Ce. Bach, EFFECTS OF A SPECIALIST HERBIVORE (ALTICA-SUBPLICATA) ON SALIX-CORDATA AND SAND DUNE SUCCESSION, Ecological monographs, 64(4), 1994, pp. 423-445
Although effects of insect herbivory on host plant growth and reproduc
tion are well documented, few studies have examined how decreases in h
ost plant fitness might influence community-level patterns, such as pl
ant succession. The purpose of this study was to examine how a special
ist flea beetle, Altica subplicata, influences (1) the growth and surv
ivorship of its host plant, sand-dune willow (Salix cordata), and cons
equently, (2) patterns of plant succession on sand dunes. Growth and m
ortality of S. cordata were compared over a 3-yr period in experimenta
l plots in which host plants were either protected from beetle feeding
(with mesh cages) or exposed to beetle feeding (without cages). The c
hanges in the abundance and biomass of other plant species in these ex
perimental plots were also monitored for the 3-yr period. Growth rates
of host plants were significantly influenced by the herbivore exclusi
on treatment. Plants protected from beetle feeding added 2.2 times as
much height and 2.0 times as much diameter as did plants exposed to be
etle feeding. Differences between growth rates of plants with and with
out beetles were greater on the west dune, which had significantly gre
ater amounts of beetle damage, than on the east dune. Furthermore, wit
hin plots exposed to beetle feeding, areas with greater amounts of bee
tle damage had significantly lower height and diameter growth rates th
an did areas with lesser amounts of beetle damage. A separate experime
nt showed that mesh cages per se did not influence height growth, diam
eter growth, or number of shoots added over a 2-yr period. Plant morta
lity was also strongly affected by the herbivore exclusion treatment.
Eighty-four percent of the plants that died over the 3-yr period of th
e study were in plots with beetles present. Mortality was 3x and 6x hi
gher for plants with beetles than for plants without beetles on the ea
st and west dune, respectively. On the west dune, which had higher bee
etle damage and plant mortality, the amount of beetle damage incurred
by plants explained 88% of the variation in plant mortality. Herbivory
on S. cordata also had significant, indirect effects on the changes i
n abundance of neighboring, non-host plants. For both monocot and dico
t herbs, there were greater increases in numbers of plants in plots wi
th beetle feeding than in plots with beetles absent, but only on the w
est dune with its greater beetle damage. Numbers of herbaceous monocot
s and herbaceous dicots increased 2.5 times and 1.5 times more, respec
tively, in plots with beetles than in plots without beetles. The chang
es in abundance of other Salix plants displayed the same trends, but d
ifferences were not statistically significant. In contrast, other wood
y plants showed significantly greater decreases in abundance in plots
with beetles than in plots without beetles on the west dune. Individua
l plant species displayed both positive and negative changes in abunda
nce in response to exclusion of herbivores from S. cordata. Biomass of
two common herbaceous monocots (Juncus spp.) increased more in plots
with beetles present, but for the most common herbaceous dicots (Aster
and Solidago spp.), increases were greater in plots with beetles abse
nt. Biomass of Salix myricoides, the most common other willow, on the
east dune increased more in plots with beetles than in plots without b
eetles. In general, species that did not show differential increases i
n abundance also did not show differential growth of already-present p
lants. After 3 yr of herbivore exclusion, there were no significant di
fferences in species richness, species diversity, or species evenness
in plots with and without beetles. Thus, herbivory on S. cordata cause
d increases in the abundance of herbaceous plants and decreases in the
abundance of other woody plants, but over 3 yr did not influence over
all species diversity. It appears that selective herbivory by Altica s
ubplicata directly decreases growth and increases mortality of S. cord
ata and that damage to this single plant species also exerts an indire
ct impact on the patterns of plant succession on sand dunes.