New York Scalewort
Frullania eboracensis
Hollywort family (Jubulaceae)
Description:
This leafy liverwort forms a low spreading mat of irregularly branched
leafy stems; it is herbaceous, evergreen, and perennial. The spreading
mat of leafy stems clings tightly to the underlying surface, where
it usually spans 0.5–10 cm. across (rarely more). Old leafy stems have
a tendency to wither away as new leafy stems develop; individual leafy
stems are only 0.8–1 mm. across. The hairless stems and leaves
are dark green or reddish green during the summer, but they become
dark red or reddish brown during the winter. Individual stems are only
0.1 mm. across and more or less terete. Along these stems, there are
alternate leaves that partially overlap each other. The upper margin of
each leaf (facing the terminal point of the stem) partially overlaps
the lower margin of the leaf that succeeds it along the stem (incubous
insertion). Like the stems, the alternate leaves lie flat along the
underlying surface. Individual leaves are bilobed-conduplicate in
structure, consisting of an upper (dorsal) lobe and a lower (ventral)
lobe. The larger upper lobes lie directly above their smaller ventral
lobes. Individual upper lobes are about 0.4 mm. long and 0.45 mm.
across; they are suborbicular in shape, toothless along their margins,
and and slightly indented (cordate) at their bases.
Individual lower lobes
are about 0.2 mm. long and 0.2 mm. across; they are folded
into helmet-shaped pockets that are somewhat flattened. The
rounded sides of each helmet-shaped pocket are folded and closed, while
its truncate bottom is open. Each pair of upper and lower lobes are
connected to each other by a short narrow stalk. Pairs of alternate
leaves are accompanied by solitary underleaves that are appressed
against the lower sides of the stems. These underleaves are about 0.2
mm. long and 0.15 mm. across; they are divided into 2 lanceolate lobes.
Each pair of these lobes extends to about one-third of the length of an
underleaf; the lateral margins of the underleaf are toothless or nearly
so. Directly across from the base of each underleaf, there is a tiny
spike, only 2-3 cells across at its base, that is linear in shape. New
York Scalewort is dioecious, forming male and female
reproductive organs on different plants. Male plants produce their
reproductive organs on short lateral branches that have several pairs
of leaf-like bracts. These bracts resemble the leaves, except they are
somewhat smaller in size. Female plants produce their reproductive
organs at the tips of stems. The female reproductive organs are
contained in bud-like perianths (one perianth per stem tip); preceding
and partly surrounding the base of each perianth, there are about 3
pairs of bracts.
Each perianth is about 1 mm. long, obovoid in
shape, 3-angled, and erect; it terminates abruptly in a short
slender beak. The perianth is keeled along its angles; these keels are
relatively smooth or only slightly roughened. The bracts preceding the
perianth resemble highly modified leaves; they are up to 0.8 mm.
long, 0.45 mm. across, and unlobed to irregularly lobed. After
fertilization occurs, spore-bearing capsules on stalks develop
during the
spring for about 2 weeks. The stalks are white, terete, translucent,
hairless, and
ascending to erect, while the small capsules are broadly ovoid to
globoid,
brown to black, and hairless. At maturity, individual capsules divide
into 4 parts, releasing their spores to the wind. Fibrous rhizoids
develop from the axils of the underleaves along each leafy stem; they
anchor the plant to the underlying surface. This leafy liverwort can
reproduce asexually by forming gemmae (miniature buds) along the
margins of its leaves. New plants can develop when the gemmae become
detached from the leaves.
Cultivation:
The preference is partial
sun to medium shade and moist to dry-mesic conditions. This leafy
liverwort normally colonizes tree bark. It has greater tolerance to
sunlight and dry heat than most leafy liverworts, but it is intolerant
of competition from other plants.
Range
& Habitat: The
native New York Scalewort (Frullania eboracensis) is
occasional in most areas of Illinois (see Distribution
Map).
This leafy liverwort undoubtedly occurs in
additional counties than what has been indicated, but it is easy to
overlook because of its small size. New York Scalewort is widely
distributed in eastern North America and adjacent areas of southern
Canada. Habitats include bark of trees on rocky bluffs, bark of trees
along wooded slopes, bark of trees in upland and lowland woodlands,
bark of trees along streams and ponds, bark of trees in
swamps, exposed roots of trees in woodlands, logs of fallen trees in
woodlands, bark of trees in cemeteries, and planks of wooden fences in
damp shaded places. This leafy liverwort has been found primarily on
the bark of deciduous trees, including oak, maple, ash, elm, hackberry,
cottonwood, beech, and musclewood. It has also been found on the bark
of Eastern Red Cedar (Juniperus virginiana).
Usually, it is found on
the trunks of trees, rather than branches or exposed roots. Older
mature trees are preferred over young trees. This leafy liverwort
prefers natural areas with trees, although sometimes it is found in
more developed areas where mature trees occur.
Faunal
Associations: Rotifers sometimes inhabit the
helmet-shaped lower lobes
of leaves. It is unclear if this causes any benefit or harm to the
plant (Puterbaugh et al., 2004). This leafy liverwort is used to a
minor extent in the construction of nests by Prothonotary Warblers
(Blem & Blem, 1994).
Photographic
Location: A fallen log
near a pool of water in Busey Woods at Urbana, Illinois. Close-up
photographs were taken indoors using a microscope.
Comments:
New York Scalewort (Frullania eboracensis) is the
most
common scalewort
(Frullania sp.) in Illinois; the remaining species
of scalewort are
uncommon to rare within the state. The common name derives from an old
Latin word (Eboracum) for York, England. But this name actually refers
to the state of New York, where this leafy liverwort was first
collected. Another name that could be applied to this plant is Common
Scalewort. Sometimes species in this genus are assigned to their own
plant family, Frullaniaceae. The structure of the alternate leaves for
this group of plants is unusual. The helmet-shaped lower lobes of these
leaves have the capacity to store water, enabling these miniature
plants to withstand dry spells for a longer period of time. New York
Scalewort can be distinguished from other species of scalewort within
the state by its small size (leafy shoots only 0.8–1 mm. across), lower
leaf lobes that are about as long as they are across, lower leaf lobes
that are about one-third to one-half the size of upper leaf lobes,
underleaves (0.15 mm. across) that are only a little wider than their
stems (0.10 mm. across), underleaves that have toothless or nearly
toothless lateral margins, perianths that have smooth to slightly
roughened keels along their sides (rather than tuberculate keels), and
leaf cell walls that are swollen
at intervals.