Wetlands Ecol Manage

https://doi.org/10.1007/s11273-021-09802-y (0123456789().,-volV)
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89().,-volV)

ORIGINAL PAPER

Effect of rotational grazing management on vegetation

of floodplain wetlands
Seimandi Gisela . Mesa Leticia . Marı́a Luz Sánchez . Saigo Miguel .
Gutiérrez Hugo

Received: 24 July 2020 / Accepted: 19 April 2021

Ó The Author(s), under exclusive licence to Springer Nature B.V. 2021

Abstract In Argentina, the expansion of soybean
production has forced the relocation of cattle to
marginal sites for agriculture, such as the wetlands
of the Parana River floodplain, constituting a threat to
the vegetation of these ecosystems. In this context, the
aims of this study were (1) to assess the effect of
different resting/grazing periods on the composition,
richness, diversity and dominance of functional
groups of wetlands vegetation assemblages and (2)
to analyze the influence of cattle exclusion on these
parameters. For these purposes, three wetlands sub-
jected to rotational grazing management included in
different paddocks with different resting and grazing
periods were selected. Exclosures were established in
upland and lowland zones. Samples of vegetation
cover were collected in these zones with quadrats of
4m2. Our results demonstrated that vegetation param-
eters did not differ as grazing and resting time period
differed. Both the wetland with short resting/long
grazing period as well as the wetland with long resting/
short grazing period had lower diversity and richness
S. Gisela  M. L. Sánchez  G. Hugo
Facultad de Humanidades y Ciencias (FHUC-UNL),
Ciudad Universitaria, Paraje El Pozo, 3000 Santa Fe,
Argentina
M. Leticia (&)  S. Miguel
Instituto Nacional de Limnologı́a (INALI-CONICET-
UNL), Ciudad Universitaria, Paraje El Pozo,
3000 Santa Fe, Argentina
e-mail: letimesa@hotmail.com
in comparison with the wetland without cattle grazing.
Grazing exclusion favoured the survival of species in
upland zones, increasing total richness and diversity
and the development of unpalatable non-forage and
shrub/subshrub species. Exclosures would be benefi-
cial to the ecological condition of the vegetation of the
wetlands, allowing grazing-sensitive species to
recover from grazing and trampling.
Keywords Floodplain wetlands  Cattle grazing 
Palatable plants  Rotational management  Parana
river system
Introduction
Wetlands are productive ecosystems that host a great
biodiversity and provide a wide variety of essential
ecosystem services for society (McInnes et al. 2017;
Moges et al. 2018). Despite their high value, ecolog-
ical conditions of wetlands continue to decline due to
increasing pressure from both direct and indirect
human activities (National Research Council 2001;
TEEB 2013).
Livestock grazing affects wetland plant communi-
ties as it significantly changes the species composition
and functional structure (Reeves and Champion 2004;
Moran et al. 2008). Through selective herbivory and
associated activities (trampling, fraying, bedding,
urination and defecation), cattle can drastically

