CASE STUDY

Wetland Cover Types and Plant Community

Changes in Response to Cattail-Control
Activities in the Palo Verde Marsh, Costa Rica
Florencia A. Trama, Federico L. Rizo-Patrón, Anjali Kumar, Eugenio González, Daniel Somma
and Michael B. McCoy C.

ABSTRACT
The Palo Verde marsh restoration was performed in Costa Rica to reduce invasive Typha domingensis stands. The invasion
of this cattail in the 1980s decreased the habitat quality, and thousands of waterbirds that used this seasonal marsh were
not returning. In 2002, the Organization for Tropical Studies decided to manage the marsh using a tractor with angle-iron-
paddle wheels to crush cattail. We evaluated the land cover changes in response to the prescribed management activities
through GIS and vegetation-assessment analysis. Crushing activities started in |uly 2002. We defined three management
plots: A and B) with crushing activities for different lengths of time and C) the control plot with passive management.
We evaluated wetland cover changes, richness, and dominance of plant species in the three plots. We found that owing
to management activities 1 ) cattail cover area decreased from 35% to 9% (Plot A) and from 62% to 7% (Plot B), while
in Plot C it remained almost constant (always greater than 60%); 2) the area of total live vegetation including cattail
decreased, giving way to open water and bare ground; 3) plant species richness was higher in the crushed plots than in
the control plot; 4) cattail dominance decreased and more species were sharing the marsh area in managed plots than
in the control plot; and 5) cattail cover was negatively related to species richness. We recommend continuing the control
of cattail by crushing in order to rehabilitate the marsh and conserve its biodiversity.
Keywords: aquatic vegetation responses, cattail {Typha domingensis), Costa Rica, Palo Verde Marsh, restoration

I n the neotropics, the richest and

most diverse biogeographical region
in the world, only eight countries have
implemented measures to restore or
rehabilitate threatened wetlands
(Ramsar 2005). Costa Ricas Palo
Verde is a neotropical marsh that has
been very important for more than
60 species of resident and migratory
waterbirds (Boza 1981, Boza and
Mendoza 1981, Meza 1991) (Figure
1). In 1979, more than 5,000 wood
storks [Mycteria americana), 30,000
black-bellied whistling ducks {Den-
drocygna autumnalis), 25,000 blue-
winged teals {Anas discors), 700 roseate
spoonbills (Ajaia ajaja), and many
Ecological Restoration Vol. 27, No. 3, 2009
ISSN 1S22-4740 E-ISSN 1543-4079
©2009 by the Board of Regents of the
University of V^/isconsin System.
Other species were registered in a single
count (McCoy 1996, McCoy and
Rodriguez 1994, Sánchez et al. 1985,
Vaughan et al. 1996).
Since the early 1500s, when the
Spanish arrived in the area, this wet-
land has contained cattle (Vaughan et
al. 1996). It was part of a huge cattle
hacienda until the mid-1970s, when
it was expropriated and declared as
Palo Verde National Wildlife Refuge.
In 1978, it was named Palo Verde
Biological Reserve by the Costa Rican
government (McCoy and Rodriguez
1994). In 1980, cattle were removed
(McCoy 1996) and a road was con-
structed, interrupting the flow of two
natural streams, one that supplied
water from adjacent hills and another
that allowed tidal flow into Palo Verde
Marsh (Calvo and Arias 2004). A very
dry period occurred between 1975
and 1980, when the total precipitation
records were lower than the annual
mean values (OTS and IUCN 2005,
Vaughan et al. 1982). Also, the gov-
ernment had granted water conces-
sions to large-scale rice farmers from
theTempisque River (adjacent to Palo
Verde Marsh), decreasing natural flow
and altering the river's hydrological
relationship with the marsh (OTS and
UICN 2005).
Cattail Invasion and
Effects on Plant
Communities
Cattail (Typha spp.) has colonized
many wetlands, and its proliferation
is recognized as a potential threat to
wetland ecosystems (Keller 2000,
Smith and Newman 2001, Waters
and Shay 1991). Cattails can be very

