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2012 SOUTHEASTERN NATURALIST 11(4):747–754
Plant Source Influence on Spartina alterniflora Survival
and Growth in Restored South Carolina Salt Marshes
Jennifer Beck1,2 and Danny J. Gustafson3,*
Abstract - Continued loss of coastal wetlands due to anthropogenic causes along the Atlantic
and Gulf coasts has increased the need to restore eroded or hardened structures along
the shoreline back to natural marsh systems. The source of plant material used in these
restoration plantings may have unintended consequences for the reestablishment of marsh
systems, especially if the plant material is not adapted to the local environment. In this
two-year field study, we tested the hypothesis that locally collected Spartina alterniflora
(Smooth Cordgrass) will have higher plant performance than non-local commercially
available plant material from native plant nurseries. We found that locally collected
S. alterniflora plants had higher survivorship, aboveground biomass, and cumulative stem
length than plants from non-local sources. There was a significant association between
plant performance and genetic similarity at the end of the first field season. Based on our
findings, we recommend the use of locally collected S. alterniflora from adjacent salt
marshes for small-scale salt marsh restoration projects; however, care should be taken to
not degrade the donor marsh during the process.
Salt marshes are a critical link between terrestrial and marine ecosystems
along the Atlantic and Gulf Coasts (Mendelssohn and Morris 2000). They account
for 41% of the total coastal wetlands in the conterminous United States,
providing important ecosystem functions like creating habitat for estuarine species
and nurseries for many marine species, reducing wave energy during storms,
fostering important biogeochemical processes, and allowing for soil accretion
in response to sea-level rise (Borja et al. 2010, Feagin et al. 2010, McKee et
al. 2004). The loss of salt marshes, due to anthropogenic and biological causes,
could result in the collapse of coastal ecosystems and associated species dependent
on these habitats (NOAA 1995).
Restoring the dominant perennial Spartina alterniflora Loisel. (Smooth Cordgrass)
is essential for reestablishing salt marsh ecosystem functions (Mendelssohn
and Morris 2000, Proffitt et al. 2003, Seliskar et al. 2002). Spartina alterniflora
colonization of intertidal substrate occurs through dispersal of seeds and clonal
growth (Elsey-Quirk et al. 2009, Proffitt et al. 2003). Variable seed production
in wild populations coupled with low seed set and germination rates of the only
available cultivar (Vermilion from Louisiana) can limit seed availability for large
restoration and erosion control projects; however, polycross S. alterniflora seed
shows promise (Utomo et al. 2010). Utomo et al. (2010) used 15 native Louisiana
parental lines that demonstrated superior growth and seed production to establish
1College of Charleston, MES Program, Charleston, SC 29424. 2Current address - 33 Montana
Avenue, Asheville, NC 28806. 3Department of Biology, The Citadel, Charleston, SC
29409. *Corresponding author - firstname.lastname@example.org.
748 Southeastern Naturalist Vol. 11, No. 4
seed production plots that produced genetically diverse, high-viability seed
which can be used in larger-scale marsh restoration. For smaller-scale salt marsh
restoration projects, planting of vegetative material can be a preferred method
because of the potential high survivorship rates and establishment of a marsh in
as little as four growing seasons (Craft et al. 1999, Dai and Weigert 1996, Proffitt
et al. 2003, Smart 1982).
Selection of plant material may have unintended consequences for any restoration
project if the planted material is poorly suited for environmental conditions
of the restoration site. Spartina alterniflora ecotypes have been documented with
genetic-based differences in morphology, physiology, and life history in comparison
with other conspecific populations (Daehler et al. 1999; Proffitt et al. 2003,
2005; Travis et.al 2002). Therefore, matching plant material with environmental
conditions at the target sites could increase the likelihood of a successful project
and long-term stability of the salt marsh ecosystem.
