Family: Lophocoleaceae

Synonyms

Chiloscyphus appalachianus (R. M. Schuster) J. J. Engel & R. M Schuster

NatureServe Conservation Status

G1G2Q

Although the NatureServe rank indicates Questionable taxonomy, Lophocolea appalachiana is accepted as a good taxon in the World Checklist of Liverworts and Hornworts (Söderström et al. 2016).

Distribution

Endemic to southeastern U.S.A. Georgia, Kentucky, North Carolina, South Carolina, Tennessee. Apparently restricted to the Appalachian Plateaus and Blue Ridge physiographic provinces. As recently as 2017 the species was reported to be known from only North Carolina and Tennessee (Stotler and Crandall-Stotler 2017; Schuster 1980).

Habitat

In shaded streams within northern hardwood forests and acidic cove forests usually with Tsuga canadensis and almost always with Rhododendron maximum; moderate elevations (1180 ft – 4000 ft.).

Occurring on wet rock (solid bedrock, wet stones projecting just above water’s surface in quite streamlets, ledges and rocks in the splash zone of waterfalls, sometimes on thin layer of soil over wet rocks) and to a lesser extent on wet woody debris (logs, limbs, and exposed roots at the water’s edge). Often in deep shade. It appears to disperse locally by spores; mats may persist in place for a few years before succession or local perturbations remove them. Though not found in the spray cliffs of the Chattooga Basin (Zartman and Pittillo 1998), Lophocolea appalachiana is a component of spray cliff communities.

Associated bryophytes include: Liverworts: Conocephalum salebrosum, Jubula pennsylvanica, Jungermannia evansii, Pellia appalachiana, Plagiochila porelloides, Riccardia multifida, Scapania undulata; Hornwort: Nothoceros aenigmaticus; and Mosses: Amblystegium tenax, Hookeria acutifolia, Hygohypnum eugyrium, Platylomella lescurii.

The southernmost known occurrence is from Amicalola Mountain in Dawson County, Georgia, occurring with Nothoceros aenigmaticus in a streamlet surrounded by a Kalmia latifolia understory in a mixed hardwood forest.

Brief Description and Tips for Identification

Plants green, forming loose mats. Individual shoots ca. 2 – 2.5 mm wide. Lateral leaves succubously arranged, apex shallowly bilobed to entire. Underleaves bilobed with a deep, U-shaped to V-shaped sinus, each lateral base of the underleaf with or without a lateral tooth; decurrent base of underleaf on one side of the stem often fused with base of a lateral leaf.

Monoicous. Androecia at apices of elongate leafy shoots. Male bracts smaller than lateral vegetative leaves, bract apices rounded, unlobed or rarely a few bracts feebly bilobed. Perianth at apex of short lateral branch or at apex of elongate leafy shoot. Perianths trigonous (with three flat sides), keels of perianth typically winged, the wings often coarsely toothed.

While data on timing of gametangia and spore production is mostly lacking, mature capsules were present in April (Tennessee) and young female shoots (archegoniate) present in July and August (North Carolina, Georgia). Presumably, sporophytes are conceived in mid- to late summer and mature the following spring.

Plants usually possess sexual branches (even if only from past seasons). Androecia offer the one, most reliable character for confident identification. Within the androecium the male bracts are unlobed with rounded apices. The apices of male bracts in the rather similar Lophocolea coadunata (=L. cuspidata) are bilobed. As both species are polymorphous (Schuster 1980), there are few absolute character distinctions between L. appalachiana and L. coadunata (=L. cuspidata). Other than the apices of male bracts, distinctions are subtle. In both species one side of the underleaf base may be connate with base of a lateral leaf; however, this condition is more frequent in L. appalachiana. There is a difference in the degree to which lateral leaf apices are lobed. Those of L. coadunata are usually sharply bilobed, those of L. appalachiana are shallowly bilobed, feebly bilobed with one or two projecting tooth-like lobe apices, or entire.

Androecia also distinguish Lophocolea appalachiana from Chiloscyphus. Regional Chiloscyphus possess lateral leaves whose apices are entire, retuse, or rarely a few leaves are bilobed. Such variation may lead one to consider L. appalachiana for specimens of Chiloscyphus; however, androecial bracts of Chiloscyphus are similar in size to vegetative leaves and perianths are not trigonous and are always born on very short lateral branches.

Salient Features

  • Male bracts much smaller that vegetative lateral leaves
  • Apices of male bracts entire, rounded
  • Lateral leaves shallowly bilobed to entire
  • High frequency of underleaf bases connate with base of lateral leaves

References

Schuster, R.M. 1980. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian. Volume IV. Columbia University Press, New York

Söderström, L., Hagborg, A., von Konrat, M., Bartholomew-Began, S., Bell, D., Briscoe, L., ... & Cooper, E. D. 2016. World checklist of hornworts and liverworts. PhytoKeys, 59(1–2): 1–828.

Stotler, R. E., and Crandall-Stotler, B. 2017. A synopsis of the liverwort flora of North America north of Mexico. Annals of the Missouri Botanical Garden, 102(4), 574-709.

Zartman, C. E., and Pittillo, J. D. 1998. Spray cliff communities of the Chattooga Basin. Castanea, 217-240.

Acknowledgment

Some text and images on this page were originally prepared for the Georgia Department of Natural Resources in 2010, contract number 605-090427 with Paul G. Davison and used here with permission.


Habitat

moss

Lophocolea appalachiana

Silver tweezers are above the mat at the left arrow. The bright colored mats near the right arrow are Plagiochila porelloides with adjacent Lophocolea appalachiana. At this scale the mats of L. appalachiana blend in with the dark green of other bryophytes on the wet rock.

liverwort

Lophocolea appalachiana

Under deep shade of Rhododendron maximum in the splash zone of a waterfall.

Habit

liverwort

Lophocolea appalachiana

Leaves are shallowly bilobed. Occuring with Hygrohypnum eugyrium, Platylomella lescurii, Plagiochila porelloides.

liverwort

Lophocolea appalachiana

An androecium is just beginning to form at the apex of the lowest, elongate shoot in the photo

liverwort

Lophocolea appalachiana

The rounded apices of male bracts at red arrows; the black arrows at lower right indicate toothed wings on the keels of the perianth.

Habit

liverwort

Lophocolea appalachiana

Androecia with bract apices rounded, entire. In the lowest inset one shoot lies atop another but appear as one at first glance. Archegonia were found after dissection of the indicated female shoot tip.

Morphology

liverwort

Lophocolea appalachiana

Variation in leaf apices from a single collection.

liverwort

Lophocolea appalachiana

Arrows indicate the narrow decurrent strip of underleaf base that fuses with the base of a lateral leaf.

liverwort

Lophocolea appalachiana

Underleaves whose bases do not fuse with base of lateral leaves.

L. appalachiana vs. L. coadunata (=L. cuspidata)

liverwort

Lophocolea appalachiana vs. Lophocolea coadunata

Compare the apices of vegetative leaves and apices of male bracts between the two species.

Lophocolea coadunata

liverwort

Lophocolea coadunata

The saccate bases of male bracts each conceal a single antheridium.

liverwort

Lophocolea coadunata

The teeth at the base of the underleaves point towards the shoot apex rather than pointing almost 90 degrees away from the stem as in L. appalachiana.

liverwort

Lophocolea coadunata

Lower left inset with a clump of rhizoids and obscuring debris. Rhizoids in the genus Lophocolea arise from the base of underleaves and adjacent stem sector.

Lophocolea coadunata

liverwort

Lophocolea coadunata

The lobes of lateral leaves are longer than those of L. appalachiana.