278 I* September 2009 ECOLOGICAL RESTORATION 27:3

 wetlands invaded by cattail (Bancroft
etal. 2002, Cominetal. 2001, Chau-
han and Gopal 2001, Shy et al. 1998).
In an attempt to control the cat-
tail invasion, the Costa Rican Wildlife
Service signed a five-year contract in
1986 with a local rancher to rein-
troduce 1,000 cattle into the marsh.
However, this management was not
effective because cattle were not
densely enough concentrated into wet,
cattail-infested areas, and these cattle
Palo Vanta HaHkinii Parti
were brought in from upland areas and
were not accustomed to marshy sites
(McCoy and Rodriguez 1994). From
1987 to 1990, initial research and
experimental management led to an
effective technique to control cattail in
Palo Verde Marsh (McCoy and Rodri-
guez 1994).Afarm tractor with angle-
Figure 1. Location of Palo Verde National Park, Palo Verde Marsh and cattail {Typho domingeníis)
treatment plots in Costa Rica; A) crushing; B) crushing and burning; C) control.
iron paddle wheels was used to crush
cattail under water to kill it (Figure
2). After opening 30 ha between 1989
aggressive, expanding their coverage Management Efforts and 1990, 20,000 black-bellied whis-
outward around 9-10 m/y by rhi- to Restore Palo Verde tling ducks and 12,000 blue-winged
zomes, contributing to both biotic Marsh: 1985-2002 teal were registered in a single count in
and abiotic alterations (Houlahan and the managed area (McCoy and Rodri-
Findlay 2004, Keller 2000, McGoy Owing to the management and hydro- guez 1994). Cattail crushing contin-
and Rodriguez 1994). Biotic changes logical alterations in Palo Verde Marsh ued until 1997, when 70 ha had been
include 1) a decrease in richness and from 1975 to 1985, the wetland opened up (McCoy and Rodriguez
diversity of plant species due to com- changed ftom a heterogeneous land- 1994). During this period, no plant
petition; 2) a rapid accumulation scape with open water, low grasses, diversity studies were carried out in
of biomass and leaf litter that limits and floating and low emergent vegeta- the marsh. However, by 2000, with
periphyton development; 3) a decrease tion with some tall emergent patches this opening in the marsh and the
of physical living space for organisms of cattail {T. domingensis) to a mono- accompanying lesser wildfire occur-
across trophic levels; and 4) rapidly typic stand of cattail (Castillo and rence, two tree species, palo verde and
invaded shallow water and decreas- Guzman 2004, McCoy 1994, McCoy michiguiste {Pithecellobium lanceola-
ing waterfowl habitat. Abiotic changes and Rodriguez 1994, Carbonell 1998, tum), increased their coverage to 267
include the alteration of dissolved Vaughan et al. 1996). In 1985, cat- ha (Castillo and Guzman 2004).
oxygen, light, depth, and hydrology tail invaded more than 55 percent of
Because of its importance as an
(Ball 1990, Gronk and Fennessy 2001, the wetland (around 692 ha). This
international site for waterbirds,
Ghiang et al. 2000, Keller 2000, Miao invasion, combined with a very dry
PVNP was registered as a Ramsar Site
et al. 2001, Miao et al. 2000, Newman period, led to several dry-season wild-
(internationally important wetland)
et al. 1998, Carbonell 1998, Smith fires during the 1980s that eliminated
other species, especially palo verde in 1991, and it was included on the
and Newman 2001, Sojda and Solberg
1993). trees [Parkinsonia aculeata), and pro- Montreux list of Ramsar wetlands that
mulgated a homogeneous stand of cat- need management activities in 1993
Moreover, cattail liberates pheno- tall . Cattails overtook almost all open (Quiros et al. 1991). In 1998, tbe
lic components that have inhibitory water and bare ground, causing a large Costa Rican government authorized
effects on the growth and propaga- reduction in waterbird species richness active management in the wetlands
tion of other plant species, and these and abundance in the marsh. In 1988, of PVNP (Presidential Decree No.
detrimental efltcts are not reversible as only 3,000 black-bellied whistling 27345). In 2001, the Organization
long as cattail remains (Gallardo et al. ducks and 500 blue-winged teals were for Tropical Studies (OTS) and the
1999, Gallardo et al. 1998). For these observed in the marsh, numbers which Ministery of Environment and Energy
reasons, plant species richness will be coincide with observations for other (MINAE) through its National Wet-
higher when cattail is managed. lands Program developed a plan to

