Epiphyllous Hepatics from Southern Alabama

Alvin R. Diamond, Jr.; Michael Woods; Hannelore Rundell

The Bryologist, Vol. 102, No. 2. (Summer, 1999), pp. 309-313.

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 The Bryologist 102(2), pp. 309-313
Copyright 0 1999 by the American Bryological and L~chenologicalSociety, Inc.

Epiphyllous Hepatics from Southern Alabama

JR., MICHAEL
ALVINR. DIAMOND, WOODS,AND HANNELORE
RUNDELL
Department of Biological Sciences, Troy State University, Troy, AL 36082, U.S.A

Abstract. Epiphyllous liverworts are reported for the jirst time from Alabama. Cololejeunea
minutissima (Sm.) Schz-. was collected on a Sabal palmetto frond at one location in south
Alabama. Cololejeunea cardiocarpa (Mont.) Steph. was collected on the leaves of a variety of taxa
in jive southern Alabama counties. Cololejeunea cardiocarpa is also reported for the jirst time as
epiphyllous on herbaceous angiosperm taxa in the continental United States. Kalmia latifolia is
reported for the second time as a host for epiphyllous liverworts.

The liverwort Cololejeunea cardiocarpa, pan-

tropical in distribution, is typically epiphyllous in
habit (Gradstein et al. 1983). In the continental The first collection of Cololejeunea cardiocarpa
United States, it occurs from Virginia south to Flor- as an epiphyll was made quite by accident. The
ida and west to Louisiana (Schuster 1980). It has sterile specimen collected 15 February, 1998 was
been reported from Geneva and Mobile Counties in attached to the adaxial leaf surface of Hexastylis
Alabama as an epicorticate species (Breil 1970; arifolia that was collected as part of a study of the
Schuster 1980). It has been previously reported as vascular flora of Ech Lake, an area within the Fort
epiphyllous from Georgia (Ellis 1971), Louisiana Rucker military reservation, in Dale County. Fur-
(Guerke 1973), and South Carolina (Davison 1997; ther searches of this area failed to locate additional
Schuster 1959). Cololejeunea minutissima has been colonies on Hexastylis. However, C. cardiocarpa
reported from Europe, Africa, and in the United was found growing on the adaxial surface of nu-
States from Tennessee and Virginia, south to Flor- merous Kalmia latifolia leaves directly overhanging
ida, and west to Texas (Schuster 1980). This species the Hexastylis. These collections were made along
has previously been reported growing as an epi- the west side of Black's Mill Creek south of Mili-
phyll from Florida (Schuster 1959, 1980) and Lou- tary Road #7 (31°23'N, 85O46'W). The canopy was
isiana (Guerke 1973). Schuster (1980) reported this composed of Fagus grandifolia, Magnolia grandi-
species from Baldwin County, ~ l a b a m agrowing flora, and Pinus glabra. The understory consisted
epicortically. This is the first report of C. cardi- of dense thickets of K. latzyolia and scattered Illici-
ocarpa and C. minutissima as epiphylls from Ala- um floridanum. Hexastylis was the most prevalent
bama. herbaceous species.
Various species of liverworts have been reported In Pike County, Cololejeunea cardiocarpa was
to occur as epiphylls in the southeastern United discovered as an epiphyll on living and dead fronds
States north of 27"301N (Table 1). These reports of Rhapidophyllum hystrix. The collection was
have come from widely scattered areas of Florida, made on a steep, north-facing, hardwood slope
Georgia, Louisiana, North Carolina, South Caroli- along Beeman Creek north of Bear Sink Creek
na, and Tennessee (Fig. 1). The majority of these Road (31°53'N, 86"11W). The canopy was com-
previous reports have been from the leaves of posed of Fagus grandifolia, Magnolia grandt$ora,
woody or semi-woody, evergreen species, including and Quercus alba. The understory consisted of
Gordonia lasianthus, Ilex cassine, 11. opaca, Kal- dense thickets of Illicium with scattered Hydrangea
mia latifolia, Leucothoe editorum, Magnolia gran- querczyolia and R. hystix. Herbaceous species in-
diflora, Persea boronia, Prunus caroliniana, Rhap- cluded Hepatica americana, Hexastylis, Polysti-
idophyllum hystix, Rhododendron maximum, Sabal chum acrostichoides, Thelypteris hexagonoptera,
palmetto, and Serenoa repens. Exceptions include and Tiarella cordifolia. The area received only par-
reports of epiphyllous liverworts on the fronds of tial sun for a portion of the day due to gaps in the
Trichomanes petersii, in Hemando County, Florida canopy caused by Hurricane Opal in 1995. There
and on a deciduous oak, Quercus nigra, in Liberty was little air movement, and the air was humid due
County, Florida (Schuster 1959). Thus, the occur- to the proximity of Beeman creek.
rence of the Cololejuenea cardiocarpa and C. min- In Henry County, Cololejeunea cardiocarpa was
utissima as epiphylls in Alabama, and the occur- collected on the adaxial leaf surface of the epi-
rence of C. cardiocarpa on the leaves of herba- phytic orchid, Epidendrum conopseum, which was
ceous angiosperm taxa, is noteworthy. growing on the trunk of Quercus lyrata located on
310 THE BRYOLOGIST [VOL. 102

