Wetlands
DOI 10.1007/s13157-013-0504-z
ARTICLE
Ecology of Native vs. Introduced Phragmites australis
(Common Reed) in Chicago-Area Wetlands
Amy L. Price & Jeremie B. Fant & Daniel J. Larkin
Received: 8 May 2013 / Accepted: 1 December 2013
# Society of Wetland Scientists 2013
Abstract Rapid spread of Phragmites australis (common reed)
in North American wetlands is widely attributed to cryptic
invasion by an introduced lineage. However, in the
Midwestern U.S., the native subspecies (subsp. americanus)
may also exhibit rapid expansion. Where both lineages occur,
wetland managers are sometimes unsure whether they should
limit management activities to the introduced lineage or control
both. We conducted field studies to contrast the ecology of
native and introduced Phragmites by pairing patches of each
with native reference vegetation. We measured each lineage’s
association with environmental conditions, their growth metrics
(stem heights, stem densities, and plant cover), and their inva-
siveness as indicated by the diversity and composition of asso-
ciated plant communities. Introduced Phragmites exhibited
more robust growth than the native, and its growth was more
positively correlated with increases in soil nutrient availability
and salinity. Areas with introduced Phragmites had lower plant
diversity and altered species composition relative to reference
vegetation. We did not observe similar evidence of invasiveness
in native Phragmites. We encourage wetland managers to dif-
ferentiate populations by lineage and, unless there is compelling
evidence to do otherwise, restrict control efforts to the intro-
duced lineage.
Keywords Invasive species . Eutrophication . Cryptic
invasion . Midwest
A. L. Price (*) : J. B. Fant : D. J. Larkin
Plant Science and Conservation, Chicago Botanic Garden,
1000 Lake Cook Road, Glencoe, IL 60022, USA
e-mail: amy.l.price@gmail.com
A. L. Price
Graduate Program in Plant Biology and Conservation, Northwestern
University, 2205 Tech Drive, O.T. Hogan Hall, Room 2-144,
Evanston, IL 60208, USA
Introduction
Phragmites australis (Cav.) Trin. ex Steud. (common reed,
hereafter Phragmites), once a minor component of North
American wetlands, began aggressively expanding in salt
marshes along the Atlantic coast over a century ago, and has
since spread from tidal to brackish and freshwater wetlands
(Chambers et al. 1999; Saltonstall 2002). Phragmites invasion
produces tall, dense, monotypic stands that reduce
microtopographic heterogeneity, displace native vegetation, and
negatively affect food webs and wildlife (Marks et al. 1994;
Chambers et al. 1999; Able and Ragan 2003; Gratton and Denno
2006). Although Phragmites is native to North America, accord-
ing to the cryptic invasion hypothesis, the introduction of
Eurasian Phragmites genotypes has driven a rapid expansion
of the species since the early 20th century (Saltonstall 2002). The
native lineage, now recognized as the distinct subspecies
P. australis subsp. americanus Saltonstall, Peterson and Soreng
(Saltonstall et al. 2004), has become rare along the Atlantic coast,
having been largely displaced by the introduced lineage, but
persists in the Midwest (Saltonstall 2002).
The spread of Phragmites may be explained by a number of
non-mutually exclusive hypotheses that have been put forth to
explain invasive plant success. Two of these hypotheses are
the “eutrophication hypothesis,” positing that excess nutrients
can promote plant invasions by stimulating plant growth
(Hobbs and Huenneke 1992; Davis et al. 2000), and the
related, “efficient-use hypothesis,” which proposes that cer-
tain species are better able to capitalize on available resources
than others (Zedler and Kercher 2004). Evidence in support of
the eutrophication hypothesis includes greater occurrence of
Phragmites in areas where anthropogenic activities have in-
creased nutrient availability, e.g. agricultural and developed
areas (Bertness et al. 2002; Silliman and Bertness 2004;
Chambers et al. 2008; Jodoin et al. 2008; Taddeo and de
Blois 2012). Experimental work has also demonstrated that