In this study, we approached small-scale salt marsh restoration from the perspective
of a natural resources manager, local or regional government official, or
any person interested in restoring S. alterniflora salt marsh. Because of the limited
seed availability and the length of time it takes to establish a salt marsh from
seed, most natural resource managers look for sources of vegetative material and
prefer those sources able to ship the plants. The hypothesis tested in this study
was that local S. alterniflora plants will have greater survivorship and growth
than plants from non-local commercially available sources. In order to determine
if differential plant performance was associated with genetic differences, we used
inter-simple sequence repeats (ISSR) genetic markers to assess genetic relationships
Common gardens were established on Kiawah Island (32°37'27"N, 80°2'29"W)
and Morgan Island (32°28'56"N, 80°29'10"W) in South Carolina. Both locations
previously supported S. alterniflora growth, but had experienced complete loss of
S. alterniflora (a.k.a. salt marsh dieback) in 1999 and 2003, respectively. The affected
dieback areas selected to establish our common gardens were 1200 m2 on Kiawah
Island and 2000 m2 on Morgan Island, with no surviving S. alterniflora ramets
within plots or seedling recruitment during the course of this study. Morgan Island is
part of the Ashepoo Combahee Edisto (ACE) Basin Reserve and is managed by the
Natural Estuarine Research Reserve System (NERRS) and the South Carolina Department
of Natural Resources. The common gardens were located approximately
28 miles apart, with similar climates, tidal inundation, and soils (Beck 2006).
Seven sources of S. alterniflora were selected to represent populations from
three distinct regions along the Gulf and Atlantic coasts. The geographic areas
represented in this study area are (1) Mid-Atlantic (New Jersey, Maryland),
(2) South Atlantic (North Carolina, South Carolina, Florida), and (3) Gulf
(Louisiana) coasts. These regions differ in key environmental conditions such as
annual temperature, winter conditions, and the duration of growing season along
2012 J. Beck and D.J. Gustafson 749
a north-to-south latitudinal gradient (Vogel et al. 2005). New England also has
extensive S. alterniflora salt marshes; however, due to the dramatically shorter
growing season and extensive ice scouring during winter (Bertness et.al 2002),
the northeastern Atlantic region sources were excluded from this study. Florida
was placed in the South Atlantic region, as opposed to the Gulf region, based on
previous research (Seneca 1994) and the fact that the Florida nursery providing
plant material was located on the Atlantic side.
Plants from the Mid-Atlantic and South Atlantic regions were purchased from
nurseries specializing in local wetland plant species. South Carolina plants were
harvested from marshes within 400 m from the common garden. The Louisiana
plant material, the only available registered S. alterniflora cultivar “Vermilion”
still in production, was obtained from the Golden Meadows Plant Material Center
in Louisiana. The “Bayshore” S. alterniflora cultivar, a registered cultivar
developed and released in 1992 from the Cape May Plant Material Center, NJ,
was from a Maryland source. “Bayshore” cultivar, however, was not kept in
production due to the consumer demand for local ecotypes over cultivated varieties
(Cape May Plant Material Center, Cape May, NJ, pers. comm.). Roots were
washed thoroughly before the plants were transplanted into gallon containers
with Pro-Mix™, and grown under greenhouse conditions from 2 to 4 weeks prior
to field planting. Prior to placing the plants in the field, 1 gram of leaf material
was collected from 18 individuals per plant source and stored at -20 °C until genomic
DNA extractions were performed.
In April 2005, S. alterniflora from each source was transplanted in a randomized
block design, with plants spaced 0.5 m apart. The number of ramets,
maximum plant height, and cumulative stem length was recorded in September
2005 and 2006. Less than 10% of the adjacent clones grew together over the 2
field seasons; however, definitive clone identification was possible by excavating
rhizomes. Individual aboveground biomass was harvested in September 2006,
washed free of debris, and dried to a constant mass.
Plant genomic DNA was extracted from approximately 0.5-g leaf material
using E.Z.N.A.® plant DNA miniprep kit (Omega Bio-Tek, Doraville, GA).
