Oecologia Australis
23(4):776-798, 2019
https://doi.org/10.4257/oeco.2019.2304.06
FLORA AND VEGETATION STRUCTURE OF VEREDA IN SOUTHWESTERN
CERRADO
Suzana Neves Moreira1*, Vali Joana Pott2, Arnildo Pott2, Rosa Helena da Silva2 & Geraldo
Alves Damasceno Junior2
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,
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y
s
1
Universidade Estadual do Mato Grosso do Sul, Rua General Mendes de Moraes, 428, Centro, CEP 79400-000, Coxim,
MS, Brazil.
2
Universidade Federal de Mato Grosso do Sul, Programa de Pós-Graduação em Biologia Vegetal, Instituto de Biociências,
Cidade Universitária, Avenida Costa e Silva, s.n., Pioneiros, CEP 79070-900, Campo Grande, MS, Brazil.
E-mails: suzannanevesmoreira@gmail.com (*corresponding author); arnildo.pott@gmail.com; vali.pott@gmail.com;
rosahellenna@gmail.com; geraldodamasceno@gmail.com
Abstract: This work was carried out aiming to evaluate phytosociological parameters and the influence of
topographic gradient and water levels on plant distribution in a vereda. That for, we sampled two sites (wet
grassland and floodable transition to pond) in two seasons (rainy and dry). Along permanent transects, every
10 m we placed transversally four quadrats of 1 m2, 2 m apart, a total of 280 plots, to estimate percentage
cover of each species and water depth. To evaluate the influence of relief and flood level on plant distribution
we performed a Principal Coordinate Analysis (PCoA) and Analysis of Permutational Multivariate Variance
(PERMANOVA). We recorded 174 species, the richest families were Poaceae (40), Cyperaceae (25) and
Asteraceae (14) and the richest genera Paspalum, Rhynchospora and Utricularia (8 each), Hyptis and Cyperus
(6), Eleocharis, Ludwigia and Xyris (5). The phytosociological parameters revealed the importance of Poaceae,
Cyperaceae and Asteraceae in these communities, corroborating other reports on veredas. We found discrete
difference in the species richness between sampling periods, in both wet grassland and transition areas. The
slope varied from 0 to 314 cm and was significant for species distribution in both seasons, with a continuum
related to waterlogging and surface water on the distinct topographic quotas. In general, we found discrete
groupments of species, particularly in lower quotas. The main indicator species were the filiform Eriochrysis
holcoides (Poales, Poaceae), Rhynchospora emaciata (Poales, Poaceae), Andropogon virgatus (Poales,
Poaceae), Paspalum erianthoides (Poales, Poaceae) and P. flaccidum (Poales, Poaceae). Some species tend to
be selective whilst others show plasticity regarding microhabitat.
Keywords: marsh grassland; phytosociology; savanna; topographic gradient; wetland.
INTRODUCTION
The Cerrado domain is characterized by a
vegetation mosaic of savanna, forest and grassland
(Felfili et al. 2004). According to Ribeiro & Walter
(1998), the vereda is one of the physiognomies. This
environment is interposed between Cerrado stricto
sensu and gallery forest (Oliveira-Filho & Martins
1986) and plays an important role in linking these
vegetation types (Carvalho 1991). Veredas are
legally defined as “physiognomy of savanna, found
on hydromorphic soils, usually with the arboreal
palm Mauritia flexuosa (Arecales, Arecaceae) –
buriti, emergent, without forming continuous
canopy, intermingled with clumps of shrubbyherbaceous species” (Brasil 2012). That rule also
Moreira et al. | 777
considers Permanent Preservation Area (PPA) a
minimum 50 m wide belt along veredas, after the
limit of the swampy or waterlogged area. Although
there is a law protecting veredas, considering
them as PPAs, they undergo anthropic processes
that may become irreversible (Oliveira et al. 2009)
since they are highly sensitive to disturbance and
little resilient environments (Carvalho 1991).
Human disturbances in wetlands can be shown by
floristic changes, causing loss of biodiversity and
consequent disruption of the ecosystem (Meirelles
et al. 2004).
The distribution of veredas is basically
conditioned by physical factors, such as wet plain
surfaces or flat valleys, permeable surface layer over
an impermeable subsoil, water saturation nearly
year-round and shallow water table (Drummond et
al. 2005), where gentle slopes favor its seasonal rise
to the surface (Oliveira-Filho & Martins 1986). The
hydromorphic soils are ill-drained (Baccaro 1994,
Ribeiro & Walter 1998), such as gleysols, planosols
and organosols (Drummond et al. 2005). Guimarães
et al. (2002) found two soil classes, Haplic Gleysol
and Melanic Gleysol, in veredas in Uberlândia, MG.
The hydrologic regime in wetlands is the
major determinant in plant community patterns
and species zonation (Casanova & Brock 2000).
Gentry (1988) suggested that habitat contributes
significantly for plant species diversity along
topographic and edaphic gradients, as well as
Rezende (2007) who found a strong influence of
soil moisture gradient on species distribution,
assorting plants into groups of dry and wet habitats
and some occurring in both conditions.
