Targionia hypophylla L.

Targionia hypophylla is a medium-sized complex thallose liverwort with thalli 1-2 cm long and 2-5 mm wide. Its dark green thalli have purplish-black margins and tend to have rather parallel sides, giving it a strap-like appearance. The upper surface is dotted with white pores, resulting in a reptilian texture. The underside of the thallus is covered with purple-black scales, and has a tuft of white, single-celled rhizoids, which anchor the plant to the substrate. Thalli branch dichotomously and ventrally (side branches develop from underside). Targionia hypophylla is unmistakeable when fertile, with its black clam-shaped involucres at the lobe tips. It can be found on soil and less frequently on rock, typically in some shade but also exposed, often along trail cuts and roadcuts.
Scroll down for photos with labeled parts and more detailed descriptions.
Targionia hypophylla
Targionia hypophylla CC BY-NC Amanda Heinrich
Detailed Descriptions with Photos (click to enlarge)

Targionia hypophylla‘s distinctive, shiny, black clam-shaped involucres on the underside of lobe tips initially contain female reproductive structures. Following fertilization, the egg develops into the sporophyte, which remains fully contained within the involucre. The black involucres are unique to the genus, and as of 2023, Targionia hypophylla is the only species reported for California.

For a review of the liverwort life cycle click here.

Targionia hypophylla
Targionia hypophylla black clam involucres. CC BY-NC Amanda Heinrich

The upper surface of Targionia hypophylla is covered with small pores leading to photosynthetic air chambers within the thallus. The pores are raised slightly above the surface and give the plants a bumpy, somewhat reptilian look. The distinctive clam-shelled involucre can be seen peeking out from underneath the lobe tip.

Targionia hypophylla
Targionia hypophylla upper surface. CC BY-NC Amanda Heinrich

The underside of Targionia hypophylla is covered with black ventral scales. The proximal region of the lobe has a generous tuft of white, single-celled rhizoids, which anchor the plant to the substrate. The shiny black bivalved involucre can be seen at the lobe tip.

Targionia hypophylla
Targionia hypophylla lower surface. CC BY-NC Amanda Heinrich

Developing involucres of Targionia hypophylla are purplish-red.

Targionia hypophylla
Targionia hypophylla immature involucres. CC BY-NC Amanda Heinrich

This photomicrograph shows a cross-section of a Targionia hypophylla involucre containing a developing sporophyte capsule (large white sphere), which in turn contains immature spores (tiny white dots).

Targionia hypophylla
Cross-section of Targionia hypophylla involucre with immature sporophyte. CC BY-NC Amanda Heinrich

This photomicrograph shows a cross-section of a Targionia hypophylla thallus. It is quite different from those of other complex thalloid liverworts such as AsterellaCryptomitrium, and Mannia species, and can be a useful reference when attempting to identify vegetative plants.

Targionia hypophylla
Cross-section of Targionia hypophylla thallus. CC BY-NC Amanda Heinrich

Targionia hypophylla is a monoicous species, with each individual plant bearing both female and male structures. The male structures, antheridia, are on small side branches, as seen in this photo. Each bump on the side branches is an ostiole, an opening leading to an antheridium within the thallus.

Targionia hypophylla
Targionia hypophylla antheridial side branches. CC BY-NC Amanda Heinrich

This cross-section of a male Targionia hypophylla branch shows an antheridium, the male organ containing sperm, and several ostioles, the openings through which sperm will depart. Each antheridium contains thousands of sperm, which must swim through liquid water to a neighboring archegonium in order to fertilize the egg. The fertilized egg will develop into a sporophyte within the bivalved involucre and eventually bear spores.

Targionia hypophylla
Cross-section of Targionia hypophylla antheridial side branch with antheridium. CC BY-NC Amanda Heinrich

This photomicrograph shows the same Targionia hypophylla antheridium and ostiole shown above, magnified 400x so that the connection between the two can be seen.

Targionia hypophylla antheridium and ostiole
Targionia hypophylla antheridium and ostiole, 400x. CC BY-NC Amanda Heinrich

When dry, the thallus margins of Targionia hypophylla roll in, revealing the black ventral scales. The involucres are held up with mouths open wide, and, if the sporophyte capsule has dehisced, the spores are free to fly into the wind. Targionia is desiccation tolerant and will remain dormant in this dried up greasy streak state throughout the dry summer and fall months, unfurling and resuming life once the winter rains return. In this photo the brown spores can be seen spilling out of the open involucres.

Targionia hypophylla
Targionia hypophylla dry, spores spilling out. CC BY-NC Amanda Heinrich

This photo was taken at the same location as the dry plants shown above, immediately following the first rain of the season. These moistened plants have unfurled and are ready to resume growth.

Targionia hypophylla
Targionia hypophylla remoistened after first rain of season. CC BY-NC Amanda Heinrich

Targionia hypophylla spores (45-85 µm in diameter) are brown with fine areolae, which give them a “frothy” look (Doyle and Stottler). Spring-like elater cells contract and expand to give the spores an extra boost and send them on their merry way.

Targionia hypophylla spores and elater
Targionia hypophylla spore and elater, 400x. CC BY-NC Amanda Heinrich
Habitat and Distribution

Targionia hypophylla grows on soil and on rocks, typically in some shade, but also exposed, often along trail cuts and road cuts, sometimes in profusion.

Targionia hypophylla
Targionia hypophylla on soil along trail cut. CC BY-NC Amanda Heinrich

Targionia hypophylla can be found on all continents except Antarctica. In the United States it is found primarily on the west coast and in the southwest.

Map generated by google maps using data from the Consortium of Bryophyte Herbaria of collections accessioned in herbaria.

Targionia hypophylla distribution

In Santa Barbara County, Targionia hypophylla appears to be as common as Asterella californica, often growing side by side.  Targionia is significantly smaller than Asterella californica and has more parallel-sided thalli.

Targionia hypophylla and Asterella californica
Targionia hypophylla with Asterella californica. CC BY-NC Amanda Heinrich

In this close-up, the more petite, strappy Targionia hypophylla is on the right, and the more robust, flared Asterella californica is on the upper left.

Targionia hypophylla and Asterella californica
Targionia hypophylla with Asterella californica. CC BY-NC Amanda Heinrich
Name: According to calflora.net, the genus Targionia was named for Cipriano Antonio Targioni (1672-1748), a physician and scientific instrument maker from Florence. The specific epithet, “hypophylla,” is Greek for “under leaf” and presumably refers to the location of the distinctive bivalved involucre on the underside of the lobe tip.
Similar Species in Santa Barbara County: When fertile, with its black clam-shell involucres, Targionia hypophylla cannot be mistaken for any other plant. Vegetative plants may be more difficult to identify with certainty, but can be differentiated from Asterella (and Mannia and Reboulia) by the single thin layer of air-chambers as viewed in a cross-section of the thallus.
References and Further Reading
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.
Targionia hypophylla, David Long, Fred Rumsey and David Holyoak, British Bryological Society Field Guide