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modify the composition, functional diversity and
dynamics of wetland plant communities, increase the
proportion of unpalatable biomass for cattle, facilitate
plant invasions, and alter water and nutrient cycles
(Jackson and Allen-Diaz 2006; Lyseng et al. 2018;
Sonnier et al. 2020). Through these effects, grazing
has negatively affected biodiversity in some ecosys-
tems, impairing erect plants and reducing woody
species cover (Painter and Belsky 1993; Fleischner
1994; Jones et al. 2011; Magnano et al. 2013; Andrew
et al. 2015; Rahmanian et al. 2019). Conversely, in
other systems, livestock can also serve as a functional
equivalent of large native herbivores and play an
essential role maintaining biodiversity, increasing
native species richness and favoring forage species
(Harrison 1999; Maestas et al. 2003; Hayes and Holl
2003; Marty 2005; Jacobo et al. 2006; Lodge and
Tyler 2020).
In Argentina, since the late twentieth century, the
expansion of soybean production has forced the
relocation of cattle into marginal sites for agriculture
such as wetlands, constituting a threat to these
ecosystems (PROSAP 2009). This situation has
increased the livestock density in floodplain systems,
about 100% in one decade (Quintana et al. 2014).
The Middle Parana River in Argentina is one of the
most important wetland ecosystems in South America
because of its location and extent, which is being
modified quite rapidly (Baigún et al. 2008). Because of
their extensive areas of vegetation for forage and high-
quality water supply, the wetlands of the Middle
Parana River system provide excellent conditions for
cattle (Mesa et al. 2015). In these systems, cattle
historically grazed wetlands and play an essential role
in vegetation composition (Burkart 1969; Biasatti
et al. 2016). Recently, grazing managers have begun to
implement rotational grazing systems in this zone to
control the frequency and intensity of grazing and rest
periods in order to optimize livestock and plant
performance (Beetz and Rinehart 1996; Teague et al.
2010; Raymond and Vondracek 2011). This strategy
improves and maintains forage resource and increase
animal production (Sovell et al. 2000). Cattle are
continually faced with younger and more nutritious
forage throughout the growing season (Kothmann
1984).
The significant increase of cattle stocking rates in
floodplain systems of the Middle Parana River, added
to the limited understanding of its effects on the
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Wetlands Ecol Manage
vegetation composition, have pointed towards a need
for identification of proper management in these areas.
Consequently, the aims of this study were (1) to assess
the effect of different resting/grazing periods on the
composition, richness, diversity and dominance of
functional groups of wetlands vegetation assemblages
and (2) to analyze the influence of cattle exclusion on
these parameters. We hypothesized that the vegetation
of the wetland with longer grazing and shorter resting
periods have lower diversity and richness. Moreover,
in this wetland, we expect a dominance of forb and
prostrate vegetation. Finally, we predict the exclusion
of cattle promotes vegetation recovery increasing total
richness, diversity and diversity of non-forage, shrub
and erect species.
Materials and methods
Study area
The Parana River drains the second largest watershed
in South America, after that of the Amazon, covering
an area of 3.1 9 106 km2. Most of its course is flanked
by a 10–50 km wide floodplain, occupying an area of
60,000 km2 (Fig. 1a). The hydrological dynamics of
the Parana River are characterized by high water
phases with flows of up to 25,000 m3 s-1 which take
place from October to March, and by low water phases
with flows of less than 16,000 m3 s-1 which occur
from April to September (Neiff 1990). The middle
stretch of the river is located in a subtropical region. It
extends from its confluence with the Paraguay River
(27° 290 S; 58° 500 W) to the city of Diamante
(Argentina; 32° 40 S; 60° 320 W). It is fringed along
its right bank by a 6–40 km-wide floodplain (13,000
km2), comprising thousands of permanent and tem-
porary waterbodies (Paira and Drago 2007). The
geomorphology of the floodplain is characterized by
channels and flood deposits (Drago 1971). The former
is represented by geomorphological units such as
sandbanks, meander plains and spills, while the latter
are found in plains of hindered drainage (Iriondo and
Drago 1972). In the Parana fluvial system, the effects
of pulse regime on the vegetation characteristics is
conditioned by the topographic position and has been a
relevant aspect addressing the relationship between
vegetation and pulse regime (Casco et al. 2010;
Marchetti and Aceñolaza 2011, 2012). The
Wetlands Ecol Manage
Fig. 1 a Map of the Parana River system (Parana Medio River
indicated). b Paddocks of the wetlands and location of upland
and lowland exclosures (white cuadrants). c Span (in days) of
hydrological regime of this river is characterized by
seasonal and successive phases of high and low water,
also known as ‘flood pulse’ (Junk et al. 1989) or ‘pulse
regime’ (Neiff 1996). The climate of the Parana River
floodplain is humid subtropical, with a mean annual
temperature of 19 °C. Rainfall ranges from 900 to
1000 mm per year, with 73% occurring mainly from
October to April (Rojas and Saluso 1987). Soils
consist of successive layers of sediment carried and
deposited by the river with each flood phase. Fluvial
soils correspond to Entisol group and Fluvent and
Aquent subgroups (Orellana and Bertoldi De Pomar
1969).
grazing periods (in grey) and resting periods (in white) for each
wetland. Days of sampling were indicated by circles. Date of
exclosure installation was also included
The study was conducted in three wetlands of the
Middle Parana River floodplain (Santa Fe city,
Argentina), included in different paddocks (Fig. 1):
the paddock W1 (318 410 0000 S, 608 310 2400 W) was
characterized by an area of 49.8 ha, a stocking rate of
361.4 AUD ha-1, long grazing period (90 days) and
short resting periods (37 days); the paddock of W2
(318 400 4500 S, 608 300 3300 W) had an area of 15 ha, a
stocking rate of 360 AUD ha-1, short grazing period
(27 days) and long resting period (97 days); whereas
the paddock W3 (318 400 9400 S, 608 300 6200 W)
comprised an area of 19.8 ha and remained without
cattle during the studied period (Fig. 1). These wet-
lands become connected with each other and with
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other aquatic systems only during high water season.
During this period, cattle were removed from these
floodplain systems. During the study, flood peak of
Parana river reached 4.5 m in the reference gauge
level Parana hydrological station, remaining at high
water levels on the floodplain (higher than 4 m) for a
period of 7 months (December 2015 to August 2016).
The present study does not have replication of the
grazing treatments, since replication is difficult to