September 2009 ECOLOGICAL RESTORATION 27:3 279


Figure 2. Tractor with paddie wheeis used to crush cattail in February 2004. Photo by Federico L
Rjzo-Patrón
recover waterbird habitat by restor-
ing 50 ha using the cattail-crushing
method in addition to cattle grazing
(McCoy and Rodriguez 1994, OTS
2002). In 2002, an additional 250
ha of the marsh were included in this
management plan.
To evaluate plant responses to the
crushing management activities, we
developed the following two objec-
tives: to analyze the wetland vegetation
cover changes in Palo Verde Marsh
using CIS and to analyze the commu-
nity-level plant diversity within two
managed plots, as well as a control plot
free from active management. We pre-
dicted that I) management activities
will reduce invasive cattail cover, and
diverse types of wetland vegetation
will develop; 2) greater plant diversity
will be found in managed plots com-
pared to the control plot; and 3) the
dominance of cattail will be reduced
in managed plots.
Methodology
Study Area and Design
Palo Verde National Park encompasses
20,000 ha and is situated in the Are-
nal-Tempisque Conservation Area in
northwestern Costa Rica (10''2r N,
280 September 2009 ECOLOGICAL RESTORATION 27:3
S5°2l"W). Palo Verde Marsh is a sea-
sonal marsh located inside the PVNP,
covering approximately 1,207 ha
(Castillo and Cuzmán 2004) (Figure
1). The seasonality of this marsh is
the result of rainwater and run-off
from bordering hills and drying by
evapotranspiration. The interacting
dynamics determine the volume and
depth of the marsh, in this case vary-
ing from 0 m to 1.5 m in the rainy
season and between 0 m and 0.4 m
in the dry season until drying up
completely between March and April
(Vaughan et al. 1982, Vaughan et al.
1996). In the dry season (March and
April), extraordinary tides can flood
part of the marsh, and the water can
enter and flood some zones in the last
stage of the dry season (Vaughan et
al. 1982, Vaughan et al. 1996). Thus
toward the beginning of the rainy
season, the floating aquatic vegeta-
tion, mainly dotleaf waterlily [Nym-
phaea ampld), water dead and awake
{Neptunia plena), and water hyacinth
{Eichhornia crassipes), starts to grow
in open-water areas. Other areas of
the marsh are dominated by emergent
vegetation, including sedges (Cyper-
aceae) and grasses (Poaceae), which
form large stands. Still other areas are
invaded by cattail and talia or bent
alligator-flag (7Aíí/íí2^^«¿c«¿2í/2). Some
submerged plant species, Cuadalupe
waternymph [Najas guadalupensis)
and prickly hornwort [Ceratophyllum
muricatum), can be found in deep and
open water areas. However, palo verde
trees, semiaquatics like yerba de jicotea
[Ludwigia erecta), and shrubs like
catsclaw mimosa {Mimosa pigra) and
other legumes (Fabaceae) can be dis-
tributed in shallow-water areas. When
the dry season begins, the aquatic and
semiaquatic plants begin to die, but
some seeds, bulbs, or rhizomes remain
in the propagule bank for the next wet
season (Chauhan and Gopal 2001,
Middleton et al. 1991). However,
cattail remains in the lagoon all year
long, since soil under the plants retains
moisture during the dry season (Crow
2002, Carbonell 1998).
Both hydroiogical rehabilitation
and vegetation management activities
started in July 2002. The two streams
that were interrupted in the 1980s
were rehabilitated to reestablish the
wetland hydrology, and water levels
were monitored by OTS (Jiménez et
al. 2003, Trama 2005). Three 80 ha
management plots were established.
In Plot A, cattail was crushed without
controlled fires from July 2002 to the
end of this study. Prior to this man-
agement plan, all of the 1987-2002
experimental management activities
were carried out in 50 ha of this plot.
In Plot B, cattail was managed with
crushing and controlled flre from Sep-
tember 2002 to the end of this study.
Prior to this study, there were no man-
agement activities in this plot. In plots
A and B, three different tractors were
used to crush cattail underwater twice
a year for 3 years In early wet season
and early dry season, when the water
levels were below 35-40 cm. Areas
with cattail were crushed two or three
times depending on density until cat-
tail was totally crushed under water. In
plot B, two controlled fires occurred in
the dry season to reduce cattail stands
that remained unmanaged. Plot C was
the control that did not receive any
active management during or before
this study (Figure 1).
Monitoring
We carried out a photo-interpretation
analysis using ArcView 3-3 (ESRI,
Redlands CA) to estimate wetland
cover changes in all plots before
and during management activities.
We analyzed two aerial photographs
and one satellite image: 1) a Terra
orthophotograph (resolution 30 m)
acquired November 1998, late wet
season/early dry season; 2) a three-
band Ikonos image (resolution 4 m)
acquired December 2002, late wet
season/early dry season, five months
after management activities began; and
3) an infrared aerial photograph (reso-
lution 2 m) from the CARTA project
of March 2003, late dry season, nine
months after management.