elevated nitrogen increases Phragmites density, height, bio-
mass, allocation to above- vs. below-ground growth, and
sexual reproduction (Minchinton and Bertness 2003; Rickey
and Anderson 2004; Kettenring et al. 2011). In support of the
efficient-use hypothesis, direct comparisons of introduced and
native Phragmites have shown that, under the same growing
conditions, the introduced lineage generally produces more
biomass than the native and is more responsive to increases in
nutrient availability (League et al. 2006; Saltonstall and
Stevenson 2007; Holdredge et al. 2010).
In addition to being better able to exploit resources, inva-
sive plants may also have broader tolerance of disturbance or
extreme environmental conditions, according to the “broader-
tolerance hypothesis” (Zedler and Kercher 2004). In particu-
lar, a broad range of tolerance to salinity may be a factor in
Phragmites invasion (Matoh et al. 1988; Wijte and Gallagher
1996). In inland systems, salinity inputs from road salt may
promote Phragmites spread (Lelong et al. 2007; Jodoin et al.
2008). The introduced lineage has been shown to produce more
shoots and grow faster at salinity levels above freshwater than
the native (Vasquez et al. 2005). Phragmites’ tolerance of a
broad range of hydrologic conditions and disturbances may also
favor its spread (Burdick and Konisky 2003; Hudon 2004).
Problematic invasive species are not always non-native.
According to the “native-invader hypothesis,” native species
can also become hyper-abundant by taking advantage of chang-
ing environmental conditions (Garrott et al. 1993; Simberloff
2011). Some of the most problematic invasive species in North
American wetlands have native genetic components, including
Typha × glauca(hybrid cattail), which has a native parent species,
and Phalaris arundinacea (reed canarygrass), which has a
circumglobal distribution with both native and non-native geno-
types in North America (Galatowitsch et al. 1999; Casler et al.
2009). Documented rapid expanses of Phragmites are not limited
to the introduced lineage; the native subspecies has also been
found to quickly spread under disturbed conditions (Lynch and
Saltonstall 2002; Weaver et al. 2010). Given the high impacts of
anthropogenic disturbances on temperate wetlands, native
Phragmites might function as a “native invader” in some systems
(sensu Simberloff 2011), exerting ecological harm irrespective of
its geographic origins.
While Phragmites invasion has been the subject of a tremen-
dous amount of research, most North American studies have
focused on the Atlantic and Gulf coasts (Meyerson et al. 2012),
areas where Phragmites populations overwhelmingly comprise
the introduced lineage (Saltonstall 2002). In contrast, native
populations persist in the upper Midwest (Saltonstall 2002),
where Phragmites has spread rapidly over the past few decades
(Larkin 2012), but where relatively little research has been
conducted (Meyerson et al. 2012). In the Chicago region, and
elsewhere where both lineages occur, this has led to uncertainty
among wetland managers regarding whether Phragmites should
be controlled in general or if the subspecies should be
Wetlands
differentiated and control efforts limited to the introduced line-
age (Hunt and Larkin, unpub. data). Clarification of this ques-
tion could help managers better allocate limited resources, and
has implications for the future of the native subspecies, which
has declined throughout much of its range (Saltonstall 2011).
We conducted field studies to compare the ecology of native
vs. introduced populations of Phragmites in the Chicago region,
contrasting the lineages’ growth metrics (height, stem density,
and cover), correlations with environmental conditions, and
associated plant communities. We expected an interaction be-
tween genotype and local environmental conditions to deter-
mine phenotype, and phenotype to determine plant-community
impacts. We hypothesized that: 1) Phragmites, and in particular
the introduced lineage, would occur in areas with higher nutrient
concentrations than reference vegetation that lacked Phragmites
(eutrophication hypothesis); 2) introduced Phragmites would
show more responsiveness in terms of growth than native
Phragmites as nutrient availability increased (efficient-use
hypothesis); 3) introduced Phragmites would occur across a