achieve in large scale studies in floodplain systems.
Our results should be interpreted with caution when
generalizing about floodplain studies.
Sampling
Vegetation assemblages were classified according to
their topographical position: upland communities (less
floodable) and lowland communities (more floodable),
respectively (Marchetti and Aceñolaza 2012). Vege-
tation sampling was conducted monthly during
spring–summer seasons after a high water period.
High water period extended from January 2016 to July
2016 and reached a maximum value of 6.5 m of the
Parana River (Parana Port gauge scale).
Sampling was conducted from October 2016 to
March 2017 according to the movement of cattle
between paddocks (Fig. 1c): before the introduction of
cattle to a paddock and after their movement to
another paddock. Grazing regime and rotation of cattle
between wetlands did not follow a fixed schedule, it
was determined according to forage supply and plant
growth. All wetlands were sampled during the same
week when cattle movement occurred in any one
wetland. Exclosures were installed on September 2015
in the three paddocks including both upland and
lowland vegetation, in order to function as exclusion
sites (inside the exclosure) and grazed sites (outside
the exclosure) (Fig. 1c). After the installation of the
enclosures, from September 2015 to December 2015
and after the high water period, the three studied
wetlands were subjected to cattle grazing. Continuous
exclosures (length = 40 m, width = 20 m, area = 800
m2) including both vegetation types were installed in
W2 and W3 (Fig. 1b). Due to the great extension of the
lowland zone in W1, this exclosure was divided
(length = 20 m, width = 20 m, area = 400 m2, for
each), including one vegetation type within each
division (Fig. 1b).
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Wetlands Ecol Manage
The minimal area of each sample unit was deter-
mined by constructing a species-area curve for each
vegetation sample (Chytrý and Otýpková 2003). The
curve was calculated in the field within the surface
occupied by each homogeneous vegetation unit. The
species-area curve was determined to ignore counts of
species from other vegetation units (Perelman et al.
2005). For assessing vegetation cover, four quadrats
(4 m2) were chosen randomly in upland and lowland
vegetation areas of the exclusion sites and grazed sites
of each wetland (n = 16). Quadrats of grazed sites
were randomly arranged next to an exclosure. Plant
identification was done in the field; species that could
not be identified in the field were collected for
taxonomic determination. Identifications were made
at the lowest taxonomic level possible (mostly
species) using the available keys (Burkart
1969, 1974, 1979,1987; Burkart and Bacigalupo
2005; Pensiero et al. 2005). Plants were classified as
palatable and unpalatable, following previous studies
conducted in the wetlands of the Parana River (Rossi
2013; Rossi et al. 2014). Direct observation of signs of
browsing in leaves and shoots were also taken into
consideration for classification of palatable species.
Data analyses
Plants were classified into functional groups using
growth habit and life form: palatable (Pal), unpalat-
able (UPal) shrub/subshrub (Shr), forb (Forb), pros-
trate (Pros), erect (Ere) and graminoids (Gr) (Table 3).
Species diversity metrics were calculated from cover
data including exclusion and grazed sites of upland
and lowland vegetation to compare wetlands with
different resting and grazing periods. Differences in
total richness (R), total diversity (Shannon–Wiener
index, H’) total dominance (Simpson index, D)
(‘‘total’’ including all plots, and richness, diversity
and dominance of functional groups were assessed
using two factors ANOVA (R core team, 2017). The
factors were the exclosure (exclusion and grazed sites)
and the wetlands (W1, W2 and W3). Sampled sizes
ranged from 67 to 71 for exclosure and from 18 to 31
for wetlands (Table 1). In order to assess differences
between exclusion and grazed sites, data of the
different sampling dates (total richness, diversity and
dominance and functional groups) were pooled. Prin-
cipal Component Analysis (PCA) (CANOCO v.4, ter
Braak and Smilauer 1998) with cover data of upland
Wetlands Ecol Manage
Table 1 Sampled sizes (n) of the two factors ANOVA relative
to upland and lowland vegetation of W1 (long grazing/short
resting period), W2 (short grazing/long rest period) and W3
(without cattle during the studied period), and considering
exclusion sites (inside the exclosure) and grazed sites (outside
de exclosure)
Upland Lowland
W1 24 24
W2 31 29
W3 20 18
Exclusion sites 68 68
Grazed sites 71 67
and lowland vegetation was performed in order to
identify major sources of variation in vegetation cover
among wetlands. Prior to analysis, data were arcsin
transformed. The Indicator Species Analysis (R pro-
gram, Dufrêne and Legendre 1997) was used to
identify the taxa most strongly related to exclusion and
grazing sites.
Results
Sixty-eight native species were identified in the three
studied wetlands (Table 3). These species belonged to
31 botanical families, being the most representative
Poaceae (12 species), Asteraceae (10 species), Faba-
ceae (5 species) and Solanaceae (5 species). Palatable,
forb and prostrate species were dominant among both
upland and lowland vegetation (relative cover [
50%), while unpalatable, shrub and erect species
showed lower values of vegetation cover (relative
cover \ 40%). Grazed leaves and shoots of Ludwigia
peploides were observed, leading to its classification
as palatable species. Dominant species in upland and
lowland sites of the three wetlands were Cynodon
dactylon, L. peploides, Persicaria punctata and
Echinochloa crus-pavonis (relative cover [ 10%)
(Table 2).
Comparison between wetlands
and exclusion/grazed sites
Total richness and total diversity of upland vegetation
were significantly higher in W3 (without cattle) and in
exclusion sites (ANOVA, P \ 0.01), whereas
dominance was higher in W1 (short resting/long
grazing period) and W2 (long resting/short grazing
period) in comparison with W3, and in grazed sites
(ANOVA, P \ 0.01) (Fig. 2). Regarding lowland
vegetation, total richness and total diversity were
higher in W1 than in W2 and W3, whereas dominance
was lower in W3 (ANOVA, P \ 0.05) (Fig. 2). We
found no significant differences between exclusion
and grazed sites in terms of richness, diversity and
dominance in lowland communities (ANOVA,
P [ 0.05) (Fig. 2).
In relation to upland vegetation, richness and
diversity of palatable, unpalatable, shrub/subshrub,
forb and prostrate species were significantly higher in
W3 with respect to the grazed wetlands (W1 and W2)
(ANOVA, P \ 0.01), whereas dominance of palat-
able, forb and prostrate vegetation were higher in W1
and W2 (ANOVA, P \ 0.01) (Fig. 3a). Richness and
diversity of unpalatable, shrub, erect and graminoid
vegetation were significantly higher in exclusion sites
(Fig. 4a), whereas dominance of erect and graminoid
plants were higher in grazed sites (Fig. 4a). In relation
