"A tree can be defined as a plant that, when undisturbed, develops a single, erect woody trunk. A shrub, on the other hand, is a woody plant that, when undisturbed, branches spontaneously at or below ground level to produce multiple stems." Due to its growth form, a tree generally is taller than a shrub. However,
many trees can grow as shrubs under certain environmental
conditions. An important growth characteristic of shrubs is
their ability to resprout from the base when cut or damaged.
Oaks also have this characteristic and may grow as shrubs
(sometimes called “oak grubs”) for periods of time before
turning into trees.
Fire top-kills
shrubs so that with frequent fire they do not grow to substantial
size. If fire is removed, shrubs may develop to large size,
often forming patches or clones which shade out grasses and
forbs. In the absence of fire for extended periods of time,
large shrub layers generally develop and dominate the understory.
Shrubs in oak
savannas may be either native or exotic (see Table below).
Both the native and exotic shrubs may be invasive, spreading
by underground rhizomes. Often, clonal growth can lead to
the production of very large patches which shade out all other
understory plants. Although most exotic shrubs do not have
a rhizomatous growth form, they often have other “devices”
for insuring their dominance in the understory. Buckthorn
and honeysuckle are especially noteworthy for their high degree
of invasiveness.
The early land
surveyors occasionally noted the presence of hazel (Corylus
americana) as part of the undergrowth. This shrub was
possibly singled out for comment because of its potential
economic value.
One of the major
tasks in restoring an oak savanna is the removal of the shrub
layer. Often this involves removal of exotic species such
as honeysuckle or buckthorn, but if the herbaceous groundlayer
is to be restored, removal of native shrubs is also necessary.
Brambles, which are native, are often a major headache in
recently restored savannas. Even if they are not present in
significant amount before restoration is begun, they generally
develop extensively as a result of increased light to the
forest floor after the canopy has been opened.
Some characteristics
of the shrub layer of degraded savannas are given here. Techniques
for dealing with invasive shrubs are covered separately.
Table.
Characteristics of the native and nonnative shrubs often found
in degraded oak savannas. Species with clonal growth present
special problems during the restoration process.
Latin
name |
Common
name |
Status |
Clonal |
Berberis
vulgaris |
European
barberry |
Nonative |
|
Cornus
alterniflora |
Pagoda
dogwood |
Native |
|
Cornus
racemosa |
Gray
dogwood |
Native |
Yes |
Cornus
stolonifera |
Red
osier dogwood |
Native |
Yes |
Corylus
americana |
Hazelnut |
Native |
?? |
Elaeagnus
umbellata |
Autumn olive |
Nonnative |
|
Lonicera
canadensis |
Fly
honeysuckle |
Native |
|
Lonicera
maackii |
Amur's
honeysuckle |
Nonnative |
|
Lonicera
morrowii |
Morrow's
honeysuckle |
Nonnative |
|
Lonicera
reticulata |
Grape
honeysuckle |
Native |
|
Lonicera
tatarica |
Tatarian
honeysuckle |
Nonnative |
|
Prunus
americana |
Wild
plum |
Native |
|
Prunus
pumilis |
Sand
cherry |
Native |
|
Prunus
serotina |
Black
cherry |
Native |
|
Prunus
virginiana |
Choke
cherry |
Native |
|
Rhamnus
cathartica |
Smooth
buckthorn |
Nonnative |
?? |
Rhamnus
frangula |
Glossy
buckthorn |
Nonnative |
?? |
Rhus
glabra |
Smooth
sumac |
Native |
Yes |
Rhus
hirta |
Staghorn
sumac |
Native |
Yes |
Rosa
multiflora |
Multiflora
rose |
Nonnative |
|
Rubus
sp. |
Collectively
called "brambles": Red and black raspberry; blackberry;
dewberry |
Native |
Yes |
Salix
discolor |
Pussy
willow |
Native |
|
Salix
exigua |
Sandbar
willow |
Native |
Yes |
Salix
humilis |
Prairie
willow |
Native |
|
Sambucus
canadensis |
American
elderberry |
Native |
|
Sambucus
racemosa |
Red
elderberry |
Native |
|
Viburnum
acerifolium |
Maple-leaf
viburnum |
Native |
|
Viburnum
lentago |
Nannyberry |
Native |
|
Viburnum
opulus |
Highbush
cranberry |
Native |
|
Viburnum
prunifolium |
Blackhaw |
Native |
|
Xanthoxlyum
americana |
Prickly
ash |
Native |
Yes |
|
|
Honeysuckle,
a common infestation in a degraded savanna. These bushes
are just budding out in early spring. When in full leaf,
very little light reaches the ground, so that the understory
is completely suppressed. |
|
Importance
of the Shrub Seedbank to Savanna Restoration
Most degraded
savannas suffer from extensive shrub growth over long
periods of time. One consequence is that the shrubs
have been flowering and setting seed for many years,
making major contributions to the "seed bank".
As long as the shrubs are present, the understory is
so shady that most any seeds that germinate quickly
die out. However, once the shrubs (as well as any overlying
trees) are removed, the light conditions at the ground
level are changed drastically. Now, shrub seedlings
can flourish. Whereas there may have been a few large
shrubs present before restoration, there now may be
dozens if not hundreds of shrub seedlings, each of which
has the potential to form a new plant. |
|
Buckthorn
seedlings thriving in a savanna area that had been cleared
of large buckthorn plants. The oak leaves give an idea
of scale. In a restored area such as this, very large
numbers of seedlings are possible. At the stage shown
here, use of herbicide works very well and causes very
little damage to native plants. |
|
|
The
Response of Shrubs to Savanna Restoration
As a result
of savanna restoration, the groundlayer is opened up
to sunlight. With any significant seed bank, infestation
by shrub seedlings, either native or nonnative, is virtually
assured. Brambles are a notorious problem when any woodland
is opened up by cutting. Commercial loggers simply cut
and leave, but savanna restorationists must remain to
control all the new growth. Fire top-kills but does
not eradicate. |
|
Bramble
patch (black raspberries, Rubus occidentalis) that
developed after a savanna was opened up to sunlight. The
Rubus seed bank has a notoriously long life in
the soil. |
|
Gray
dogwood (Cornus
racemosa), an attractive shrub which is a favorite
of birds and butterflies. Unfortunately, it is also fiercely
clonal and has the potential for making very large patches
which shade out all understory plants. It is best to keep
a few dogwood clones for the wildlife and eradicate all
the rest. |
|
|
Techniques
for dealing with invasive shrubs are covered separately. |
|