broader range of soil moisture conditions and salinities
(broader-tolerance hypothesis); and 4) plant communities occu-
pied by Phragmites of either lineage would be less diverse than,
and distinct in composition from, areas lacking Phragmites. For
this last hypothesis, we expected this effect to be stronger in
introduced Phragmites—reflecting its more robust growth and
responsiveness to resource availability—but to also be observ-
able in the native subspecies (native-invader hypothesis).
Materials and Methods
Site Selection and Identification of Lineage
In spring 2010, we consulted wetland managers and made
field visits to identify Phragmites-containing Lake Michigan
coastal and interior wetlands for study in Lake, Cook, Kane,
and DuPage counties in Illinois and in Lake and Porter
counties in Indiana (Fig. 1). We haphazardly sampled 30
Phragmites leaves for genetic analyses, including differentiation
of native and introduced lineages, from each of 20 Phragmites
patches identified. Leaves were stored in a freezer at −20 °C until
DNA could be extracted using Qiagen DNeasy Kits (Qiagen
Inc., Valencia, CA, USA). A subset of six leaf samples from each
patch were analyzed using the PCR-RFLP technique for
Phragmites chloroplasts (Saltonstall 2003), with the remaining
samples reserved for future analyses. We found that 11 patches
were the introduced lineage, 9 were the native, and no patches
contained both subspecies (though patches of both types often
co-occurred within the same wetlands or wetland complexes).
From these 20 sites, we selected a subset of 6 sites (3 native,
3 introduced) on which to perform intensive sampling of plant-
community and environmental variables (Fig. 1). Intensively
sampled sites were limited to natural wetlands that had not
Wetlands
Fig. 1 Map of study sites sampled in northeastern Illinois and northwestern Indiana, USA
undergone Phragmites management and that had fairly large
patches of well-established Phragmites (minimum patch size of
~250 m2 and with relatively dense growth of Phragmites). For
five of the six intensive sites, Phragmites patches were paired
with adjacent uninvaded patches of wetland, not colonized by
Phragmites or any other invasive plant species, as reference
vegetation. Suitable reference vegetation was not present in one
site as an area that appeared uninvaded had dense growth of
other invasive species by late summer.
Environmental Data
Environmental data were collected from intensively sampled
sites in August and September 2010. Within each Phragmites
patch and uninvaded area, we randomly selected 6 of the 30
quadrats used for vegetation sampling (see below). From each
of these, we took one 20-cm deep soil core for nutrient and
salinity analyses, collected all litter from a 0.25-m2 subplot,
and measured water depth.
Litter mass of subplots was determined following drying in
a 44 °C herbarium drier for ≥ 24 h. Soil cores were refrigerated
until the following day and then homogenized, after which
nutrients were extracted from a 5 g subsample in 2 M KCl
using standard methods (APHA 2005). We determined
Site type
IntroducedPhragmites
NativePhragmites
Intensively sampled
concentrations of ammonium (NH4), nitrate/nitrite (NOx),
and orthophosphate (PO4) using a SEAL AQ2+ discrete ana-
lyzer (SEAL Analytical, Mequon, WI, USA). We determined
gravimetric soil moisture by oven-drying 10 g of soil for ≥
24 h at 105 °C. Soil salinity was estimated using a Eutech
Instruments electrical conductivity meter; 50 mL of distilled
water was added to 10 g of dry soil, soil solutions were shaken
for 2 h, and solutions were allowed to settle for 0.5 h.
Measures of conductivity (μ S cm−1) were converted to salin-
ity (ppm) as per APHA (2005).
Phragmites Growth Metrics and Plant-Community
Composition
In July and August 2010, we collected data on Phragmites’
growth metrics at all sites and collected vegetation data from
intensively sampled sites. At sites that were not intensively
sampled, maximum height and density of live Phragmites
stems were recorded in six 1-m2 quadrats haphazardly placed
within each patch of Phragmites, several meters away from
the patch edge. In intensively sampled sites, 30 1-m2 quadrats
spaced evenly along transects were placed within each
Phragmites patch and within each paired uninvaded area.
Transects were of varying length and number per site,