to lowland vegetation, richness and diversity of
palatable, unpalatable, shrub/subshrub, forb, prostrate
and erect species were significantly higher in W1
(ANOVA, P \ 0.001) (Fig. 3b). Dominance of palat-
able, forb and erect vegetation were significantly
higher in W3, whereas dominance of unpalatable and
shrub/subshrub vegetation were higher in W1 and W2
(ANOVA, P \ 0.001) (Fig. 3b). Richness and diver-
sity of forb and prostrate lowland vegetation were
significantly higher in W1, whereas dominance of forb
vegetation was higher in W3 (ANOVA, P \ 0.05)
(Fig. 4b).
Ordination analysis
The PCA ordination of upland vegetation sorted the
paddocks with different grazing and resting periods
along Axis 1. In the positive side of this axis, L.
peploides and Poa lanigera characterized W1 (short
resting/long grazing period), whereas C. dactylon was
associated with W2 (long resting/short grazing period)
on the negative side. Along axis 2, samples of W2 and
Setaria parviflora, P. punctata and Leersia hexandra
species characterized the positive side, whereas L.
peploides and P. lanigera were associated to the
negative side of this axis (Fig. 5a). A group of 24
species situated at the center of this ordination were
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Table 2 Dominant upland and lowland species (relative cover [ 10%) in exclusion and grazed sites relative to W1 (long grazing/
short resting period), W2 (short grazing/long rest period) and W3 (without cattle during the studied period)
Upland
W1
Exclusion Poa lanı´gera (12.7%); Cynodon dactylon (10.9%);
Ludwigia peploides (10.7%)
Grazing L. peploides (20.2%); P. lanı´gera (18.1%); C. dactylon
(10.2%)
W2
Exclusion P. punctata (19.2%)
Grazing C. dactylon (28.2%); P. punctata (13.1%)
W3
Exclusion L. peploides (19.4%); C. dactylon (17.5%)
Grazing C. dactylon (14.5%); L. peploides (12.3%)
common for the three wetlands (Fig. 5a). Concerning
lowland vegetation (Fig. 5b), L. peploides and Myrio-
phyllum aquatiqum related with W1 characterized the
positive side of axis 1, whereas P. punctata was related
to the negative side of this axis. In the positive side of
axis 2, E. crus-pavonis characterized samples of W2,
whereas Hymenachne amplexicaulis and Sesbania
virgata characterized the negative side of this axis.
Indicator species analysis
Regarding upland vegetation, Panicum prionitis,
Cyperus entrerianus, Cissus palmata and
Sapium haematospermum were the most strongly
related to grazed sites, while Mimosa tweedieana, E.
crus-pavonis, Muehlenbeckia sagittifolia, Myriophyl-
lum aquaticum and Cayaponia podantha were related
to exclusion sites (Table 4). In relation to lowland
vegetation, S. parviflora, P. punctata, Ambrosia
artemisiifolia, C. entrerianus and L. hexandra were
associated to grazed sites and Solanum glaucophyllum
and L. peploides were related to exclusion sites
(Table 4).
Discussion
Our results demonstrated that rotational cattle grazing
had a significant effect on the composition of upland
vegetation of the studied floodplain wetlands,
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Wetlands Ecol Manage
Lowland
Sesbania virgata (14.7%); Persicaria punctata (12.7%);
Echinochloa crus-pavonis (10%)
P. punctata (24.6%); L. peploides (15.6%); S. virgata
(10.8%)
P. punctata (39.7%); L. peploides (18.7%)
P. punctata (48.8%); E. crus-pavonis (16%)
P. punctata (36.4%); E. crus-pavonis (22.1%); L. peploides
(20.5%)
L. peploides (25.7%); P. punctata (22%); E. crus-pavonis
(13%)
decreasing total richness and diversity, and increasing
the dominance of palatable, forb and prostrate species.
Likewise, in contrast with our first hypothesis, our
results demonstrated that vegetation parameters did
not differ as grazing and resting time period differed.
Both the wetland with short resting/long grazing
period (W1) as well as the wetland with long resting/
short grazing period (W2) had lower diversity and
richness in comparison with the wetland without cattle
grazing (W3). Indeed, paddocks W1 and W2 subjected
to different resting and grazing periods did not differ in
richness and diversity of upland vegetation. W1 was
subjected to a short rest periods and long grazing
periods, but its greater area, heterogeneity and number
of patches of vegetation may have buffered the effect
of cattle. On the other hand, the stocking rate of W2
could have been compensated by the shorter grazing
and rest periods, diminishing cattle effect (Sterk et al.
2013).
In relation to the second hypothesis, grazing
produced a simplification of upland species of W1
and W2, with a dominance of palatable and prostrate
species with the ability for regrowth after damage (e.g.
Cynodon dactylon). Negative relationships between
species richness and dominance in grazing systems are
common in grassland (McNaughton 1983; Koerner
et al. 2014, 2018) and wetland ecosystems (Tanner
1992; Anderson and Calov 1996; Touzard and
Klement 2001; Rebergen 2002; Hornung and Rice
2003). This relation was also observed in different
Wetlands Ecol Manage
Fig. 2 Total richness, Shannon diversity and dominance index
of upland and lowland vegetation between (a) wetlands, W1
(black, long grazing/short resting period), W2 (grey, short
grazing/long rest period), W3 (white, without cattle during the
wetlands and mesic grasslands of East Africa and
America, where cattle generate extensive grazing
lawns in which a few grazing-tolerant grasses with-
stand high densities of large herbivores (McNaughton
1983; Jacobo et al. 2006; Diaz et al. 2007; Jones et al.
2011). In contrast, several studies of grassland
ecosystems (Collins et al. 1998; Hayes and Holl
2003; Koerner et al. 2014) and wetlands of North
America (e.g., Haines 1995; Jutila 1999; Rebergen
2002; Marty 2015), New Zealand (Buxton et al. 2001;
studied period); and (b) exclosure (white) and grazed (black)
sites. Significant differences of ANOVA test (P \ 0.05) are
show with different letters: ‘‘a’’, ‘‘b’’
Middleton 2002; Miller and Wells 2003) Africa
(Dahwa et al. 2014) and Australia (Mavromihalis
et al. 2013) showed that grazing reduced the domi-
nance of palatable species, with positive effects on
diversity and richness of upland vegetation.
Regarding W1 and W2, in both grazed wetlands,
palatable, forbs and prostrate species were detected as
dominant species. These species were r-strategists,
and are able to reproduce by vegetative adaptations as
rhizomes and stolons, with positive response, to
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Fig. 3 Mean values (± SD) of richness, Shannon diversity and
dominance index of each functional groups of upland (a) and
lowland (b) vegetation of W1 (black, long grazing/short rest
period), W2 (grey, short grazing/long rest periods) and W3
(white, without cattle grazing during the studied period).
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Wetlands Ecol Manage
References: palatable (Pal), unpalatable (UPal), shrub/subshrub
(Shr), forb (Forb), prostrate (Pros), erect (Ere) and graminoids
(Gr). Significant differences of ANOVA test (P \ 0.05) are
show with different letters: ‘‘a’’, ‘‘b’’
Wetlands Ecol Manage