We delineated eight wetland cover
classes depending on the season
and management plot (Crow 2002,
Hernández 1990, Hernández and
Gómez 1993, Trama 2005): 1) cattail
stands; 2) crushed cattail; 3) emer-
gent plants, including grasses, sedges,
legumes, and spurges (Euphorbia-
ceae); 4) floating vegetation, includ-
ing water hyacinths (Pontederiaceae),
wateriilies (Nymphaceae), duck-
weeds (Lemnaceae), and tape-grasses
(Hydrocharitaceae); 5) open water
areas; 6) palo verde trees; 7) exposed
ground, delineated during dry season
and including dry or wet mud, typi-
cally free of aquatic vegetation, but
in some cases covered by some dry-
season plant species; and 8) other veg-
etation cover, including emergent and
floating vegetation for the 1998 aerial
photograph.
We defined a minimum mapping
unit (mmu) of 30 m for the 1998
photograph based on the resolution
of imagery used. The mtnu defined
for the 2002 and 2003 analyses was 5
m, making the two years directly com-
parable to each other. Accuracy assess-
ment of classified areas was performed
using data provided by managers and
by ground truthing in the marsh.
We sampled vegetation 11 times
in the three management plots from
August 2003 to July 2004, including
both a dry and a wet season, using six Results
25 m X 2 m transects (300 m^ total)
for each plot every month. Plant spe- CIS Analysis
cies were identified using Crow (2002)
In 1998, baseline wetland cover in
and the Palo Verde Biological Station
the three plots consisted of four cover
herbarium collection. Each plant spe-
types: palo verde, open water, cattail,
cies was counted along each transect,
and other vegetation. In December
and its percent cover was estimated.
2002, six cover types were differenti-
To facilitate the estimation of percent
ated: palo verde, open water, cattails,
cover, we used a 2 m x 2 m quadrat
crushed cattails, emergent vegetation,
subdivided into four sections. For each
and floating vegetation. By March
transect, we used 12.5 quadrats (75 per
2003 in late dry season, another wet-
management plot). We estimated the
land cover class was identified in the
total cover of every plant species in all
crushed areas: exposed ground (Figure
quadrats along each transect. A value
3). During the first five months of
of 1% was given to those species that
crushing activities, cattail cover was
covered 1 % or less of the area sampled.
reduced from 34.7% (baseline) to
Clonal reproduction of many wetland
23.9% in plot A and from 61.5% to
plant species prevented calculation of
52.2% in plot B, while in the control
abundance (Crow 2002).
plot it increased by 5.1 %. Afterward,
Data Analysis from December 2002 to March 2003
cattail cover decreased by 25.5% in
We carried out a descriptive GIS plot A and 54.5% in plot B. How-
analysis comparing the wetland cover ever, plot C, the unmanaged plot, was
classes before the restoration initia- always more than 60% covered by
tive started (baseline at 1998) and cattail.
five months (2002) and eight months
A change in the area occupied by
(2003) after management activities
TLVA due to cattail-crushing manage-
began. The estimated coverages were
ment in plots A and B was observed
compared between years and manage-
during 2002-2003 (Figure 3). In
ment plots. We calculated the Total
1998, TLVA, including cattail, was
Live Vegetation Area (TLVA), includ-
95.1% of plot A, 99.9% of plot B, and
ing cattail and other vegetation, as well
98.7% of plot C. By December 2002,
as other wetland cover classes as a total
TLVA decreased to 85.4% in plot A,
for each year or plot analysis.
and to 67.6% in plot B, but still cov-
The six transects were considered
ered 99.9% of the control plot (C).
replicates for each plot. The mean spe-
In March 2003, TLVA was 28.2% of
cies richness was compared between
plot A, 22.6% for plot B, and 97.9%
the three plots by one-way ANOVA
of the control plot. The remaining
(LSD interval for equal samples) using
area in all cases was occupied by open
Statgraphics 3.1 (Manugistics, Rock-
water, exposed ground, and managed
ville MD). For the seasonal distribu-
sectors with crushed cattail.
tion of species, we tested interactions
During management activities,
between dates and all three plots.
emergent and floating vegetation
Using the vegetation sampling data,
could be delineated, and several sea-
we estimated the percentage of TLVA
sonal changes were observed in the
and other wetland covers. Dominance
plots. In 1998, open water occupied
of cattail and other plant species was
4.9% of plot A and 0.1% of plot B.
compared between the three plots.
In December 2002 after five months
We defined a plant as dominant when
of tnanagement, the percentage of
it covered more than 10% of the
open water increased to 9.5% and
sampled area within a plot on at least
16.1% in plots A and B, respectively.
one sampling date. Finally, a linear
Additionally, we observed that in the
regression analysis determined if cat-
dry season (from December 2002 to
tail cover influenced species richness.
September 2009 ECOLOGICAL RESTORATION 27:3 ** 281
 1998

