Wetlands Ecol Manage

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

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modify the composition, functional diversity and vegetation composition, have pointed towards a need
dynamics of wetland plant communities, increase the for identification of proper management in these areas.
proportion of unpalatable biomass for cattle, facilitate Consequently, the aims of this study were (1) to assess
plant invasions, and alter water and nutrient cycles the effect of different resting/grazing periods on the
(Jackson and Allen-Diaz 2006; Lyseng et al. 2018; composition, richness, diversity and dominance of
Sonnier et al. 2020). Through these effects, grazing functional groups of wetlands vegetation assemblages
has negatively affected biodiversity in some ecosys- and (2) to analyze the influence of cattle exclusion on
tems, impairing erect plants and reducing woody these parameters. We hypothesized that the vegetation
species cover (Painter and Belsky 1993; Fleischner of the wetland with longer grazing and shorter resting
1994; Jones et al. 2011; Magnano et al. 2013; Andrew periods have lower diversity and richness. Moreover,
et al. 2015; Rahmanian et al. 2019). Conversely, in in this wetland, we expect a dominance of forb and
other systems, livestock can also serve as a functional prostrate vegetation. Finally, we predict the exclusion
equivalent of large native herbivores and play an of cattle promotes vegetation recovery increasing total
essential role maintaining biodiversity, increasing richness, diversity and diversity of non-forage, shrub
native species richness and favoring forage species and erect species.
(Harrison 1999; Maestas et al. 2003; Hayes and Holl
2003; Marty 2005; Jacobo et al. 2006; Lodge and
Tyler 2020). Materials and methods
In Argentina, since the late twentieth century, the
expansion of soybean production has forced the Study area
relocation of cattle into marginal sites for agriculture
such as wetlands, constituting a threat to these The Parana River drains the second largest watershed
ecosystems (PROSAP 2009). This situation has in South America, after that of the Amazon, covering
increased the livestock density in floodplain systems, an area of 3.1 9 106 km2. Most of its course is flanked
about 100% in one decade (Quintana et al. 2014). by a 10–50 km wide floodplain, occupying an area of
The Middle Parana River in Argentina is one of the 60,000 km2 (Fig. 1a). The hydrological dynamics of
most important wetland ecosystems in South America the Parana River are characterized by high water
because of its location and extent, which is being phases with flows of up to 25,000 m3 s-1 which take
modified quite rapidly (Baigún et al. 2008). Because of place from October to March, and by low water phases
their extensive areas of vegetation for forage and high- with flows of less than 16,000 m3 s-1 which occur
quality water supply, the wetlands of the Middle from April to September (Neiff 1990). The middle
Parana River system provide excellent conditions for stretch of the river is located in a subtropical region. It
cattle (Mesa et al. 2015). In these systems, cattle extends from its confluence with the Paraguay River
historically grazed wetlands and play an essential role (27° 290 S; 58° 500 W) to the city of Diamante
in vegetation composition (Burkart 1969; Biasatti (Argentina; 32° 40 S; 60° 320 W). It is fringed along
et al. 2016). Recently, grazing managers have begun to its right bank by a 6–40 km-wide floodplain (13,000
implement rotational grazing systems in this zone to km2), comprising thousands of permanent and tem-
control the frequency and intensity of grazing and rest porary waterbodies (Paira and Drago 2007). The
periods in order to optimize livestock and plant geomorphology of the floodplain is characterized by
performance (Beetz and Rinehart 1996; Teague et al. channels and flood deposits (Drago 1971). The former
2010; Raymond and Vondracek 2011). This strategy is represented by geomorphological units such as
improves and maintains forage resource and increase sandbanks, meander plains and spills, while the latter
animal production (Sovell et al. 2000). Cattle are are found in plains of hindered drainage (Iriondo and
continually faced with younger and more nutritious Drago 1972). In the Parana fluvial system, the effects
forage throughout the growing season (Kothmann of pulse regime on the vegetation characteristics is
1984). conditioned by the topographic position and has been a
The significant increase of cattle stocking rates in relevant aspect addressing the relationship between
floodplain systems of the Middle Parana River, added vegetation and pulse regime (Casco et al. 2010;
to the limited understanding of its effects on the Marchetti and Aceñolaza 2011, 2012). The