Fig. 4 Mean values (± SD) of richness, Shannon diversity and shrub/subshrub (Shr), forb (Forb), prostrate (Pros), erect (Ere)
dominance index of each functional group of upland (a) and and graminoids (Gr). Significant differences of ANOVA test
lowland (b) vegetation relative to exclosure (white) and grazed (P \ 0.05) are show with different letters: ‘‘a’’, ‘‘b’’
sites (black). References: palatable (Pal), unpalatable (UPal),

grazing (Diaz et al. 2007). In particular, palatable and and palatable species such Panicum prionitis, Cyperus
prostrate species such as C. dactylon and L. hexandra entrerianus, S. parviflora are generally associated to

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Fig. 5 Principal Components Analysis ordination of upland
(a) and lowland (b) vegetation. Sites ordination: W1 (black,
long grazing periods and short resting periods), W2 (grey, short
highly disturbed and overgrazed environments (Alte-
sor et al. 2005; Gutierrez 2014; Quintana et al. 2014;
Magnano 2017). These strategies would avoid defo-
liation of the aboveground biomass and allow species
to quickly colonize bare ground (Dong and de Kroon
1994). Palatable species registered in this study were
the same such as those found in studies performed in
the lower Parana River (Rossi 2013; Magnano 2017).
An exception is Ludwigia peploides which, although it
may not be preferred by cattle, it could serve as an
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Wetlands Ecol Manage
grazing periods and longer rest periods) and W3 (white, without
cattle); species ordination. References of species were shown in
Table 3
alternative resource when more palatable species are
scarce (Massa et al. 2018).
According to our third hypothesis, the result of this
work showed that grazing exclusion favoured the
colonization, regeneration and survival of upland
species, increasing richness and diversity of unpalat-
able, shrub, erect and graminoid species. Others
studies have shown the positive effects of cattle
exclusion on vegetation attributes (Cooper and
McCann 2011; Shaltout et al. 1996; Fujita et al. 2009;
Wetlands Ecol Manage

Table 3 Species list relative to upland (U) and lowland grazing/long rest period; W3: without cattle during the studied
(L) communities identified in the three studied wetlands (W1: period), status (N: native) and characterization of functional
long grazing period/short resting period; W2: short groups (X)
Family Species Abbrev Status Type of Functional groups
vegetation
For NFor Shr Forb Pros Ere Gr

Alismataceae Sagittaria montevidensis Cham. & Sagi N L X X X

Schltdl
Amaranthaceae Alternanthera philoxeroides (Mart.) Alte N U–L X X X
Griseb
Apiaceae Cyclospermum leptophyllum (Pers.) Cyclo N U X X X
Sprague
Araliaceae Hydrocotyle bonariensis Comm. ex Hydro N U X X X
Lam
Asteraceae Ambrosia artemisiifolia L AmbA N U–L X X X
Aspilia silphioides (Hook. & Arn.) Aspi N U X X X
Benth. ex Benth. & Hook.f
Bidens laevis (L.) ‘‘Britton, Sterns & Bi N U–L X X X
Poggenb
Eclipta prostrata (L.) L Ecli N U X X X
Enydra anagallis Gardner Eny N U–L X X X
Eupatorium hecatanthum (DC.) Baker Eupa N U X X X
Gymnocoronis spilanthoides (D.Don Gym N U–L X X X
ex Hook. & Arn.) DC
Melanthera latifolia (Gardner) Mela N U X X X
Cabrera
Mikania periplocifolia Hook. & Arn MikP N U X X X
Vernonanthura lucida (Less.) H.Rob Ver N U X X X X
Brassicaceae Lepidium bonariense L Lepi N U X X X
Cleomaceae Cleome houtteana Schltdl Cleo N U X X X
Commelinaceae Commelina erecta L Com N U X X X
Convolvulaceae Aniseia argentina (N.E. Br.) O’Donell Ani N U–L X X X
Cuscuta sp. Cus N L X X X
Ipomoea nil (L.) Roth Ipo N U X X X
Cucurbitaceae Cayaponia podantha Cogn Caya N U X X X
Cyperaceae Cyperus entrerianus Boeckeler CypE N U–L X X X
Cyperus odoratus L CypO N L X X X
Cyperus pohlii (Nees) Steud CypP N L X X X
Isolepis cernua (Vahl) Roem. & Iso N L X X X
Schult
Euphorbiaceae Sapium haematospermum Müll.Arg Sapi N U X X X
Caperonia castaneifolia (L.) A.St.-Hil Cape N L X X X
Fabaceae Lathyrus nigrivalvis Burkart Lat N U X X X
Mimosa pigra L MPi N U X X X
Mimosa tweedieana Barneby ex MTw N U–L X X X
Glazier & Mackinder
Senna pendula (Willd.) H.S.Irwin & Sen N U X X X
Barneby
Sesbania virgata (Cav.) Pers Ses N U–L X X X