2002

I I Floating Vegetation
•I Crushed Cattail
n3 Exposed Ground
^1 Emergent Vegetation
^1 Open Water
• Palo Verde
^1 Other Vegetation
im Cattail

2003

900 1600 Meters

Figure 3. Wetland cover by treatment plot, before (1998) and during (2002, 2003) management activities.

282 September 2009 ECOLOGICAL RESTORATION 27:3

Table 1. Plant species recorded in each treatment plot: A) plot managed since 1987 with cattail crushing; B) plot
managed since 2002 with cattail crushing and burning; and C) control plot without active management.
Plot
Family Species

Alismataceae Echinodorus paniculatus X

Araceae Pistia stratiotes X
Asteraceae Edipta prostrata X
Boraginaceae Heliotropium sp. X
Cannaceae Canna glauca X
Ceratophyllaceae Cerataphyllum muricatum X
Convoivulaceae Aniseia martinicensis X
Ipomoea carnea X
Cucurbitaceae Luffa operculata X
Cyperaceae Cyperus articulatus X
C. digitatus X
C tria X
C. tenerrimus X
Eleocharis elegans
E. interstincta X
E. mutata X
Fimbristylis spadicea (= F. thermalis)
Oxycaryum cúbense X
Elatinaceae Sergio capensis X
Euphorbiaceae Caperonia castaneifolia X
Crotón argenteus X
Euphorbia sp. X
Fabaceae Aeschynomene sensitiva X
Mimosa pigra X
Neptunio natons X
Parkinsonia aculeato X
Sesbonio emerus X
Hydrocharitaceae Limnobium iaevigotum X
Nojas guodalupensis X
Hydrophyllaceae i-lydraieo elatior
Lemnaceae Lemna aequinoctioHs X
Wolffieila welwitschii X
Lentibulariaceae Utricuiaria foliosa X
U. gibbo X
Malvaceae Molochro fasciota X
Molvostrum americanum X
Marantaceae Tholia geniculota X
Marsileaceae Morsiieo deflexa
Nymphaeaceae Nymphoeo omazonum X
Nymphaeo ampio X
Onagraceae Ludwigio erecto X
Ludwigio indinota X
Parkeriaceae Cerotopteris pteridoides X
Poaceae Echinochloa colono X
Hymenachne omptexicaulis X
Leersia hexondra
Paspolidium geminatum X
Pospalum repens X
Pospolum sp. X
Polygonaceae Polygonum hispidum
P. segetum X
Pontederiaceae Eichbornio crassipes X

September 2009 ECOLOGICAL RESTORATION 27:3 283

 Table 1., continued

Plot
Family Species

Heteranthera limosa
Portulacaceae Portulaca sp.
Rubiaceae Borrerio assurgens
Azolla mexicana
Salviniaceae Salvinia minima
Bacopa axillaris
Scrophulariaceae B. repens
Physalis angulota
Solanaceae Solanum campechiense
Typha domingensis
Typhaceae Phyla nodiflora
Verbenaceae Unidentified 1
Malvacea Unidentified 2
Verbenacea Unidentified 3
Asteraceae Unidentified 4
Unidentified 5
Unidentified 6

March 2003), che area of open water
decreased owing to desiccation, to
8.5% in plot A and 0.4% in plot B.
However, in plot C, open water was
always around 2% cover because that
part of the marsh is more than one
meter deep and does not dry up until
around April. Also, the floating veg-
etation cover decreased from 24% to
7.6% in plot A and 0.9% to 0% in
plot B owing to the normal seasonal
change of the marsh. We did not
detect floating vegetation in plot C.
At the same time, exposed ground was
appearing in the dry season when open
water areas and floating vegetation was
decreasing, covering 32.9% in plot A
and 17.8% in plot B. Exposed ground
was not detected on the control plot in
any of the analysis years, as this cover
appears only after open water areas dry
up (Figure 3).
Vegetation Sampling
Ihe total plant species richness of the
marsh was 69; we found 59 species
in plot A, 61 in plot B, and 20 in
plot C (Table 1). The mean species
richness was higher in plots A (13.2)
and B (11.8) than in plot C (4.2)
{F= 104.94, df = 2.197,/- < 0.0001).
Additionally, mean species richness
varied temporally during this study
( f = 3 . 3 1 , df= 20,197,;'< 0.0001)
(Figure 4).
According to our vegetation tran-
sects, cattail cover was always less than
20% in plot A, less than 40% in plot
B, and greater than 70% in plot C
during this study [F = 884.55, df =
2,197,p < O.OOOl) (Figure 4). Total
live vegetation area varied seasonally
in the managed plots but remained
almost constant in the control plot
(> 85%) (Figure 5). Additionally, in
the dry season (January to May), the
percentage of TLVA was lower in the
managed plots. Moreover, we found
that dominance was shared among
more species in the managed plots
than in the control plot. In plots A
and B, we found cattail and tnore
than 50 other plant species with
greater than 10% cover, whereas in
plot C, cattail was clearly the domi-
nant species, and less than 20% of
the total area was shared with 19
species (Figure 6). Furthermore, the
regression analysis showed a nega-
tive relationship between cattail cover
and plant species richness by transect
(Figure 7). Fire did not have a sig-
nificative effect on cattail in plot B.
Cattail started growing again 30 days
after the burn. In the wet season, the
tractor had to pass over the burned
area to reduce this cover.
Discussion and
Recommendations
Cattail in tropical and some temper-
ate wetlands is considered an invasive
plant that reduces habitat availability
for wildlife and plant communities
(Farnsworth and Meyerson 2003,
Crace and Wetzel 1981, Houlahan
and Findlay 2004, Lee et al. 1996).
Monotypic cattail stands reduce over-
all habitat value, affecting some species
of wildlife such as waterfowl, water-
birds, fish, amphibians, and inver-
tebrates (Lee et al. 1996, Sojda and
Solbergl993).
The results of this study show that
crushing activities have significantly
reduced cattail stands and improved
habitat for other plant species. In 1998,
the difference in cover classes between
plot A and plots B and C is striking.
By then, plot A had already experi-
enced more than ten years of experi-
mental active management (McCoy
and Rodriguez 1994). Plot A had less
cattail cover and the rest of the plot
was occupied by other vegetation and
open water. However, in plots B and C,
cattail was present over more than 60%
of the area owing to its invasiveness
and the lack of management. Between
2002 and 2003, plots A and B showed
an increase in plant cover types and

