Reference: Biol. Bull. 205: 238 –239. (October 2003)
© 2003 Marine Biological Laboratory
Growth of a Salt Marsh Invertebrate on Several Species of Marsh Grass Detritus
A. M. Agnew
1
, D. H. Shull
1,
*, and R. Buchsbaum
2
1
Gordon College, Wenham, MA
2
Massachusetts Audubon Society, Wenham, MA
Salt marshes are important and productive ecosystems. Marsh
grasses fuel coastal ecosystem production, and marsh invertebrates
convert abundant decomposing marsh grasses into biomass avail-
able to higher trophic levels. Changes in climate, land use, nutrient
input, and introduced species potentially threaten this ecosystem,
however. An accelerated rate of sea-level rise has allowed cord
grass (Spartina alterniflora) to migrate shoreward (1). Meanwhile,
the high-marsh invasive reed Phragmites australis has expanded
seaward, reducing the extent of indigenous high-marsh grasses
such as Spartina patens and Distichlis spicata (2). Although
changes in invertebrate community structure have been observed
following Phragmites invasion (3), less is known about its effect
on ecosystem function. How would changes in the species com-
position of marsh grasses affect food supply for higher trophic
levels? Are all species of marsh grass equally nutritious for the
invertebrates that feed on them? These questions are closely re-
lated to some of the primary goals of the Plum Island Ecosystem
Long Term Ecological Research (PIE-LTER) program, which is
concerned with the processing of organic matter within the salt
marsh.
To address some of the above questions, our study investigated
the growth rates of the salt marsh amphipod Orchestia grillus, one
of the most abundant and best-studied detritivores at our study site
(4), feeding on four species of marsh grass. Two sites were studied,
Clubhead Creek and Greenwood Creek; the latter is nutrient en-
riched due to its proximity to a sewage outfall. The purpose of our
research was to examine the nutritional quality of the detritus in
terms of O. grillus growth rates and nitrogen and carbon content.
The four plant species studied were Spartina alterniflora (cord
grass), Spartina patens (marsh hay), Distichlis spicata (spike
grass), and Phragmites australis (common reed). S. alterniflora is
the dominant low-marsh species, S. patens and D. spicata are
native high-marsh species, and P. australis is a high-marsh inva-
sive.
Samples of one-year-old standing dead grasses and young new-
growth grasses were collected from each site from 2 to 6 June
2003. We sampled two differently aged grasses to assess how the
quality of detritus changes over time. Old samples were sorted, cut,
and frozen at -20 °C. Young grass samples were dried at 70 °C
for 24 h, soaked in seawater for 63 h, and then dried again at 70 °C
for 24 h before freezing, to simulate the formation of fresh detritus.
Organisms collected from the marshes were allowed to acclimate
to laboratory conditions in a large tank containing marsh wrack for
3 days. At the start of the experiment on 22 June 2003, single
organisms were weighed and placed into petri dishes containing a
single type of grass from each site. Ten replicates were designated
for each grass-species/age/site treatment. These were placed at
random over a numbered grid on a laboratory bench. Grasses were
changed about once a week, and dishes were kept wet by the
addition of filtered seawater from the marsh every few days.
Growth data for O. grillus were collected weekly by removing,
patting dry, and weighing individuals, and then returning them to
their petri dish. These data were normalized by dividing measure-
ments by the initial size of each individual. Mortality data were
collected daily, but all organisms that died before the end of the
experiment were eliminated from growth data sets. The experiment
was allowed to run for 38 days.
Within a few weeks, nearly 80% of the organisms feeding on
fresh grass were dead, and it became clear that those individuals
remaining alive in these treatments were not growing. This pattern
may have been due to the presence of high phenolic concentrations
in the fresh marsh grass detritus. Phenolic concentrations (deter-
mined by absorbance of methanol extracts at 320 nm) were sig-
nificantly higher in the fresh detritus, as demonstrated by two-
factor ANOVA with grass species and age as fixed factors
( F
[1,24]
= 331, P 0.0001), and Orchestia mortality rates were
significantly correlated with phenolic concentrations ( R
2
= 0.56,
P = 0.01). For these reasons, we focused our analysis of growth
rates on the year-old detritus.
The mean growth rates for O. grillus consuming year-old detri-
tus varied with marsh grass species. Rates were highest for am-
phipods consuming D. spicata (O. grillus increased in size by 40%
to 50%) and decreased in the order of S. alterniflora, S. patens, and
P. australis (Fig. 1A). Growth rates were determined from the
slope of organism size versus time plots by linear regression, and
rates among sites and species were compared by two-factor anal-
ysis of variance, with collection site and marsh grass species as
fixed factors. Data were log-transformed prior to analysis to cor-
rect for heteroscedasticity. No differences in growth rates between
the two sites were found ( F
[1,49]
= 0.088, P = 0.77). Variation
in the growth rates of O. grillus on different marsh grass species
was highly significant ( F
[3,49]
= 5.7, P = 0.002), with no
significant interaction ( F
[3,49]
= 0.12, P = 0.95). Post hoc
comparisons indicated that the growth rates of O. grillus on P.
australis were significantly lower than those on S. alterniflora and
D. spicata (Scheffe ´ F-test, P 0.01).
Variation in O. grillus growth rate was not correlated with the
percent carbon content of the detritus ( R
2
= 0.052, P = 0.59).
However, linear regression analysis indicated that nitrogen content
in year-old detritus accounted for approximately 70% of the vari-
ation in growth rates (Fig. 1B, F
[1,6]
= 13.7, P = 0.01).
Our results indicate that, when compared to the three native
marsh plant species studied, the invasive species, P. australis, is a
relatively poor food source for O. grillus. A diet of salt marsh * Corresponding author: dshull@gordon.edu
238 REPORTS FROM THE MBL GENERAL SCIENTIFIC MEETINGS