123
 Wetlands Ecol Manage

Table 3 continued

Family Species Abbrev Status Type of Functional groups

vegetation
For NFor Shr Forb Pros Ere Gr

Haloragaceae Myriophyllum aquaticum (Vell.) Myri N U–L X X X

Verdc
Lamiaceae Salvia pallida Benth Salv N U X X X
Malpighiaceae Stigmaphyllon calcaratum N.E. Br Sti N U X X X
Malvaceae Hibiscus striatus Cav Hibi N U X X X
Modiolastrum malvifolium (Griseb.) Modio N U X X X
K. Schum
Marantaceae Thalia geniculata L Tha N L X X X
Onagraceae Ludwigia peploides (Kunth) Lud N U–L X X X
P.H.Raven
Oxalidaceae Oxalis corniculata L OxC N U X X X
Passifloraceae Passiflora misera Kunth Passi N U X X X
Plantaginaceae Plantago myosuros Lam Plan N U X X X
Poaceae Cynodon dactylon (L.) Pers Cyn N U–L X X X X
Echinochloa crus-pavonis (Kunth) EchiCP N U–L X X X X
Schult
Echinochloa sp. Echi N U–L X X X X
Hymenachne amplexicaulis (Rudge) Hym N L X X X X
Nees
Leersia hexandra Sw Lee N U–L X X X X
Panicum elephantipes Nees ex Trin Pele N U–L X X X X
Panicum prionitis Nees PPri N U X X X X
Paspalum acuminatum Raddi PasA N U–L X X X X
Paspalum notatum Flüggé PasN N L X X X X
Phalaris angusta Nees ex Trin Pha N U–L X X X X
Poa lanigera Nees Poa N U X X X X
Setaria parviflora (Poir.) SetP N U–L X X X X
M.Kerguelen
Polygonaceae Muehlenbeckia sagittifolia (Ortega) Mueh N U X X X
Meisn
Persicaria punctata (Elliott) Small Per N U–L X X X
Polygonum paraguayense Wedd Pol N L X X X
Rumex argentinus Rech. f RuA N U X X X
Pontederiaceae Eichhornia crassipes (Mart.) Solms Eich N L X X X
Rubiaceae Spermacoce verticillata L Sper N U X X X X
Sapindaceae Cardiospermum halicacabum L Cardio N U X X X
Solanaceae Jaborosa integrifolia Lam Jabo N U X X X
Salpichroa origanifolia (Lam.) Baill Salpi N U X X X X
Solanum amygdalifolium Steud SAmy N U X X X
Solanum glaucophyllum Desf SGla N U–L X X X
Solanum nigrescens M. Martens & Snig N U X X X
Galeotti
Verbenaceae Lippia alba (Mill.) N.E.Br. ex Lip N U X X X
Britton & P.Wilson
Vitaceae Cissus palmata Poir Ciss N U X X X

For forage species, NFor non-forage species, Shr shrub/subshrub, Forb forb, Pros prostrate, Ere erect, Gr graminoid

123
Wetlands Ecol Manage

Minchinton et al. 2019). Some of the direct impacts of Table 4 Significant results of Indicator Species Analysis.
cattle grazing (trampling, urine) could restrict the Species and P values for exclusion (P Exclusion) and grazed
(P Grazed) sites are provided
establishment and survival of certain sensitive unpalat-
able upland species (Champion et al. 2001; Becerra P Exclusion P Grazed
2006; Liu et al. 2015). The improvement in soil
Upland
conditions (temperature, moisture, nutrient cycling) in
Panicum prionitis 0.009
excluded sites could favour the regeneration and
Cyperus entrerianus 0.0007
development of unpalatable plants, perhaps more sen-
Cissus palmata 0.008
sitive to cattle impact than palatable species (Yates et al.
Sapium haematospermum 0.02
2000; Pei et al. 2008; Rossi et al. 2014). Among
Mimosa tweedieana 0.0001
unpalatable species, Mimosa tweedieana, Muehlen-
Echinochloa crus-pavonis 0.0007
beckia sagittifolia, Myriophyllum aquaticum, Cayapo-
Muehlenbeckia sagittifolia 0.01
nia podantha, Ambrosia artemisiifolia and Solanum
glaucophyllum were found as indicators of grazed Myriophyllum aquaticum 0.02
excluded sites in floodplain wetlands of the Parana Cayaponia podantha 0.02
River. Lowland
In lower topographical zones such as lowland S. parviflora 0.0005
ecosystems, the duration of flooding would represent P. punctata 0.01
the main factor driving the resilience and composition Ambrosia artemisiifolia 0.02
of lowland vegetation (Franceschi et al. 2010; L. hexandra 0.01
Marchetti el al. 2013). In lowlands, species adapted L. peploides 0.04
to waterlogged conditions were dominant (i.e. P. Solanum glaucophyllum 0.04
punctata, L. peploides, Sesbania virgata and E. crus-
pavonis) (Malvárez 1997; Kandus et al. 2010; Rossi
et al. 2014). The dominance of P. punctata in the
Further studies are necessary to deepen the under-
studied wetlands would be related to its ability to
standing of cattle effects on floodplain wetlands.
survive in disturbed sites (Rossi et al. 2014), deter-
mining its high cover and dominance in wetlands
subjected to cattle grazing. In these wetlands, cattle
Conclusion
preferred the less floodable plant community, with
significant effects on upland vegetation. Indeed, in the
The fact that wetlands with different resting/grazing
studied wetlands, cattle had a significant effect on
period did not differ in vegetation parameters suggest
water quality, incorporating organic and dissolved
that longer resting period could be necessary to ensure
nutrients by manure and urine (Mesa et al. 2015). The
the recovery of the vegetation community. In this
increase in nutrient concentration along with the
context, further research including manipulative field
floods, which can supply new species to the floodplain
experiments could be very valuable to determine the
through the import of propagules, would represent
optimum resting period for plant community recovery.
factors with significant effects on diversity and
Cattle management is a very important issue to
richness of lowland vegetation, as was observed in
study given the change in land use in floodplain
W1. This paddock had a more extensive and hetero-
systems of the Middle Parana. Moreover, at a world-
geneous lowland ecosystem that offers a high number
wide scale there is an urgent need to develop
of ecological niches for species colonization.
sustainable cattle production with the least possible
Although the present study does not have replica-
impact. Grazing management plans and subsequent
tion of the grazing treatments, the results of this work
operational decisions should include both economic
give significant information of the impact of cattle on
and biological considerations. The simplification of
vegetation of wetlands of the Middle Parana River
wetland ecosystems by grazing needed to be assessed
floodplain. Our results should be interpreted with
in order to reverse the loss of function and benefit from
caution when generalizing about floodplain studies.
the services they provide. Integrating protection by