284 September 2009 ECOLOGICAL RESTORATION 27:3

 dormant, and exposed ground replaces
all the other areas except cattail. This
is because cattail does not vary season-
ally like other vegetation in a seasonal
Rot marsh (Crow 2002). By decreasing
I — A cattail cover and, consequently, the
er percentage of TLVA, management
B
activities have permitted the reestab-
- C lishment of the wetland heterogeneity.
CO
ö
o r \.H. I From the beginning of cattail-
crushing activities, the vegetation
assessment showed that plant species
richness varied both between treat-
ments and from seasonal changes
c o c o
during the year. However, species rich-
99 ness was always higher in the managed
plots when compared to the control
plot. In plots A and B, there were
SarrpJingcËte many dominant species compared to
the one dominant plant (cattail) in
the control plot. Additionally, 60%
110 of the variation seen can be explained
by the relationship between cattail per-
90 cent cover and species richness. This
implies that management is necessary
Rat to improve and maintain habitat in
— A Palo Verde Marsh.
if 50 We assume that the return of the
B
m wetland heterogeneity is related to the
C recovery of flora, fauna, and ecologi-
Ü '*' cal processes (Ruiz-Jaen and Mitchell
2005, Young 2000). In complemen-
10 tary assessments, greater species rich-
ness and numbers of waterbirds were
observed in plots A and B than in plot
C. Of 62 bird species recorded in the
study, 54 were registered in plot A, 52
in plot B, and 22 in plot C, with only
Ssrrplingdate 12 in the cattail cover. Moreover, more
than 35,000 black-bellied whistling
Figure 4. Seasonal variation in mean plant species richness (top) and percent cover of cattail
(Jypha domingensii) (bottom) in three experimental plots: A) plot managed since 1987 with cat-
ducks and about 9,000 blue-winged
tail crushing; B) plot managed since 2002 with cattail crushing and burning; and C) control plot teals were observed in the managed
without active management. plots in a single count (Trama 2005).
Open water and exposed ground were
plant species diversity compared with activities is related to larger and more the two wetland covers preferred by
the control plot, which demonstrates numerous areas of shallow water or waterfowl and other waterbirds,
that this particular type of manage- exposed ground in the dry season, which have used the wetland during
ment is successfial. The small changes conditions that are normal in this sea- the crushing activities (Colweli and
in wetland cover in the control plot sonal marsh (McCoy and Rodriguez Taft 2000, Green 1996, Trama 2005).
observed during the study could be due 1994). After cattail was crushed in Additionally, more and different taxa
to variations in resolution. In spite of plots A and B, these areas were occu- of invertebrates have been recorded in
this, control plot C always had more pied by open water or other wetland managed plots than in the control plot
than 60% of cattail coverage. cover classes. During dry-season desic- (Rizo-Patrón et al. 2009).
The reduction of total live vegeta- cation, the open water decreases, the However, after almost 20 years of
tion area (TLVA) due to the crushing seeds of floating vegetation become invasion by cattail, this plant is part of

September2009 ECOLOGICAL RESTORATION 27:3 285

 PIOtA
grow back very fast (McGoy 1994).
Instead of being eradicated, most of
the cattail stands were reduced, so the
risk of teinvasion remained (McCoy
and Rodriguez 1994, SER 2004). In
cases where water levels were low ( 15
cm) during crushing, some cattail
resprouted. We recommend manu-
ally cutting this sparse regrowth to
reduce the need for future crushing
in the same area. Otherwise, cattail
returns quickly to original densities. It
PlotB is necessary to continue the manage-
ment every year to guarantee cattail
is controlled. Also, it is important to
note that management activities with
the tractor were directly associated
with more open areas free of cattail.
TLVA These areas were colonized by other
Other habitats plant species that behaved like normal
seasonal plant communites. In this
case, seasonality was responsible for
the reduction of floating vegetation
and open water and their replacement
by exposed ground from December
2002 to March 2003 (Trama 2005).
Fire as a management tool was not
successful because it occurred in the
dry season. Cattail started to grow
very fast and there was no water in
the marsh to allow the tractor to enter
and control it. Additionally, fire was
very bad for wildlife that use cattail
stands as a refuge in the dry season
(Castañeda 2003).