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Fig. 1 a Map of the Parana River system (Parana Medio River grazing periods (in grey) and resting periods (in white) for each
indicated). b Paddocks of the wetlands and location of upland wetland. Days of sampling were indicated by circles. Date of
and lowland exclosures (white cuadrants). c Span (in days) of exclosure installation was also included

hydrological regime of this river is characterized by The study was conducted in three wetlands of the
seasonal and successive phases of high and low water, Middle Parana River floodplain (Santa Fe city,
also known as ‘flood pulse’ (Junk et al. 1989) or ‘pulse Argentina), included in different paddocks (Fig. 1):
regime’ (Neiff 1996). The climate of the Parana River the paddock W1 (318 410 0000 S, 608 310 2400 W) was
floodplain is humid subtropical, with a mean annual characterized by an area of 49.8 ha, a stocking rate of
temperature of 19 °C. Rainfall ranges from 900 to 361.4 AUD ha-1, long grazing period (90 days) and
1000 mm per year, with 73% occurring mainly from short resting periods (37 days); the paddock of W2
October to April (Rojas and Saluso 1987). Soils (318 400 4500 S, 608 300 3300 W) had an area of 15 ha, a
consist of successive layers of sediment carried and stocking rate of 360 AUD ha-1, short grazing period
deposited by the river with each flood phase. Fluvial (27 days) and long resting period (97 days); whereas
soils correspond to Entisol group and Fluvent and the paddock W3 (318 400 9400 S, 608 300 6200 W)
Aquent subgroups (Orellana and Bertoldi De Pomar comprised an area of 19.8 ha and remained without
1969). 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. The minimal area of each sample unit was deter-
During this period, cattle were removed from these mined by constructing a species-area curve for each
floodplain systems. During the study, flood peak of vegetation sample (Chytrý and Otýpková 2003). The
Parana river reached 4.5 m in the reference gauge curve was calculated in the field within the surface
level Parana hydrological station, remaining at high occupied by each homogeneous vegetation unit. The
water levels on the floodplain (higher than 4 m) for a species-area curve was determined to ignore counts of
period of 7 months (December 2015 to August 2016). species from other vegetation units (Perelman et al.
The present study does not have replication of the 2005). For assessing vegetation cover, four quadrats
grazing treatments, since replication is difficult to (4 m2) were chosen randomly in upland and lowland
achieve in large scale studies in floodplain systems. vegetation areas of the exclusion sites and grazed sites
Our results should be interpreted with caution when of each wetland (n = 16). Quadrats of grazed sites
generalizing about floodplain studies. were randomly arranged next to an exclosure. Plant
identification was done in the field; species that could
Sampling not be identified in the field were collected for
taxonomic determination. Identifications were made
Vegetation assemblages were classified according to at the lowest taxonomic level possible (mostly
their topographical position: upland communities (less species) using the available keys (Burkart
floodable) and lowland communities (more floodable), 1969, 1974, 1979,1987; Burkart and Bacigalupo
respectively (Marchetti and Aceñolaza 2012). Vege- 2005; Pensiero et al. 2005). Plants were classified as
tation sampling was conducted monthly during palatable and unpalatable, following previous studies
spring–summer seasons after a high water period. conducted in the wetlands of the Parana River (Rossi
High water period extended from January 2016 to July 2013; Rossi et al. 2014). Direct observation of signs of
2016 and reached a maximum value of 6.5 m of the browsing in leaves and shoots were also taken into
Parana River (Parana Port gauge scale). consideration for classification of palatable species.
Sampling was conducted from October 2016 to
March 2017 according to the movement of cattle Data analyses
between paddocks (Fig. 1c): before the introduction of
cattle to a paddock and after their movement to Plants were classified into functional groups using
another paddock. Grazing regime and rotation of cattle growth habit and life form: palatable (Pal), unpalat-
between wetlands did not follow a fixed schedule, it able (UPal) shrub/subshrub (Shr), forb (Forb), pros-
was determined according to forage supply and plant trate (Pros), erect (Ere) and graminoids (Gr) (Table 3).
growth. All wetlands were sampled during the same Species diversity metrics were calculated from cover
week when cattle movement occurred in any one data including exclusion and grazed sites of upland
wetland. Exclosures were installed on September 2015 and lowland vegetation to compare wetlands with
in the three paddocks including both upland and different resting and grazing periods. Differences in
lowland vegetation, in order to function as exclusion total richness (R), total diversity (Shannon–Wiener
sites (inside the exclosure) and grazed sites (outside index, H’) total dominance (Simpson index, D)
the exclosure) (Fig. 1c). After the installation of the (‘‘total’’ including all plots, and richness, diversity
enclosures, from September 2015 to December 2015 and dominance of functional groups were assessed
and after the high water period, the three studied using two factors ANOVA (R core team, 2017). The
wetlands were subjected to cattle grazing. Continuous factors were the exclosure (exclusion and grazed sites)
exclosures (length = 40 m, width = 20 m, area = 800 and the wetlands (W1, W2 and W3). Sampled sizes
m2) including both vegetation types were installed in ranged from 67 to 71 for exclosure and from 18 to 31
W2 and W3 (Fig. 1b). Due to the great extension of the for wetlands (Table 1). In order to assess differences
lowland zone in W1, this exclosure was divided between exclusion and grazed sites, data of the
(length = 20 m, width = 20 m, area = 400 m2, for different sampling dates (total richness, diversity and
each), including one vegetation type within each dominance and functional groups) were pooled. Prin-
division (Fig. 1b). cipal Component Analysis (PCA) (CANOCO v.4, ter
Braak and Smilauer 1998) with cover data of upland