123

excluding cattle in some floodplain systems could be
considered an effective strategy to restore native
vegetation, maintaining the heterogeneity and diver-
sity of the wetlands.
Acknowledgements This research was supported by the
Consejo Nacional de Investigaciones Cientı́ficas y Técnicas
(CONICET), and Agencia de Promoción Cientı́fica y
Tecnológica (PICT 2017-2982; PICT 2018-02203). We would
also like to thank Luciana Montalto, Diana Alberto, Gisela
Mayora and Ernesto Massa for help with field sampling and
laboratory analysis.
Funding This study was funded by Consejo Nacional de
Investigaciones Cientı́ficas y Técnicas (CONICET), and
Agencia de Promoción Cientı́fica y Tecnológica: PICT 2017-
2982, PICT 2018-02203.
References
Altesor A, Oesterheld M, Leoni E, Lezama F, Rodrı́guez C
(2005) Effect of grazing on community structure and pro-
ductivity of a Uruguayan grassland. Plant Ecol 179:83–91
Anderson UV, Calov B (1996) Long term effects of sheep
grazing on giant hogweed (Hercaleum mantegazzianum).
Hydrobiologia 340:277–284
Andrew SM, Totland Ø, Moe SR (2015) Spatial variation in
plant species richness and diversity along human distur-
bance and environmental gradients in a tropical wetland.
Wetl Ecol Manag 23:395–404
Baigún CRM, Puig A, Minotti PG, Kandus P, Quintana RD,
Vicari R, Bó RF, Oldani NO, Nestler JA (2008) Resource
use in the Parana river delta (Argentina): moving away
from an ecohydrological approach? Ecohydrol Hydrobiol
8:245–262
Becerra JAB (2006) Grazing Intensity, plant diversity, and
rangeland. In: Spehn EM, Liberman M, Körner C (eds)
Land use change and mountain biodiversity. Tylor &
Francis Group, Londres, pp 153–166
Beetz AE, Rinehart L (1996) Rotational grazing. Attra.
Biasatti NR, Rozzatti JC, Fandiño B, Pautaso A, Mosso E,
Marteleur G et al (2016) Las ecoregiones, su conservación
y las áreas naturales protegidas de la provincia de Santa Fe.
Ministerio de Medio Ambiente, Santa Fe
Burkart A (1969) Flora Ilustrada de Entre Rı́os (Argentina).
Tomo VI, Parte 2 ed. Colecc. Ci. Inst. Nac. Tecnol. Ago-
pecu., Buenos Aires
Burkart A, (1974). Flora ilustrada de Entre Rı́os (Argentina)
(No. 581.98221 584.9). Instituto Nacional de Tecnologı́a
Agropecuaria (Argentina).
Burkart A. (1979). Flora ilustrada de Entre Rı́os (Argentina).
Parte V. Dicotiledóneas Metaclamı́deas. Generali-
dades;(Gamopétalas). Colección Cientı́fica del INTA.
Tomo VI-V. Bs. As.
Burkart A (1987) Flora ilustrada de entre rı́os (Argentina). Tomo
VI, Parte 3 ed. Colecc. Ci. Inst. Nac. Tecnol. Agopecu.,
Buenos Aires
123
Wetlands Ecol Manage
Burkart A, Bacigalupo N (2005) Flora ilustrada de entre rı́os
(Argentina). Tomo VI, Parte 4 ed. Colecc. Ci. Inst. Nac.
Tecnol. Agopecu., Buenos Aires
Buxton RP, Timmins SM, Burrows LE, Wardle P (2001) Impact
of cattle on department of conservation grazing leases in
south wetland: results from monitoring 1989–99, and rec-
ommendation. Sci Conserv 179:1–64
Casco SL, Neiff JJ, Poi de Neiff A (2010) Ecological responses
of two pioneer species to a hydrological connectivity gra-
dient in riparian forests of the lower Parana river. Plant
Ecol 209:167–177
Champion PD, Beadel SM, Dugdale TM (2001) Turf commu-
nities of lake whangape and experimental assessment of
some potential management techniques. Sci Conserv
186:1–51
Chytrý M, Otýpková Z (2003) Plot sizes used for phytosocio-
logical sampling of European vegetation. J Veg Sci
14:563–570
Collins SL, Knapp AK, Briggs JM, Blair JM, Steinauer EM
(1998) Modulation of diversity by grazing and mowing in
native tallgrass prairie. Science 280:745–747
Cooper A, McCann T (2011) Cattle exclosure and vegetation
dynamics in an ancient Irish wet oakwood. Plant Ecol
212:79–90
Dahwa E, Kativu S, Hungwe T, Mudzengi CP, Poshiwa X,
Murungweni C, Shoko MD (2014) Influence of grazing
intensity on plant community diversity, structure and
composition in Dambo wetlands of communal Zimbabwe.
GJBAHS 3:259–266
Diaz S, Lavorel S, McIntyre S, Falczuk V, Casanoves F,
Milchunas DG, Skarpel C, Rusch G, Sternberg M, Noy-
Meir I, Landsberg J, Zhang W, Clark H, Campbell BD
(2007) Plant trait responses to grazing–a global synthesis.
Glob Change Biol 13:313–341
Dong M, de Kroon H (1994) Plasticity in morphology and
biomass allocation in Cynodon dactylon, a grass species
forming stolons and rhizomes. Oikos 70:99–106
Drago E (1971) Génesis y evolución de la llanura alluvial del
Parana Medio y los cuerpos de agua contenidos en ella.
Instituto Nacional de Limnologı́a CNICT, Santo Tomé
inédito
Dufrêne M, Legendre P (1997) Species assemblages and indi-
cator species: the need for a flexible asymmetrical
approach. Ecol Monogr 67:345–366
Haines M (1995) The Mt Barker ephemeral tarns—A wetland
’’management experiment‘‘. Centre for Resource Man-
agement, Lincoln University. Report to Department of
Conservation.
Fleischner TL (1994) Ecological costs of livestock grazing in
western North America. Conserv Biol 8:629–644
Franceschi EA, Torres PS, Lewis JP (2010) Diversidad de la
vegetación durante su recuperación tras una creciente
extraordinaria del Rı́o Parana Medio (Argentina). Rev Biol
Trop 58:707–716
Fujita N, Amartuvshin N, Yamada Y, Matsui K, Sakai S,
Yamamura N (2009) Positive and negative effects of
livestock grazing on plant diversity of Mongolian nomadic
pasturelands along a slope with soil moisture gradient.
Grassl Sci 55:126–134
Wetlands Ecol Manage
Gutiérrez HF (2014) Botánica sistemática de las plantas con
semillas: principales familias de Gimnospermas y Mono-
cotiledóneas. Ediciones UNL, Santa Fe
Harrison S (1999) Native and alien species diversity at the local
and regional scales in a grazed California grassland.
Oecologia 121:99–106
Hayes GF, Holl KD (2003) Cattle grazing impacts on annual
forbs and vegetation composition of mesic grasslands in
California. Conserv Biol 17:1694–1702
Hornung JP, Rice CL (2003) Odonata and wetland quality in
Southern Alberta, Canada: a preliminary study. Odonato-
logica 32:119–129
Iriondo M, Drago E (1972) Descripción cuantitativa de dos
unidades geomorfológicas de la llanura aluvial del rı́o
Parana medio. Argentina Rev Asoc Geol Argent
27:143–160
Jackson RD, Allen-Diaz B (2006) Spring-fed wetland and
riparian plant communities respond differently to altered
grazing intensity. J Appl Ecol 43:485–498
Jacobo EJ, Rodrı́guez AM, Bartoloni N, Deregibus VA (2006)
Rotational grazing effects on rangeland vegetation at a
farm scale. Rangel Ecol Manag 59:249–257
Jones WM, Fraser LH, Curtis PJ (2011) Plant community
functional shifts in response to livestock grazing in inter-
mountain depressional wetlands in British Columbia.
Canada Biol Conserv 144:511–517
Junk WJ, Bayley PB, Sparks RE (1989) The flood pulse concept
in river-floodplain systems. Can Spec Publ Fish Aquat Sci
106:110–127
Jutila H (1999) Effect of grazing on the vegetation of shore
meadows along the Bothnian Sea Finland. Plant Ecol
140:77–88
Kandus P, Morandeira N, Schivo F (2010) Bienes y Servicios
Ecosistémicos de los Humedales del Delta del Parana.
Fundación Humedales/Wetlands International, Buenos
Aires
Koerner SE, Burkepile DE, Fynn RW, Burns CE, Eby S,
Govender N, Hagenah N, Matchett KJ, Thompson DI,
Wilcox KR, Collins SL, Kirkman KP, Knapp AK, Smith
MD (2014) Plant community response to loss of large
herbivores differs between North American and South
African savanna grasslands. Ecology 95:808–816
Koerner SE, Smith MD, Burkepile DE, Hanan NP, Avolio ML,
Collins SL et al (2018) Change in dominance determines
herbivore effects on plant biodiversity. Nat Ecol Evol
2:1925–1932
Kothmann MM (1984) Concepts and principles underlying
grazing systems: a discussant paper
Liu J, Feng C, Wang D, Wang L, Wilsey BJ, Zhong Z (2015)
Impacts of grazing by different large herbivores in grass-
land depend on plant species diversity. J Appl Ecol
52:1053–1062
Lodge KA, Tyler AC (2020) Divergent impact of grazing on
plant communities of created wetlands with varying
hydrology and antecedent land use. Wetl Ecol Manag
28:797–813
Lyseng MP, Bork EW, Hewins DB, Alexander MJ, Carlyle CN,
Chang SX, Willms WD (2018) Long-term grazing impacts
on vegetation diversity, composition, and exotic species
presence across an aridity gradient in northern temperate
grasslands. Plant Ecol 219:649–663
Maestas JD, Knight RL, Gilgert WC (2003) Biodiversity across
a rural land-use gradient. Conserv Biol 17:1425–1434
Magnano A (2017) Efectos de las actividades ganaderas sobre
un ecosistema de humedal del Delta del rı́o Parana. Tesis de
doctorado. Universidad Nacional de San Martin, Buenos
Aires
Magnano A, Vicari R, Astrada E, Quintana RD (2013) Gana-
derı́a en humedales: respuestas de la vegetación a la
exclusión del pastoreo en tres tipos de ambientes en un
paisaje del Delta del Parana. ASADEP 4:137–148
Malvárez I (1997) Las comunidades vegetales del Delta del Rı́o
Parana: su relación con factores ambientales y patrones del
paisaje. Tesis de doctorado. Universidad de Buenos Aires,
Buenos Aires
Marchetti ZY, Aceñolaza PG (2011) Vegetation communities
and their relationship with the pulse regime on islands of
the Middle Parana River, Argentina. Iheringia. Ser Bot
66:209–226
Marchetti ZY, Aceñolaza PG (2012) Pulse regime and vegeta-
tion communities in fluvial systems: the case of the Parana
River floodplain, Argentina. Flora 207:795–804
Marchetti ZY, Latrubesse EM, Pereira MS, Ramonell CG
(2013) Vegetation and its relationship with geomorpho-
logic units in the Parana River floodplain Argentina. J S
Am Earth Sci 46:122–136
Marty JT (2005) Effects of cattle grazing on diversity in
ephemeral wetlands. Conserv Biol 19:1626–1632
Marty JT (2015) Loss of biodiversity and hydrologic function in
seasonal wetlands persists over 10 years of livestock
grazing removal. Restor Ecol 23:548–554
Massa ES, González R., Benavidez H. (2018). Productividad de
un ‘‘verdolagal’’ (Ludwigia peploides) en islas del rı́o
Parana entre dos crecientes sucesivas. IV Congreso del
Mercosur sobre Manejo de Pastizales Naturales, La Rioja
Mavromihalis JA, Dorrough J, Clark SG, Turner V, Moxham C
(2013) Manipulating livestock grazing to enhance native
plant diversity and cover in native grasslands. Rangel J
35:95–108
McInnes RJ, Simpson M, Lopez B, Hawkins R, Shore R (2017)
Wetland ecosystem services and the ramsar convention: an
assessment of needs. Wetlands 37:123–134
McNaughton SJ (1983) Serengeti grassland ecology: the role of
composite environmental factors and contingency in
community organization. Ecol Monogr 53:291–320
Mesa L, Mayora G, Saigo M, Giri F (2015) Nutrient dynamics in
wetlands of the middle Parana River subjected to rotational
cattle management. Wetlands 35:1117–1125
Middleton BA (2002) The flood pulse concept in wetland
restoration. In: Middleton BA (ed) Flood pulsing in wet-
lands: Restoring the natural hydrological balance. Wiley,
New York, pp 1–10
Miller C, Wells A (2003) Cattle grazing and the regeneration of
totara (Podocarpus totara var. waihoensis) on river ter-
races, south Westland New Zealand. N Z J Ecol 27:37–44
Minchinton TE, Shuttleworth HT, Lathlean JA, McWilliam RA,
Daly TJ (2019) Impacts of cattle on the vegetation structure
of mangroves. Wetlands 39:1–9
Moges A, Beyene A, Triest L, Ambelu A, Kelbessa E (2018)
Imbalance of ecosystem services of wetlands and the per-
ception of the local community towards their restoration
123