TABLE1. Liverworts reported as epiphylls from the southeastern United States, north of 27"30'N.

Liverwort species Host species Location Reference

Cheilolejeunea rigidula Magnolia grandiflora Tangipahoa Parish, LA Guerke 1973
Cololejeunea biddlecomiae Rhododendron maximum Scott Co., TN Davison 1997
Cololejeunea cardiocarpa Sabal palmetto Conecuh Co., AL This paper
Cololejeunea cardiocarpa H e x a s ~ l i sarifolia, Kalmia Dale Co., AL This paper
latifolia
Cololejeunea cardiocarpa Rhapidophyllum hystrix Escambia Co., AL This paper
Pike Co., AL
Cololejeunea cardiocarpa Epidendrum conopseum Henry Co., AL This paper
Cololejeunea cardiocarpa Magnolia grandzflora, Persea Clay Co., FL Schuster 1959
borbonia, Prunus carolini-
ana, Raphidophyllum hys-
trix, Serenoa repens
Cololejeunea cardiocarpa Z1e.r cassine Lake Co., FL Schuster 1959
Cololejeunea cardiocarpa Kalmia latifolia, Quercus ni- Liberty Co., F1 Schuster 1959

Cololejeunea cardiocarpa Gordonia lasianthus, Zlex Marion Co., FL Schuster 1959

cassine, Serenoa repens,
Sabal palmetto
Cololejeunea cardiocarpa Raphidophyllum hystrix Seminole Co., FL Schuster 1959
Cololejeunea cardiocarpa Zlex opaca, Magnolia grandi- Thomas Co.. GA Ellis 1971
,flora, Persea borbonia,
Serenoa repens
Cololejeunea cardiocarpa Leucothoe editorum, Rhodo- Oconee Co., SC Schuster 1959
dendron maximum
Cololejeunea cardiocarpa Rhododendron maximum Oconee Co., SC Davison 1997
Cololejeunea minutissima Sabal palmetto Conecuh Co., AL This paper
Diplasiolejeunea rudolphi- Serenoa repens Hillsborough Co., FL Schuster 1959
ana
Diplasiolejeunea sp. Zlex cassine Lake Co., FL Schuster 1959
Drepanolejeunea appalachi- Rhododendron maximum Burke Co., NC Davison 1997
ana Oconee Co., SC
Pickens Co., SC
Drepanolejeunea bidens Leucothoe editorum, Rhodo- Oconee Co.. SC Schuster 1959
dendron maximum
Frullania asagrayana Rhododendron maximum Graham Co., NC Davison 1997
Frullania obcordata Magnolia grandijora Tangipahoa Parish, LA Guerke 1973
Frullania obcordate Leucothoe editorum, Rhodo- Oconee Co., SC Schuster 1959
dendron maximum
Frullania obcordata Rhododendron maximum Oconee Co., SC Davison 1997
Frullania sp. Rhododendron maximum Monroe Co., TN Davison 1997
Lejeunea cardoti Sabal sp. Jefferson Parish, LA Schuster 1959
Lejeunea laetevirens Magnolia grandiflora Iberia Parish, LA Guerke 1973
Lejeunea ulicina Leucothoe editorum Pickens Co., SC Davison 1997
Lejeunea ulicina Rhododendron maximum Graham Co., NC Davison 1997
Burke Co., NC
Pickens Co., SC
Leucolejeunea conchifolia Rhododendron marimum Graham Co., NC Davison 1997
Oconee Co., SC
Leucolejeunea unciloba Serenoa repens Thomas Co.. GA Ellis 1971
Metzgeria myriopoda Magnolia grandiflora Tangipahoa Parish, LA Guerke 1973
Rectoleieunea maxonii Magnolia grandiflora Tangipahoa Parish, LA Guerke 1973