reflecting the variation in patch dimensions, ranging from 3 to
5 transects, each 30 to 50 m in length. For each quadrat, we
recorded Phragmites stem density and maximum height, the
height of the four tallest plants of any species, and the cover of
each species using an arcsine-square root cover class system
(McCune and Grace 2002).
Analyses
For the intensively sampled sites, we grouped data from vegeta-
tion plots with their nearest environmental plot and calculated
means, yielding 6 samples of Phragmites-growth metrics, vege-
tation, and environmental data for each patch. For each of these
six data points, we calculated the abundance of each species as
the sum of its relative frequency and relative cover, plant species
richness and Shannon’s index (H′) (excluding Phragmites itself),
mean canopy height, and Phragmites maximum height and stem
density where applicable. As an overall indicator of Phragmites
growth, we calculated an importance value for Phragmites
(see Curtis and McIntosh 1951; Mozdzer et al. 2008) by taking
the mean of relative cover, relative stem density, and relative
height and multiplying the result by 100 to convert to a percent
scale. Possible values of this index range from 0 %
(no Phragmites) to 100 % (if the highest values of all three
components of the index were found in a single location).
We compared environmental and plant-community measures
among intensively sampled native-Phragmites, introduced-
Phragmites, and uninvaded patches using linear mixed-effects
(LME) models (Pinheiro and Bates 2000) to account for our
nested sampling design (areas with and without Phragmites
were paired within the same sites). For variables with significant
main effects of plant community type, we performed a priori
contrasts (Crawley 2005) to test for significant differences be-
tween areas with and without Phragmites and then between
areas with native vs. introduced Phragmites.
We used ANOVA to test for differences in growth metrics
between native and introduced Phragmites. Comparisons of
Phragmites stem density and height were made using data
from all 20 sites. Tests for differences in Phragmites cover and
importance value were made using data from the six inten-
sively sampled sites. We used regressions to test whether
Phragmites importance values were correlated with environ-
mental measures, and whether plant diversity differed with
Phragmites importance. Prior to ANOVA, LME, and regres-
sion analyses, continuous variables were ln-transformed to
better meet assumptions of normality and homoscedasticity.
We characterized plant-community composition using
non-metric multidimensional scaling (NMDS) ordination.
Phragmites itself was removed from community data so that
it could be treated as a potential explanatory variable. Species
occurring in fewer than 5 % of plots were also excluded to
control against disproportionate effects of rare taxa (McCune
and Grace 2002). Data were relativized by sample and species
Wetlands
maxima and the ordination was based on Bray-Curtis dissim-
ilarity using three axes.
We tested whether environmental factors or Phragmites
importance values were correlated with community composi-
tion using a permutation test (10,000 permutations) and over-
laid significant vectors on the ordination to show the direction
and magnitude of correlations. Differences in community
composition by plant community type were tested using per-
mutational multivariate analysis of variance (PERMANOVA,
a multivariate analog of ANOVA; Anderson 2001), which was
performed using Bray-Curtis dissimilarities, with P-values
calculated by a Monte-Carlo procedure with 10,000 permuta-
tions. Because one patch of introduced Phragmites was not
paired with an uninvaded area, this could have exaggerated
differences in composition associated with introduced
Phragmites, so we repeated tests with the unpaired patch
removed. Having found significant differences in composition
by plant community type, we used similarity percentage
(SIMPER) analysis to evaluate the contributions of each plant
species to Bray-Curtis dissimilarity between groups (Clarke
and Warwick 2001).
All analyses were performed in R 2.15.3 (R Core Team
2013), with the vegan and nlme packages used for multivar-
iate and LME analyses, respectively (Oksanen et al. 2013;
Pinheiro et al. 2013). Significance was assessed as α<0.05,
but marginal differences (P<0.10) are noted.
Results
Areas of different plant community type significantly differed
in concentrations of soil NH4, NOx, and PO4; marginally
differed in soil moisture; and did not differ in salinity
(Table 1). In general, values of these properties were highest
in areas with native Phragmites and lowest in areas with
introduced Phragmites.
Introduced Phragmites had more robust growth than native
Phragmites, growing taller, more densely, and with higher cover
(Table 1). Importance values of native Phragmites were nega-
tively correlated with soil NOx, positively correlated with PO4,
and unrelated to soil moisture, salinity, or NH4 (Fig. 2).
Importance values for introduced Phragmites increased with soil
moisture, salinity, and PO4; marginally increased with NH4; and
were unrelated to NOx (Fig. 2). Areas with Phragmites of either
subspecies had taller vegetation than uninvaded areas and intro-
duced Phragmites grew taller than native Phragmites (Table 1).
Litter mass was lowest in uninvaded areas and highest in areas
with native Phragmites (Table 1).
Plant species richness and H′ differed by plant community
type (Table 1). Both were lowest in areas with introduced
Phragmites and similar between areas that were uninvaded
and had native Phragmites (inclusion of Phragmites itself would
have resulted in areas with native Phragmites having the highest
Wetlands