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Table 1 Sampled sizes (n) of the two factors ANOVA relative dominance was higher in W1 (short resting/long
to upland and lowland vegetation of W1 (long grazing/short grazing period) and W2 (long resting/short grazing
resting period), W2 (short grazing/long rest period) and W3
period) in comparison with W3, and in grazed sites
(without cattle during the studied period), and considering
exclusion sites (inside the exclosure) and grazed sites (outside (ANOVA, P \ 0.01) (Fig. 2). Regarding lowland
de exclosure) vegetation, total richness and total diversity were
Upland Lowland
higher in W1 than in W2 and W3, whereas dominance
was lower in W3 (ANOVA, P \ 0.05) (Fig. 2). We
W1 24 24 found no significant differences between exclusion
W2 31 29 and grazed sites in terms of richness, diversity and
W3 20 18 dominance in lowland communities (ANOVA,
Exclusion sites 68 68 P [ 0.05) (Fig. 2).
Grazed sites 71 67 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)
and lowland vegetation was performed in order to (ANOVA, P \ 0.01), whereas dominance of palat-
identify major sources of variation in vegetation cover able, forb and prostrate vegetation were higher in W1
among wetlands. Prior to analysis, data were arcsin and W2 (ANOVA, P \ 0.01) (Fig. 3a). Richness and
transformed. The Indicator Species Analysis (R pro- diversity of unpalatable, shrub, erect and graminoid
gram, Dufrêne and Legendre 1997) was used to vegetation were significantly higher in exclusion sites
identify the taxa most strongly related to exclusion and (Fig. 4a), whereas dominance of erect and graminoid
grazing sites. plants were higher in grazed sites (Fig. 4a). In relation
to lowland vegetation, richness and diversity of
palatable, unpalatable, shrub/subshrub, forb, prostrate
Results and erect species were significantly higher in W1
(ANOVA, P \ 0.001) (Fig. 3b). Dominance of palat-
Sixty-eight native species were identified in the three able, forb and erect vegetation were significantly
studied wetlands (Table 3). These species belonged to higher in W3, whereas dominance of unpalatable and
31 botanical families, being the most representative shrub/subshrub vegetation were higher in W1 and W2
Poaceae (12 species), Asteraceae (10 species), Faba- (ANOVA, P \ 0.001) (Fig. 3b). Richness and diver-
ceae (5 species) and Solanaceae (5 species). Palatable, sity of forb and prostrate lowland vegetation were
forb and prostrate species were dominant among both significantly higher in W1, whereas dominance of forb
upland and lowland vegetation (relative cover [ vegetation was higher in W3 (ANOVA, P \ 0.05)
50%), while unpalatable, shrub and erect species (Fig. 4b).
showed lower values of vegetation cover (relative
cover \ 40%). Grazed leaves and shoots of Ludwigia Ordination analysis
peploides were observed, leading to its classification
as palatable species. Dominant species in upland and The PCA ordination of upland vegetation sorted the
lowland sites of the three wetlands were Cynodon paddocks with different grazing and resting periods
dactylon, L. peploides, Persicaria punctata and along Axis 1. In the positive side of this axis, L.
Echinochloa crus-pavonis (relative cover [ 10%) peploides and Poa lanigera characterized W1 (short
(Table 2). resting/long grazing period), whereas C. dactylon was
associated with W2 (long resting/short grazing period)
Comparison between wetlands on the negative side. Along axis 2, samples of W2 and
and exclusion/grazed sites Setaria parviflora, P. punctata and Leersia hexandra
species characterized the positive side, whereas L.
Total richness and total diversity of upland vegetation peploides and P. lanigera were associated to the
were significantly higher in W3 (without cattle) and in negative side of this axis (Fig. 5a). A group of 24
exclusion sites (ANOVA, P \ 0.01), whereas 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 Lowland