the east bank of the East Fork Choctawhatchee Riv-
er south of Alabama Highway 10 (31°36'N,
85"201W). Canopy cover was complete, and the
plants received only filtered sunlight. The canopy
was composed of Carpinus caroliniana, Q. lyrata,
and Taxodium distichum. The understory was thin
due to the deep shade with Sabal minor being the
most common shrub. The herbaceous layer was
mostly absent, except for a few Carex species and
numerous bryophytes.
In Conecuh County, Cololejeunea cardiocarpa
was collected on the adaxial leaf surface of a Sabal
palmetto, which had been planted by Diamond in
1992. The palm is located on the west side of Co-
necuh County Highway 77, approximately 160 m
north of U.S. Highway 31 along the south side of
a small creek (31°26'N, 86'56'W). Growing on the
same frond were trace amounts of Cololejeunea
minutissima. The canopy consisted primarily of
Nyssa sylvatica with scattered Diospyros virgini-
 DIAMOND ET AL.: EPIPHYLLOUS HEPATICS FROM ALABAMA

FIGURE 1. Reports of epiphyllous liverworts from the southeastern United States, north of 27"301 N. Circles-
Previous literature reports; Triangles-Localities newly reported here.

ana, Pinus taeda, and Platanus occidentalis. Un- was collected on the adaxial leaf surface of Rhap-
derstory vegetation included various species of idophyllum hystrix. The Rhapidophyllum plants
ferns, rushes, and grasses. were growing on a high terrace along the east side
In Escambia County, Cololejeunea cardiocarpa of the Conecuh River about 3.2 km south of U.S.
312 THE BRYOLOGIST [VOL. 102