Twenty-five inter simple sequence repeats (ISSR) primers were surveyed, with 4
primers selected for this study (sequence, number of bands: (GT)6-RG, 8 bands;
(CA)8-RG, 7 bands; (GA)8-YC, 10 bands; (CT)9-G, 9 bands). ISSR polymerase
chain reaction (PCR) protocol followed that of Wolfe et al. (1998): 94 °C for 1
min 50 sec, 40 cycles of 94 °C for 40 sec, 43 °C for 45 sec, and 72 °C for 1 min
50 sec, followed by a final extension at 72 °C for 5 min. PCR profiles were visualized
in 1.5% agarose gels and stained with ethidium bormide. Images were
captured using a digital camera (Olympus C-4000 Zoom, Melville, NY) and
converted to a negative image, and fragment size was estimated based on a DNA
marker (Benchtop pGEM, #G7521, Promega, Madison, WI). Fragment sizes
were used to assign loci for each primer, and bands were scored as diallelic for
each locus (1 = band present, 0 = band absent).
750 Southeastern Naturalist Vol. 11, No. 4
Plant survivorship at the end of the first and second seasons was analyzed using
c2 test of independence between plant source (local vs. non-local) and survivorship
(alive or dead) (Gotelli and Ellison 2004). Two-way analysis of variance (ANOVA)
was used to analyze end of the second growing season plant performance (log
transformed) with site (common garden) and origin of plants (local vs. non-local;
source) main effects, and all interactions (SAS Enterprise Guide, version 4.3, SAS
Institute Inc., Cary, NC). Block effects were not significant and were removed from
all analyses presented. Genetic relationships among sources were estimated using
unweighted pair group means cluster analysis (UPGMA) cluster analysis of Jaccard’s
distances based on band frequency data. Mantel’s test using the Monte Carlo
approach with 1000 randomized runs was used to test the null hypothesis of no relationship
between genetic similarity and plant performance matrices (PC-Ord, ver.
4.2, MjM Software Design, Gleneden Beach, OR).
Local plants had significantly higher survivorship than non-local plants after
the first season (c2 = 42.2, P < 0.0001) and across the first two seasons (c 2 = 16.3,
P < 0.0001). Approximately twice as many local plants survived compared to the
non-local plants (Fig. 1).
When analyzing plant performance at the end of the second season, local
plants produced significantly greater biomass (F[1,33] = 7.9, P < 0.01) and cumulative
stem length (F[1,33] = 4.9, P = 0.04) than non-local plants (Fig. 2). There was
a significant site effect for cumulative stem length (F[1,33] = 9.2, P < 0.01), with
plants producing approximately twice the cumulative stem length at the Morgan
Island common garden (Fig. 3). Morgan Island also had significantly higher ramet
production (F[1,33] = 46.9, P < 0.0001) than plants grown at the Kiawah Island
site (Fig. 3). There were no differences in maximum plant height at the end of
the 2006 season. This means that differences in biomass and cumulative stem
length between local and non-local sources was a function of ramet production
and growth, despite no significant difference in ramet production (F[1,33] = 2.1,
P = 0.16) between local and non-local sources.
Figure 1. Local Spartina
alterniflora plants had
higher survivorship than
non-local plants over both
2012 J. Beck and D.J. Gustafson 751
Figure 2. Local Spartina alterniflora plants produced more total aboveground biomass
(left) and cumulative stem length (right) than plants from non-local sources. Bars represent
means ± 1 SE after 2 years of growth in South Carolina salt marshes.
Figure 3. Spartina alterniflora plants growing on Morgan Island produced more ramets
(left) and higher cumulative stem length (right) than plants growing on Kiawah Island.
Bars represent means ± 1 SE after two years of growth in South Carolina salt marshes.