W1
Exclusion Poa lanı´gera (12.7%); Cynodon dactylon (10.9%); Sesbania virgata (14.7%); Persicaria punctata (12.7%);
Ludwigia peploides (10.7%) Echinochloa crus-pavonis (10%)
Grazing L. peploides (20.2%); P. lanı´gera (18.1%); C. dactylon P. punctata (24.6%); L. peploides (15.6%); S. virgata
(10.2%) (10.8%)
W2
Exclusion P. punctata (19.2%) P. punctata (39.7%); L. peploides (18.7%)
Grazing C. dactylon (28.2%); P. punctata (13.1%) P. punctata (48.8%); E. crus-pavonis (16%)
W3
Exclusion L. peploides (19.4%); C. dactylon (17.5%) P. punctata (36.4%); E. crus-pavonis (22.1%); L. peploides
(20.5%)
Grazing C. dactylon (14.5%); L. peploides (12.3%) L. peploides (25.7%); P. punctata (22%); E. crus-pavonis
(13%)

common for the three wetlands (Fig. 5a). Concerning decreasing total richness and diversity, and increasing
lowland vegetation (Fig. 5b), L. peploides and Myrio- the dominance of palatable, forb and prostrate species.
phyllum aquatiqum related with W1 characterized the Likewise, in contrast with our first hypothesis, our
positive side of axis 1, whereas P. punctata was related results demonstrated that vegetation parameters did
to the negative side of this axis. In the positive side of not differ as grazing and resting time period differed.
axis 2, E. crus-pavonis characterized samples of W2, Both the wetland with short resting/long grazing
whereas Hymenachne amplexicaulis and Sesbania period (W1) as well as the wetland with long resting/
virgata characterized the negative side of this axis. short grazing period (W2) had lower diversity and
richness in comparison with the wetland without cattle
Indicator species analysis grazing (W3). Indeed, paddocks W1 and W2 subjected
to different resting and grazing periods did not differ in
Regarding upland vegetation, Panicum prionitis, richness and diversity of upland vegetation. W1 was
Cyperus entrerianus, Cissus palmata and subjected to a short rest periods and long grazing
Sapium haematospermum were the most strongly periods, but its greater area, heterogeneity and number
related to grazed sites, while Mimosa tweedieana, E. of patches of vegetation may have buffered the effect
crus-pavonis, Muehlenbeckia sagittifolia, Myriophyl- of cattle. On the other hand, the stocking rate of W2
lum aquaticum and Cayaponia podantha were related could have been compensated by the shorter grazing
to exclusion sites (Table 4). In relation to lowland and rest periods, diminishing cattle effect (Sterk et al.
vegetation, S. parviflora, P. punctata, Ambrosia 2013).
artemisiifolia, C. entrerianus and L. hexandra were In relation to the second hypothesis, grazing
associated to grazed sites and Solanum glaucophyllum produced a simplification of upland species of W1
and L. peploides were related to exclusion sites and W2, with a dominance of palatable and prostrate
(Table 4). species with the ability for regrowth after damage (e.g.
Cynodon dactylon). Negative relationships between
species richness and dominance in grazing systems are
Discussion common in grassland (McNaughton 1983; Koerner
et al. 2014, 2018) and wetland ecosystems (Tanner
Our results demonstrated that rotational cattle grazing 1992; Anderson and Calov 1996; Touzard and
had a significant effect on the composition of upland Klement 2001; Rebergen 2002; Hornung and Rice
vegetation of the studied floodplain wetlands, 2003). This relation was also observed in different