and management in Jimma highlands, southwestern
Ethiopia. Wetlands 38:1081–1095
Moran J, Skeffington MS, Gormally M (2008) The influence of
hydrological regime and grazing management on the plant
communities of a karst wetland (Skealoghan turlough) in
Ireland. Appl Veg Sci 11:13–24
National Research Council (2001) Compensating for wetland
losses under the clean water act. National Academy Press,
Washington DC
Neiff JJ (1990) Ideas para la interpretación ecológica del Parana.
Interciencia 15:424–441
Neiff JJ (1996) Large rivers of South America: toward the new
approach. Vereinigung für theoretische und angewandte
Limnologie 6:167–180
Orellana, J. A., & Bertoldi De Pomar, H. (1969). Introducción al
estudio de los suelos isleños del Paraná Medio. Actas del a
5a RACS. Asociación Argentina de Ciencias del Suelo,
Santa Fe, 482–490
Painter EL, Belsky AJ (1993) Application of herbivore opti-
mization theory to rangelands of the western United States.
Ecol Appl 3:2–9
Paira AR, Drago EC (2007) Origin, evolution, and types of
floodplain water bodies. In: Iriondo MH, Paggi JC, Parma
MJ (eds) The middle Parana river: limnology of a sub-
tropical wetland. Springer, Berlin, pp 53–81
Pei S, Fu H, Wan C (2008) Changes in soil properties and
vegetation following exclosure and grazing in degraded
Alxa desert steppe of Inner Mongolia, China. Agric Eco-
syst Environ 124:33–39
Pensiero J, Gutierrez H, Luchetti A, Exner E, Kern V, Brnich E,
Oakley L, Prado D, Lewis J (2005) Flora vascular de la
provincia de Santa Fe. Claves para el reconocimiento de las
familias y géneros. Catálogo sistemático de las especies.
Ediciones UNL
Perelman SB, Batista WB, León RJC (2005) El estudio de la
heterogeneidad de la vegetacion, Fitosociologı́ay técnicas
relacionadas. In: Oesterheld M, Aguiar MR, Ghersa CM,
Paruelo JM (eds) La heterogeneidad de la vegetacion de los
agroecosistemas. Facultad de Agronomı́a, Buenos Aires,
pp 321–350
PROSAP (2009) Hacia un manejo integrado del Agua de Riego
en Argentina. Programa de Servicios Agrı́cola Provin-
ciales, Ministerio de Agricultura, Ganaderı́a y Pesca,
Buenos Aires
Quintana RD, Bó RF, Astrada E, Reeves C (2014) Lineamientos
para una ganaderı́a ambientalmente sustentable en el Delta
del Parana. Fundación para la Conservación y el Uso
Sustentable de los Humedales, Buenos Aires
Rahmanian S, Hejda M, Ejtehadi H, Farzam M, Memariani F,
Pyšek P (2019) Effects of livestock grazing on soil, plant
functional diversity, and ecological traits vary between
regions with different climates in northeastern Iran. Ecol
Evol 9:8225–8237
Raymond KL, Vondracek B (2011) Relationships among rota-
tional and conventional grazing systems, stream channels,
and macroinvertebrates. Hydrobiologia 669:105–117
R Core Team (2017) R: A language and environment for sta-
tistical computing. https://www.R-project.org/
Rebergen A (2002) Survey, monitoring and management of the
native grass Amphibromus fluitans, especially in relation to
123
Wetlands Ecol Manage
the exotic grass Paspalum distichum (Mercer grass) at
Lake Wairarapa wetlands. Informe inédito. Department of
Conservation, Wairarapa
Reeves PN, Champion PD (2004) Effects of livestock grazing
on wetlands: literature review. NIWA, Hamilton
Rojas AE, Saluso JH (1987) Informe climático de la Provincia
de Entre Rı́os. INTA, Parana
Rossi C (2013) Composición florı́stica y caracterización nutri-
cional de las especies forrajeras en el sistema silvopastoril
del Delta del Parana. Tesis doctoral. Universidad Nacional
de Cordoba.
Rossi CA, De Magistris AA, González GL, Carou NE, De Loof
EP (2014) Plantas de interés ganadero de la región del Bajo
Delta del Parana (Argentina). Editorial UNLZ, Buenos
Aires
Shaltout KH, El Halawany EF, El Kady HF (1996) Conse-
quences of protection from grazing on diversity and
abundance of the coastal lowland vegetation in Eastern
Saudi Arabia. Biodivers Conserv 5:27–36
Sonnier G, Quintana-Ascencio PF, Bohlen PJ, Fauth JE, Jenkins
DG, Boughton EH (2020) Pasture management, grazing,
and fire interact to determine wetland provisioning in a
subtropical agroecosystem. Ecosphere 11:e03209
Sovell LL, Vondracek AB, Frost JA, Mumford KG (2000)
Impacts of rotational grazing and riparian buffers on
physicochemical and biological characteristics of South-
eastern Minnesota, USA, streams. Environ Manag
26:629–641
Sterk M, Gort G, Klimkowska A, Van Ruijven J, Van Teeffelen
AJA, Wamelink GWW (2013) Assess ecosystem resi-
lience: linking response and effect traits to environmental
variability. Ecol Indic 30:21–27
Tanner CC (1992) A review of cattle grazing effects on lake
margin vegetation with observations from dune lakes in
Northland, New Zealand. N Z Nat Sci 19:1–14
Teague WR, Dowhower SL, Baker SA, Ansley RJ, Kreuter UP,
Conover DM, Waggoner JA (2010) Soil and herbaceous
plant responses to summer patch burns under continuous
and rotational grazing. Agric Ecosyst Environ
137:113–123
TEEB (2013) The economics of ecosystems and biodiversity for
water and wetlands. Institute for European Environmental
Policy (IEEP) & Ramsar Secretariat, London and Brussels
terBraak CJ, Smilauer P (1998) CANOCO reference manual and
user’s guide to canoco for windows: software for canoni-
cal: software for canonical community ordination (version
4). Microcomputer Power, Ithaca
Touzard B, Clement B (2001) Plant diversity dynamics in an
eutrophic alluvial reed bed after experimental small-scale
disturbances. Bot Helv 111:45–58
Yates CJ, Norton DA, Hobbs RJ (2000) Grazing effects on plant
cover, soil and microclimate in fragmented woodlands in
south-western Australia: implications for restoration.
Austral Ecol 25:36–47
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