The key to conservation of inland

wetland biodiversity is to discourage
the spread of community dominants
Sampling date
(Houlahan and Findlay 2004). Thus it
Figure 5. Percentage of Total Live Vegetation Area (TLVA) and other cover types (inciuding is necessary to continue with manage-
crushed cattail, open water, and exposed ground) during this study: A) plot managed since ment activities to maintain the Palo
1987 with cattail crushing; B) piot managed since 2002 with cattail crushing and burning; and Verde wetland habitat by controlling
C) control plot without active management.
invasive species. Crushing with the
tractor is the first step and should be
the Palo Verde marsh and several spe- Management intensity varied complemented by grazing to control
cies of birds, reptiles, and invertebrates depending on availability of the cat- other species that appear after cat-
use this cover, mainly for resting and tail-crushing tractor and water levels. tail elimination, such as West Indian
protection from predators (Castañeda The tractor could not enter when marsh grass {Hymenachne amplexi-
2003, Trama 2005, Vaughan et al. water was deeper than 45 cm. In some caulis) and Egyptian panicgrass {Pas-
1996). Therefore, leaving cattail paths areas, the tractor had to pass two or palidium geminatum), both of which
or corridors in some areas and near the three times to complete the cattail produce an effect similar to cattail as
fence line has been suggested to help crushing. Because cattail is rhizoma- far as decreasing habitat quality for
supply the seasonal requirements of tous, if it is not eradicated within 15 waterfowl. Grasses can also be a very
wildlife (Vaughan et al. 1996). days of management activities, it can dangerous fuel source for wildfire

286 September 2009 ECOLOGICAL RESTORATION 27:3

 Plot A
40-
PlolB
PlotC
Sampling date
Figure 6. The relative cover of the most dominant plants during this study: A) plot managed since
1987 with cattail crushing; B) plot managed since 2002 with cattail crushing and burning; and
C) control plot without active management.
in the dry season (Barboza 2005,
Vaughan et al. 1996). Palo Verde
National Park administration should
improve grazing supervision and take
advantage of the cattle present at the
park. Cattle should he concentrated in
smaller areas with fencing and rotated
between fenced parcels. Cattle density
•*T'
\.\ )
HymenactiiTv I
N»pliinienaians
II I ' Nymphaoe amBiormm
(B39 TttaKa geniculata
^ B LimnotMum faanoatum
• • O'ycaryum cubens«
t^^-^ Eictionia crasspos
ODwr 59 epeom
is usually too low to produce an
impact on vegetation, unless restricted
by fencing. Recent research shows that
proper cattle grazing with fencing does
control cattail and grasses and increase
plant diversity (M. McCoy, pers. obs.).
Additionally, some activities to
restore hydrology have occurred
September 2009 ECOLOGICAL RESTORATION
(Calvo and Arias 2004). The results
and implications of these activities
are still being analj'zed. However, it is
believed that the reestablishment of
the two previously rerouted streams
will permit the wetland to fill with
rainwater more quickly. Cattail and
aquatic grasses cannot survive as easily
if the wetland is filling quickly with
rainwater and therefore will likely
establish in fewer areas. With crushing
activities and hydrological restoration,
it is predicted that Palo Verde Marsh
will be rehabilitated to a self-regulat-
ing ecosystem with conditions similar
to those prior to the cattail invasion.
Although this study only considered
the management responses in Palo
Verde Marsh, similar responses could
be obtained in other wetlands invaded
by cattail. Inside PVNP, La Bocana
Wetland also has problems with cattail
invasion (Rizo-Patrón 2004), while
Varillal and Piedra Blanca wetlands
have problems with catsclaw acacia
{Mimosapigra). We recommend evalu-
ating possible management activities
in these other wetlands to recover and
conserve these habitats and their bio-
diversity. After the management done
in Palo Verde, thousands of ducks and
other waterbirds were observed at the
wetland, and the historic abundance
of individuals recorded before the
cattail invasion for some species has
recovered (Trama 2005). In this way,
including several sites in a restoration
strategy to recover wetlands inside and
around the national park would ensure
the conservation of these habitats and
biodiversity at a landscape level.
Habitat improvement to benefit
wildlife was the main objective of the
restoration initiative in Palo Verde.
Few data exist on waterbirds' pref-
erences for cover class areas in Palo
Verde Marsh; however, a more het-
erogeneous wetland and more shal-
low-water or exposed-ground areas
could imply more feeding and resting
habitats for more than 60 waterbird
species (McCoy 1996, Trama 2005).
We recommend studies on waterbirds'
preferences for each vegetation type.
27:3 287
 100 —L bv¿ g • —
Cronk, J.K. and M.S. Fennessy. 2001. \^et-
Ban D o
- X. ^& D^\n u• B
a §
\ -
- land Plants: Biology and Ecobgy. Boca
- \.^~N. Raton FL: CRC Press.
50 • \ -
• ^
v Crow, G.E. 2002. Plantas Acuáticas del
\ Q \
Parque Nacional Palo Verde. Heredia,
•