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Fig. 2 Total richness, Shannon diversity and dominance index studied period); and (b) exclosure (white) and grazed (black)
of upland and lowland vegetation between (a) wetlands, W1 sites. Significant differences of ANOVA test (P \ 0.05) are
(black, long grazing/short resting period), W2 (grey, short show with different letters: ‘‘a’’, ‘‘b’’
grazing/long rest period), W3 (white, without cattle during the

wetlands and mesic grasslands of East Africa and Middleton 2002; Miller and Wells 2003) Africa
America, where cattle generate extensive grazing (Dahwa et al. 2014) and Australia (Mavromihalis
lawns in which a few grazing-tolerant grasses with- et al. 2013) showed that grazing reduced the domi-
stand high densities of large herbivores (McNaughton nance of palatable species, with positive effects on
1983; Jacobo et al. 2006; Diaz et al. 2007; Jones et al. diversity and richness of upland vegetation.
2011). In contrast, several studies of grassland Regarding W1 and W2, in both grazed wetlands,
ecosystems (Collins et al. 1998; Hayes and Holl palatable, forbs and prostrate species were detected as
2003; Koerner et al. 2014) and wetlands of North dominant species. These species were r-strategists,
America (e.g., Haines 1995; Jutila 1999; Rebergen and are able to reproduce by vegetative adaptations as
2002; Marty 2015), New Zealand (Buxton et al. 2001; rhizomes and stolons, with positive response, to

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Fig. 3 Mean values (± SD) of richness, Shannon diversity and References: palatable (Pal), unpalatable (UPal), shrub/subshrub
dominance index of each functional groups of upland (a) and (Shr), forb (Forb), prostrate (Pros), erect (Ere) and graminoids
lowland (b) vegetation of W1 (black, long grazing/short rest (Gr). Significant differences of ANOVA test (P \ 0.05) are
period), W2 (grey, short grazing/long rest periods) and W3 show with different letters: ‘‘a’’, ‘‘b’’
(white, without cattle grazing during the studied period).

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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 grazing periods and longer rest periods) and W3 (white, without
(a) and lowland (b) vegetation. Sites ordination: W1 (black, cattle); species ordination. References of species were shown in
long grazing periods and short resting periods), W2 (grey, short Table 3

highly disturbed and overgrazed environments (Alte- alternative resource when more palatable species are
sor et al. 2005; Gutierrez 2014; Quintana et al. 2014; scarce (Massa et al. 2018).
Magnano 2017). These strategies would avoid defo- According to our third hypothesis, the result of this
liation of the aboveground biomass and allow species work showed that grazing exclusion favoured the
to quickly colonize bare ground (Dong and de Kroon colonization, regeneration and survival of upland
1994). Palatable species registered in this study were species, increasing richness and diversity of unpalat-
the same such as those found in studies performed in able, shrub, erect and graminoid species. Others
the lower Parana River (Rossi 2013; Magnano 2017). studies have shown the positive effects of cattle
An exception is Ludwigia peploides which, although it exclusion on vegetation attributes (Cooper and
may not be preferred by cattle, it could serve as an McCann 2011; Shaltout et al. 1996; Fujita et al. 2009;

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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

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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

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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

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Acknowledgements This research was supported by the ommendation. Sci Conserv 179:1–64
Consejo Nacional de Investigaciones Cientı́ficas y Técnicas Casco SL, Neiff JJ, Poi de Neiff A (2010) Ecological responses
(CONICET), and Agencia de Promoción Cientı́fica y of two pioneer species to a hydrological connectivity gra-
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