Highway 29 in the Conecuh National Forest TABLE2. Additional evergreen angiosperm taxa in
southern Alabama that could serve as a potential host for
(31°9'N, 86'47'W). Species in the overstory includ- epiphyllous liverworts.
ed Nyssa sylvatica, Pinus glabra, Platanus occi-
dentalis, and Quercus nigra. Herbaceous taxa in- Arundinaria gigantea
cluded Chasmanthium latifolium and Hexastyalis. Baccharis halimifolia
Bignonia capreolata
Gelsemium rankinii
Gelsemium sempervirens
Ilex coriacea
In Dale County, the Hexastylis leaf that con- Ilex glabra
Ilex myrtifolia
tained the Cololejeunea cardiocarpa colonies ap- Ilex vomitoria
peared to be at least two years old based upon color, Illicium JIoridanum
texture, and leaf deterioration. Previously, Colole- Leucothoe axillaris
jeunea had been reported as an epiphyll on Kalmia Leucothoe populifolia
Ligustrum japonica
latifolia from Liberty County, Florida (Schuster
Ligustrum sinense
1959, 1980). Davison (1997), in a study of epi- Lyonia lucida
phyllous liverworts in the Southern Appalachians, Magnolia virginiana
did not observe any taxa as epiphylls on K. latifol- Myrica cerifera
ia. Kalmia latifolia, because of its wide range of Myrica heterophylla
Myrica inodora
distribution, from New Brunswick to southern On- Osmanthus americana
tario and south to the western Florida Panhandle Persea palustris
and Louisiana (Godfrey 1988), should be consid- Pieris phillyreifolia
ered a likely host for epiphyllous liverworts at other Quercus laurifolia
Quercus hemisphaerica
sites. Quercus virginiana
In the southeastern United States, palm fronds Rhododendron minus
seem to be a particularly suitable host for epiphyl- Sabal minor
lous liverworts. In both Escarnbia and Pike Coun- Smilax bona-nox
Smilax laurifolia
ties, the liverwort was common on the adaxial sur- Smilax smallii
faces of the older (two to four years) living and
dead Rhapidophyllum fronds, and was not restricted
to the leaf tips or major drainage patterns. No liv-
erwort colonies were discovered on fronds less than be investigated as a potential host for epiphyllous
two years of age. The plants appeared healthy and liverworts in other parts of its range.
possessed both gemmae and spore capsules. Colo-
lejeunea cardiocarpa has previously been reported
as epiphyllous on the fronds of Serenoa repens in Epiphyllous liverworts are undoubtedly more
Thomas County, Georgia (Ellis 1971). The Cone- common, and occur on a wider range of host spe-
cuh County collection, from Sabal palmetto fronds, cies, than has been reported to date. Numerous oth-
contained not only C. cardiocarpa, but also trace er evergreen taxa occur in suitable habitats in
amounts of C. minutissima. Other epiphyllous liv- southern Alabama and should be investigated as
erwort species have been reported on S. palmetto potential hosts for epiphyllous liverworts (Table 2).
and S. repens from Hillsborough and Marion Coun- In particular, Cololejeunea cardiocarpa appears to
ties, Florida; S. repens in Clay County, Florida; R. be adapted to this habitat, having been found on a
hystrix in Seminole and Clay Counties Florida; and number of species, both woody and herbaceous,
on Sabal in Jefferson Parish, Louisiana (Schuster from the Appalachian mountains to sub-tropical
1959). Louisiana. The occurrence of C. cardiocarpa on
At the Henry County site, Epidendrum conop- Hexastylis may represent a rare colonization from
seum was growing laterally on the south side of the overhanging leaves of Kalmia latifolia that sup-
Quercus lyrata that leaned over the river. The Ep- ported numerous colonies. Other species of liver-
idendrum was attached to the trunk approximately worts that typically occur as epicorticates have been
2.5 m above the water surface. Although the Epi- reported as epiphyllous, apparently representing
dendrum colonies contained hundreds of leaves, fleeting colonization attempts by reproductive
only a few leaves had the liverwort growing on structures from adjacent sources. Apparently, the
them, and they were all at least two years of age main factors required for the colonization of this
as revealed by the presence of remnant flowering species are evergreen leaves, which are retained for
stalks. Epidendrum conopseum occurs from North two to many years, and a habitat that has consis-
Carolina west to Louisiana (Luer 1972). It should tently high humidity.
19991 DIAMOND ET AL.: EPIPHYLLOUS HEPATICS FROM ALABAMA
Specimens examined.-Cololejeunea cardiocarpa
(Mont.) Steph. U.S.A. ALABAMA. Conecuh Co., west side
of Conecuh County Highway 77 approximately 160 m
north of U.S. Highway 3 1 along the south side of a small
creek (31°26'N, 86"56'W), Diamond I1332 (TROY);Dale
Co., west side of Black's Mill Creek south of Military
Road #7 (31°23'N, 85"46'W), Rundell 428; Woods, Dia-
mond & Rundell 7963 (TROY);Escambia Co., high terrace
along the east side of the Conecuh River about 3.2 krn
south of U.S. Highway 29 in the Conecuh National Forest
(31°9'N, 86"47'W), Diamond I1333 (TROY);Henry Co.,
east bank of the East Fork Choctawhatchee River south
of Alabama Highway 10 (31°36'N, 85"20fW), Diamond
11309 (TROY);Pike Co., north facing, hardwood slope
along Beeman Creek north of Bear Sink Creek Road
(31°53'N, 86"11W), Diamond 11305 (TROY).-Cololejeu-
nea minutissima (Sm.) Schiffn. U.S.A. ALABAMA. Cone-
cuh Co., west side of Conecuh County Highway 77 ap-
proximately 160 m north of U.S. Highway 31 along the
south side of a small creek (31°26'N, 86"56'W), Diamond
11332A (TROY).
We wish to thank Russell Bush for preparing Figure 1
and Paul G. Davison and Raymond E. Stotler for their
critical reviews and constructive comments of this man-
uscript.
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ELLIS,E. A. 1971. Epiphyllous hepaticae in southwest
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