Table 1 Summary data and statistical comparisons for areas differing in plant community type

Plant community type (mean ± s.e.) Main effects Contrasts (P-values)

Variables None Native Introduced df F P None vs. Native vs.

either lineage introduced

Environmental factors
Gravimetric soil moisture (%) 62.9±2.0 65.4±2.0 55.6±2.5 2, 57 2.4 0.097 − −
Salinity (ppm) 467±44 531±50 355±33 2, 57 2.2 0.12 − −
NH4 (μg g−1 dry soil) 11.3±0.7 11.7±0.8 8.7±0.5 2, 57 3.8 0.028 0.17 0.014
NOx (μg g−1 dry soil) 1.4±0.3 3.7±0.9 0.7±0.1 2, 57 6.3 0.003 0.11 0.004
PO4 (μg g−1 dry soil) 0.5±0.1 0.9±0.1 0.92±0.3 2, 57 6.8 0.002 0.002 0.27
Phragmites growth
Maximum height (cm) − 246±6.2 306±8.3 1, 117 29.3 <0.0001 − −
Stem density (stems m−2) − 22.3±1.8 31.9±2.1 1, 117 9.7 0.002 − −
Cover (%) − 14.6±0.9 98.4±0.9 1, 33 70.7 <0.0001 − −
Importance value (%) − 37.4±2.6 63.9±4.2 1, 33 27.0 <0.0001 − −
Vegetation metrics
Litter mass (g m−2) 14.4±2.4 29.4±4.2 20.9±4.5 2, 57 6.7 0.002 0.001 0.59
Canopy height (cm) 162±7 214±7 258±18 2, 57 23.1 <0.0001 <0.0001 0.18
Species richness (per m2) 7.1±0.3 6.9±0.6 3.9±0.5 2, 57 7.4 0.001 0.001 0.031
Shannon’s index (per m2) 1.9±0.05 1.8±0.1 1.2±0.13 2, 57 6.1 0.004 0.005 0.028

(bold text indicates significant differences)

diversity). There was no relationship between importance of Discussion

native Phragmites and H′, but there was a strong, negative
relationship in the case of introduced Phragmites (Fig. 3).
NMDS ordination and PERMANOVA showed significant
differences in plant community composition by plant commu-
nity type (Fig. 4). Areas with no Phragmites and native
Phragmites only marginally differed in pairwise comparisons
of composition, but areas with introduced Phragmites showed
highly significant differences with both other groups. These
patterns were unchanged when one introduced Phragmites
patch not paired with an uninvaded area was removed from
the analysis. Plant-community composition was most strongly
correlated with Phragmites importance; vectors for soil mois-
ture, nutrients, and salinity were also significant.
Community dissimilarity of areas with introduced
Phragmites, relative to areas with native or no Phragmites,
was driven by: 1) lower abundances of Carex stricta (tussock
sedge), Calamagrostis canadensis (bluejoint grass), Euthamia
graminifolia (grass-leaved goldenrod), Polygonum amphibium
var. stipulaceum (water smartweed), Schoenoplectus acutus
(hardstem bulrush), and Typha spp. (cattails); and 2) higher
abundances of Onoclea sensibilis (sensitive fern), Impatiens
capensis (jewelweed), Phalaris arundinacea (reed canarygrass),
Lysimachia thyrsiflora (tufted loosestrife), and Pilea pumila
(Canada clearweed). Collectively, these 11 species (10 % of
all species identified) accounted for 50 % of the total
Bray-Curtis dissimilarity between areas with and with-
out introduced Phragmites.
Despite being closely related, native and introduced
Phragmites differed in their ecology in our Chicago-area study
sites. Contrary to expectations, introduced Phragmites tended
to occur in areas with equivalent or lower nutrient and salinity
concentrations than native Phragmites or reference vegetation.
However, the growth of introduced Phragmites, which was
greater than the native by all measures, showed stronger, more
positive correlations with nutrients, soil moisture, and salinity.
Wetland plant communities associated with the tall, dense
growth of introduced Phragmites had low species diversity
and were characterized by tolerant, generalist species. In con-
trast, areas with native Phragmites did not differ from refer-
ence vegetation in terms of diversity, and only marginally
differed in species composition.
Evidence in support of the eutrophication hypothesis was
mixed. In general, we did not find that Phragmites tended to
occur in areas with elevated soil nutrient concentrations or that
introduced Phragmites was associated with more eutrophic
conditions than native Phragmites. However, one exception is
that Phragmites of either lineage was found under significant-
ly higher PO4 concentrations than reference vegetation
(Table 1). Evidence from the literature is also mixed. For
example, Tulbure and Johnston (2010) did not find significant
differences in N or P availability between plots with and
without Phragmites in Great Lakes coastal wetlands, while
Silliman and Bertness (2004) found that N availability was a
 Wetlands