N X.
8
60 -

-
N
\
X n
G ^
Costa Rica: INBio.
Farnsworth, E.J. and L.A. Meyerson. 2003.
\
Comparative ecophysiology of four wet-
attail

AQ D
D •
\ ^ 5 - land plant species along a continuum
o \
X ofinvasiveness. U^f¿íní¿ 23:750—762.
20 -
° \ . • Gallardo, M.T, J. Ascher, M.J. Collier. B.B.
\

\ Martin and D.E Martin. 1999. Effect

Q D
N •
-

of cattail ( Typha domingensis) extracts,

0 0 -^ -
- , . . . 1
leachates, and selected phenolic com-
10 15 20 25 30 pounds on rates of oxygen production
Plant species richness by salvinia {Salvinia minima). Journal of
Aquatic Plant Management 37:80-82.
Figure 7. Relationship between cattail cover and plant species richness, (f = 268.65, df = 1,112, Gallardo, M . T , B.B. Martin and D.F.
R* = 0.60, p < 0.0001). Martin. 1998. Inhibition of water
fern Salvinia minima hy cattail ( lypha
domingensis) extracts and by 2-chloro-
Acknowledgments Boza, M.A. and R. Mendoza. 1981. The
phenol and salicylaldehyde. Journal of
Our sincerest thanks to Organization for National Parks of Costa Rica. Madrid:
Chemical Ecology 24:1483-1490.
Tropical Studies (OTS), the Instituto en INCAFO.
Grace, J.B. and R.G. Wetzet. 1981. Hab-
Conservación y Manejo de Vida Silvestre Calvo, J.C.A. and O.R. Arias. 2004. Res- itat partitioning and competitive dis-
ICOMVIS (formerly PRMVS), and the tauración hidrológica del humedal Palo
placement in cattails: Experimental
Ministerio de Ambiente y Energía (MINAE) Vtiáe. Ambientico 129:7-8.
field studies of intensity of competi-
for making possible this project. To Ducks Carbonell. M. 1998. Sitio Ramsar Parque tion. American Naturalist 118:463-474.
Unlimited, the United States Fish and Wild- Nacional Palo Verde, Costa Rica. Misión
Green, A.J. 1996. Analyses of globally
life Service (USFWS). Costa Rica-United Ramsar de Asesoramiento Report
threatened anatidae in relation to
States Foundation (CRUSA), The Ramsar No. 39. Gland, Switzerland: Ramsar
threats, distribution, migration patterns,
Convention, AVINA. and IDEA WILD Convention Office.
and habitat use. Conservation Biology
for supporting the Palo Verde restoration Castañeda, E. 2003. Informe del impacto 10:1435-1445.
project. Also, we would like to thank the de las quemas en la laguna Palo Verde.
Hernández, D.A.E. 1990. Cambios anu-
OTS administration that helped co get the Guanacaste, Costa Rica: Organization
ales en la composición y distribución
research and collecting permit (No. 30856). for Tropical Studies.
de la vegetación acuática en el humedal
Thanks go to my friend Montserrat Car- Castillo, M. andJ.A. Guzman. 2004. Cam- estacional de Palo Verde, Costa Rica.
bonell, who has always encouraged me with bios en cobertura vegetal en Palo Verde
Heredia, Costa Rica: Organization for
her advice; to Mauricio Castillo and José según SIG. Ambientico 129:4-6.
Tropical Studies.
Guzman for technical support with GIS; Chauhan, M. and B. Gopal. 2001. Biodi- Hernández, E.D. and J. Gómez. 1993. La
and a very special acknowledgment to Ken versity and management of Keoíadeo
Flora Acuática del Humedal Palo Verde.
Kriese and Dawn Browne for their editing National Park (India)—A wetland of
Heredia, Costa Rica: EUNA.
and advice. international importance. Pages 217-
Houlahan, J.E. and C.S. Findlay. 2004.
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