Figure 4. Genetic relationships
plant performance (below)
of Spartina alterniflora
from local and
non-local sources grown
in South Carolina salt
marshes. Genetic relationships
populations were based
on UPGMA of Jaccard’s
distances. Plant performance
of ramets, total plant
height, and maximum
height) relative to source
populations were investigated
cluster analysis of relative
752 Southeastern Naturalist Vol. 11, No. 4
Genetic relationships among source populations follow a roughly geographic
pattern (Fig. 4). Groupings followed the Gulf coast (Florida and Louisiana), then
South Carolina and Maryland, and the North Carolina and New Jersey sources.
If the pattern of genetic relationships strictly followed source origins, then we
would have predicted Maryland to group with New Jersey. There was significant
(r = 0.69, Z = 0.02, P = 0.03) concordance between genetic relationships among
sources and the end of the 2005 season plant performance (average number of
ramets, cumulative stem length, and maximum height; Fig. 4). Inclusion of the
2005 survivorship data decoupled this relationship (r = -0.15, Z = 0.14, P = 0.32).
Restoring S. alterniflora to southeastern intertidal salt marshes can have beneficial
effects on native snail, fish, mammals and bird species (Galleher et al. 2009,
Gawlik 2002, Hotaling et al. 2010, Pung et al. 2008, Stolen et al. 2009), as well
as providing broader ecosystem functions (Leonard and Croft 2006, Seliskar et
al. 2002). The primary objective of this research was to determine if locally collected
S. alterniflora would outperform commercially available S. alterniflora in
two southeastern salt marshes. From this 2-year field study, we found that locally
collected S. alterniflora plants had higher survivorship, aboveground biomass
and cumulative stem length in 2 South Carolina salt marshes. There was a general
latitudinal pattern for genetic similarity and select plant performance measures
(average number of ramets, cumulative stem length, and maximum height) at the
end of the first field season; however, low survivorship rates of the purchased
North Carolina and Florida plants during the second season decoupled this pattern.
We did observe a consistent pattern of local plants outperforming non-local
plants in this two-season, two-common-garden field study.
In a study of early regeneration dynamics in restored and native salt marshes,
Elsey-Quirk et al. (2009) found that S. alterniflora seedling recruitment was enhanced
with open habitat at higher marsh elevations, while seedling recruitment
was facilitated in the presence of established plants at lower marsh elevations. Unlike
the strong diurnally tidal fluctuations of our common garden locations (Kiawah
Island, ≈1.8 m; Morgan Island, ≈2.4 m), water levels in the Louisiana marshes are
more influenced by wind direction than tidal flooding (Edwards and Proffitt 2003).
Mechanical stresses like tides and wind have been shown to influence clonal plant
growth (Puijalon and Bornette 2006, Puijalon et al. 2005), which could account for
an increase in S. alterniflora ramet density and cumulative stem length differences
between the 2 field sites. However, the potential effect of tidal flow velocity on
S. alterniflora morphogenesis is beyond the scope of this current research.
In conclusion, our research shows that S. alterniflora collected from adjacent
salt marshes will outperform plants purchased from non-local nurseries. This
pattern of local ecotypes outperforming non-local sources is well documented
in many species, including S. alterniflora (Daehler et al. 1999; Gustafson et al.
2005, 2008; Proffitt et al. 2003 2005; Travis et al. 2002). We recommend using
plant material collected from adjacent salt marshes for small-scale salt marsh
restoration projects; however, care should be taken to not degrade the local donor
marsh during the process.
2012 J. Beck and D.J. Gustafson 753
We would like to thank Charlie Zemp, Norm Shea, Will Chapman, Shane Kersting,
Ross Garner, and the 2005 Citadel Ecology class for help in the field. We acknowledge
the thoughtful comments and suggestions by two reviewers that improved this manuscript.
This project was made possible with grants and logistical support from the South
Carolina Department of Natural Resources, Ashepoo Combahee Edisto (ACE) Basin
NERRS Program, and The Citadel Foundation to D.J. Gustafson.
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