Fig. 2 Regressions of native vs.

introduced Phragmites phenotype
as a function of soil properties.
Solid and dashed lines indicate
trends that were significant at
P<0.05 and P<0.10, respectively.
a Soil moisture, Native: not
significant, Introduced: ln(y)=
0.988+0.782×ln(x). b Salinity,
Native: not significant,
Introduced: ln(y)=1.4028+
0.4347×ln(x). c Soil NH4, Native:
not significant, Introduced: ln(y)=
3.042+0.504×ln(x). d Soil NOx,
Native: ln(y)=3.705–0.160×ln(x),
Introduced: not significant. e
Orthophosphate, Native: ln(y)=
3.647+0.1625×ln(x), Introduced:
ln(y)=4.226+0.1608×ln(x)

key driver of Phragmites invasion in New England coastal
marshes. An important caveat of ours and other observational
studies is the ‘chicken or egg’ problem of invasive species
both responding to and potentially influencing nutrient avail-
ability. Phragmites invasion can be driven by elevated nutri-
ents but it can also influence nutrient availability through both
direct uptake and alteration of soil microbial processes
(Templer et al. 1998; Meyerson et al. 2000; Ehrenfeld 2003;
Windham and Meyerson 2003). In observing a lack of
differences in N availability between areas with and
without Phragmites or P availability between areas with
native vs. introduced Phragmites, we cannot rule out the
possibility that robust growth of Phragmites is itself
altering nutrient availability.
Wetlands

Fig. 3 Plant diversity (excluding Phragmites) as a function of the im-
portance values of native vs. introduced Phragmites. Native: not signif-
icant, Introduced: ln(y)=3.810–0.780×ln(x)
Overall, introduced Phragmites reached greater height, stem
density, and cover than native Phragmites (Table 1), and intro-
duced Phragmites’ growth was more positively correlated with
Soil moisture
1.0
nutrient availability than the native (Fig. 2), which supports the
efficient-use hypothesis (Zedler and Kercher 2004). These
patterns have been observed in introduced Phragmites in a
variety of systems (e.g., League et al. 2006; Holdredge et al.
2010), suggesting that the introduced lineage may more effi-
ciently utilize nutrients. Native Phragmites also showed greater
growth with increasing soil P; however, introduced Phragmites
outgrew the native across all levels of P. Holdredge et al. (2010)
similarly observed increases in aboveground biomass with
elevated P for both lineages, but found that this effect was
stronger in the introduced lineage.
We did not find that Phragmites occurred under different
soil moisture or salinity conditions than reference vegetation
or that introduced Phragmites occurred in areas of different
moisture or salinity than native Phragmites, which would
have been consistent with the broader-tolerance hypothesis
(Zedler and Kercher 2004). However, growth of introduced
(but not native) Phragmites was positively correlated with
increases in both moisture and salinity (Fig. 2), which has
been found in previous studies (Vretare et al. 2001; Vasquez
et al. 2005). We encountered only narrow ranges of these
factors, probably due to site-selection criteria that excluded
habitats like deeper water detention basins or wetlands adja-
cent to roads that could be subjected to high salt inputs.
We found no evidence of native Phragmites functioning as a
native invader. In fact, areas with vigorous stands of native
Phragmites were indistinguishable from uninvaded areas in
terms of plant diversity and showed only marginal differences
in composition. As expected, occurrence of introduced
Phragmites was associated with reduced plant diversity

