade, where plants are greener. Plants may grow upright or more prostrate and may form loose or dense patches.
Scroll down for photos with labeled parts and more detailed descriptions.
Marsupella bolanderi
Marsupella bolanderi (with Cephaloziella divaricata) CC BY-NC Amanda Heinrich
Detailed Descriptions with Photos (click to enlarge)

Individual Marsupella bolanderi plants are up to 6 mm long. Upper leaves are often reddish and densely overlapping, while lower leaves, less exposed to the sun, are often greener, as well as more distantly spaced along the stem.

Marsupella bolanderi plants
Marsupella bolanderi plants, CAH #1237. CC BY-NC Amanda Heinrich

Marsupella bolanderi has two rows of leaves, which are transversely inserted into the stem, that is, they stick out at a right angle. The leaves are slightly concave and spreading. Whereas leaves at the apex of the shoot overlap one another, those below are remotely distributed along the stem such that the stem and the colorless rhizoids are clearly visible when viewed under a microscope. In contrast to many other leafy liverwort species, Marsupella bolanderi lacks a third row of smaller underleaves.

Marsupella bolanderi stems
Two Marsupella bolanderi plants at 40x, CAH #700. CC BY-NC Amanda Heinrich

The leaves of Marsupella bolanderi are bilobed, with the sinus between the lobes extending up to a third of the length of the leaf. The tips of the lobes are acute to obtuse, resulting in triangular to ovate lobes.

Marsupella bolanderi stems
Marsupella bolanderi bilobed leaves, CAH #700. CC BY-NC Amanda Heinrich

Individual Marsupella bolanderi leaves are bilobed, with the sinus extending up to 0.3 the length of the leaf. Cells along the leaf margins are often radially elongate, that is, wider than high.

Marsupella bolanderi leaf
Marsupella bolanderi leaf, 100x, CAH #1237. CC BY-NC Amanda Heinrich

In this photomicrograph of a Marsupella bolanderi leaf magnified at 400x, one can see the reddish (vinaceous) cell walls and the rather small, triangular spaces between cells (trigones). This is in contrast to M. emarginata (not reported in Santa Barbara County), which has large, conspicuous trigones. The oil bodies within the cells are also visible, resembling funny little potatoes.

Marsupella bolanderi trigones
Marsupella bolanderi leaf cells with trigones, CAH #700. CC BY-NC Amanda Heinrich

Occasionally one will enjoy the privilege of finding Marsupella bolanderi with sporophytes. Sporophytes consist of a black spherical capsule upon a flimsy, translucent seta.

Marsupella bolanderi sporophyte
Marsupella bolanderi with a single sporophyte, CAH #1237. CC BY-NC Amanda Heinrich

Marsupella bolanderi sporophytes are quite transient, with capsules opening along 4 lines of dehiscence to release spores, and watery setas collapsing shortly thereafter. The sporophytes shown here are now draped over the long-lived, persistent gametophytes.

Marsupella bolanderi sporophytes
Marsupella bolanderi sporophytes, CAH #1237. CC BY-NC Amanda Heinrich

Marsupella bolanderi sporophytes consist of a spherical black capsule atop a pale flimsy seta emerging from the perianth at the apex of the female plant.  

Marsupella bolanderi sporophyte micrograph
Marsupella bolanderi sporophyte at 40x, CAH #1237. CC BY-NC Amanda Heinrich

At maturity, Marsupella bolanderi sporophyte capsules open along 4 lines of dehiscence to release the spores.

Marsupella bolanderi open sporophyte micrograph
Marsupella bolanderi sporophyte capsule opening along 4 lines of dehiscence, at 40x, CAH #1237. CC BY-NC Amanda Heinrich

Spring-like elater cells help to disperse Marsupella bolanderi spores.

Marsupella bolanderi spores and elaters
Marsupella bolanderi spores and elaters, at 400x, CAH #1237. CC BY-NC Amanda Heinrich

Marsupella bolanderi is a dioicous species, with separate male and female plants. Antheridia, the male sex organs, appear as green spheres nestled amongst the upper leaves of male plants.  

Marsupella bolanderi male plants with antheridia
Marsupella bolanderi male plants with spherical antheridia amongst upper leaves, CAH #1237. CC BY-NC Amanda Heinrich


Upon closer inspection, one can see that the Marsupella bolanderi antheridia are in fact attached to the plant stem via their own tiny stems. Each antheridium contains thousands of sperm, which must swim to an archegonium on a female plant to fertilize the egg. The fertilized egg then develops into the sporophyte.

Marsupella bolanderi antheridium
Marsupella bolanderi antheridium containing sperm, at 400x, CAH #1237. CC BY-NC Amanda Heinrich

Though individual Marsupella bolanderi plants are quite small, they can form fairly extensive dense patches, visible from a distance as a dark maroon turf when moist and somewhat exposed. Less exposed plants are green.

Marsupella bolanderi
Marsupella bolanderi moist in March, CAH #732. CC BY-NC Amanda Heinrich

Marsupella bolanderi is often found with an even tinier, stringier leafy liverwort growing amongst it, Cephaloziella divaricata.

Marsupella bolanderi with Cephaloziella divaricata
Marsupella bolanderi with Cephaloziella divaricata, dry in April. CC BY-NC Amanda Heinrich

In this photo, Marsupella bolanderi and Cephaloziella divaricata are seen together again, this time moist, and a little greener. Cephaloziella divaricata are the skinnier, stringier plants often lying atop the more bulbous Marsupella bolanderi plants.

Marsupella bolanderi and Cephaloziella divaricata
Marsupella bolanderi with Cephaloziella divaricata, moist in February. CC BY-NC Amanda Heinrich
Habitat and Distribution

Marsupella bolanderi can be found on soil, particularly sandy soil, as well as on sandstone outcrops or on exposed rocks.

Marsupella bolanderi habitat
Large patch of Marsupella bolanderi on sandy soil over sandstone in chaparral. CC BY-NC Amanda Heinrich

Marsupella bolanderi is said to occur from 65m-850m in elevation in California, Oregon and Washington, though a search of North American herbaria shows collections only in California.

Marsupella bolanderi distribution
Map generated by google maps using data from the Consortium of Bryophyte Herbaria of collections accessioned in herbaria.
Similar Species in Santa Barbara County: Marsupella emarginata (not yet found in Santa Barbara County) is reportedly similar but can be differentiated by its large trigones.
Name: According to Nadezhda A. Konstantinova in the online Bryophyte Flora of North Americca, the genus name Marsupella apparently refers to the brood pouch (marsupium = Latin for pouch). Mary Lincoln, author of Liverworts of New England, writes that marsupium is Latin for money bag, a reference to the mouth of the perianth which is “drawn up like a pouch.” The specific epithet bolanderi honors the German-American botanist, Henry Bolander (1831-1897).
References and Further Reading
Gymnomitriaceae, Bryophyte Flora of North America, Provisional Publication, Missouri Botanical Garden
Contributions toward a bryoflora of California III. Keys and annotated species catalogue for liverworts and hornworts, William T. Doyle and Raymond E. Stotler, Madroño, A West American Journal of Botany, Volume 53, Number 2. This issue can be ordered here here.