(Table 1, Fig. 3) and Phragmites importance was a strong

Phragmites
importance value predictor of local species composition (Fig. 4). Based on
0.5

Salinity
SIMPER analysis, key species differences in areas with intro-
NOx duced Phragmites in our study included greater abundances of
PO4
NMDS2

more terrestrial and weedier species (e.g., Impatiens capensis

0.0

and Phalaris arundinacea) and lower abundances of native

wetland obligates (e.g., Carex stricta and Calamagrostis
-0.5

canadensis) compared with reference plant communities and

those containing native Phragmites.
Important caveats of our study include its observational
-1.0

Phragmites
None
Native
nature and limited geographic scope and sample size. Also,
Introduced the differences we observed between introduced and native
-1.5

Phragmites are probably conservative due to our site-selection

-1.0 -0.5 0.0 0.5 1.0 1.5
NMDS1
Fig. 4 NMDS ordination of plant-community composition (excluding
Phragmites) by plant community type (three axes, stress=9 %). Ellipses
depict one standard deviation around the centroid of each group. Composi-
tion significantly differed by plant community type overall (PERMANOVA:
P<0.0001). In pairwise comparisons, areas with native Phragmites and no
Phragmites only marginally differed (P=0.058) while areas with introduced
Phragmites significantly differed from both other groups (P<0.0001). Over-
laid vectors were significantly correlated with community composition
(Phragmites importance value: P<0.0001, PO4: P<0.0001, Salinity:
P=0.011, Soil moisture: P=0.015, NOx: P=0.025)
2.0
criteria: we did not include roadsides, detention basins, farm
fields, or other anthropogenic habitats where introduced
Phragmites often occurs (Lelong et al. 2007; Jodoin et al.
2008)—locations likely to have highly altered hydrology,
elevated nutrient and salinity concentrations, and depauperate
plant communities. In addition, we excluded locations that
had only small patches or scattered stems of Phragmites. This
increased our chances of observing invasive-like behavior in
native Phragmites as we only sampled areas where it grew
especially vigorously.

The patterns we found are fully consistent with Saltonstall’s
(2002) cryptic-invasion hypothesis, i.e., that the invasiveness of
Phragmites in North America is driven by introduction and
spread of non-native genotypes—as has been found elsewhere
in the Midwest, Gulf Coast, and western U.S. (e.g., Kulmatiski
et al. 2010; Meyerson et al. 2010; Tulbure and Johnston 2010).
We encourage wetland managers to identify Phragmites to
subspecies using morphological and/or genetic markers.
While there may be exceptions where the native subspecies
exhibits aggressive growth, this possibility should be weighed
against the substantial costs associated with control efforts and
the need to conserve a native subspecies that has been displaced
throughout much of its range (Saltonstall 2011).
Acknowledgments This research was supported by funding from
Illinois-Indiana Sea Grant and a Plant Biology and Conservation Student
Research Award. We thank the following agencies for granting us per-
mission to conduct research on their lands and helping us to identify
suitable study sites: Chicago Department of the Environment, Forest
Preserve District of Cook County, Forest Preserve District of DuPage
County, Forest Preserve District of Kane County, Illinois Department of
Natural Resources, Illinois Nature Preserves Commission, Indiana De-
partment of Natural Resources, Indiana Dunes National Lakeshore, Lake
County Forest Preserve District, and United States Army Corps of Engi-
neers. The following research assistants and volunteers helped with field
and lab work: Adewale Adeoba, Jeb Boyer, Cat Collins, David Ford,
Thomas Graan, and Dara Wise. Stuart Wagenius and Eric Lonsdorf
provided guidance on statistical analyses and other aspects of the project.
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