In the study area, the drainage from the wet
grassland provides water to an adjacent pond
(Moreira et al. 2015). Consequently, areas near the
pond (transitional areas) remain with more water
throughout the year when compared with wet
grassland. The hypothesis we want to test is that
topographic gradient and consequent different
water levels influence plant species distribution
patterns and abundance in vereda vegetation.
Therefore, the study was divided into secondary
objectives: (1) present the floristic composition
of the whole vereda; (2) evaluate species richness
and abundance in the areas (wet grassland and
transitional areas) and in the seasons (dry and
rainy); (3) analyze, in each season, if the topography
and different water levels influence plant species
distribution; (4) compare the floristic similarity
between areas and seasons; (5) compare species
richness between areas.
MATERIAL AND METHODS
Study area
Our work was done in a vereda (20º33’24.1” S,
54º47’23.6” W, datum SAD69) at the Fazenda
Modelo, municipality of Terenos, Mato Grosso do
Sul state, Brazil. The area is at 550 m altitude and the
seasonal climate is Aw in the classification of Peel et
al. (2007). Monthly rainfall data were provided by a
weather station, 30 km away, at the headquarters of
Embrapa Beef Cattle Research Center. The original
vegetation was Cerrado woodland of which a
fragment remains at the wetland headwater, near
an outcrop of laterite we believed that underlies
this vereda. Close to the headwater is a stand of M.
flexuosa. Surrounded by pasture (mainly Paspalum
notatum (Poales, Poaceae) and Urochloa spp.),
the study area is grazed and trampled by cows and
horses, mainly in the dry season and on the edges,
not sampled.
Fieldwork and data analysis
Samplings were performed in the rainy and in the
dry season. We placed four permanent transects
to sample vegetation, being two on the floodable
transitional zone, with 170 m (68 plots) and 140 m
(56 plots) and two on the wet grassland, one 150
m long (60 plots) and another one 240 m long (96
plots), a total of 280 plots. The transition site, so
called because it is located near a pond (described
by Moreira et al. 2011), is seasonally flooded.
In each transect, at every 10 m, we placed four
plots of 1 m 2 transversally to the transect line
and 2 m apart, a total of 280 plots. To visually
estimate cover, we imagined a cross line dividing
the plot in four quarters, and so on (Figure 1). The
percentage cover (vertical projection) per plant
species within each plot was visually estimated,
always by the same two persons (S. N. M. and V.
J. P.), who recognized vegetative plants. Some
grasses flowered where we trampled them.
Fertile plant specimens were botanized for the
Herbarium CGMS of the Universidade Federal de
Mato Grosso do Sul, while sterile ones were taken
to the greenhouse until flowering. Taxonomic
identification was achieved comparing vouchers
Oecol. Aust. 23(4): 776–798, 2019
778 | Vegetation structure of vereda
Figure 1. Study area and scheme of distribution of the plots in the place at the Fazenda Modelo, municipality
of Terenos, Mato Grosso do Sul state, Brazil.
in the herbarium and consulting bibliography and
specialists.
We calculated the phytosociological parameters
Absolute Frequency (AF), Relative Frequency
(RF), Absolute Cover (AC) and Relative Cover (RC)
(Brower & Zar 1984) and the Sørensen Similarity
Index (Mueller-Dombois & Ellenberg 1974). For that
analysis we used percentage of cover as a measure of
abundance instead of number of individuals, once
defining an individual is very difficult for bunchy
herbaceous species (Kent & Coker 1992), generally
filiform.
We used an optical level to do the topographic
survey on the flood free part and we measured
water depth per plot with a ruler (in cm) on flooded
grassland to determine the altimetric position of
each plot along transects. To evaluate the influence
of the topography on the pattern of species
distribution in the wet grassland and in transition
areas, we utilized the spreadsheets with the
percentages of cover of the species in relation to the
topographic quotas of the terrain. In the analyses,
the species cover percentages were distributed
according to the respective topographic quotas,
on a scale of 15 cm per quota. The plots within the
limits of the topographic quotas were grouped for
the analyses.
Oecol. Aust. 23(4): 776–798, 2019
To compare the data between both seasons, we
utilized the Principal Coordinates Analysis (PCoA),
which allows to explore and visualize similarities
or differences between data sets and reveals a
centralized matrix decomposed in its eigenvalues
and eigenvectors. Next, the groups identified
in the PCoA were submitted to a Permutational
Multivariate Analysis of Variance (PERMANOVA)
to verify significance between the groupments of
topographic quotas. The analyses were performed
utilizing the package Vegan (Oksanen et al. 2014) of
the program R (R Core team 2017).
Regarding indicator species (Dufrêne & Legendre
1997), we utilized the function indval() of the
package labdsv (David, 2016), also in R. An indicator
species has specificity to a particular niche; thus, its
presence or abundance was utilized to indicate a
particular environment (Cáceres et al. 2010).
RESULTS
Overall floristic data
The overall data shows a high plant richness in the
studied vereda. In the wet grassland plus transition
areas, we sampled 174 species of 97 genera and 46
families. Poaceae (40 species), Cyperaceae (25),
Asteraceae (14) were the richest families, adding
up 45% of the species, while 21 families showed
Moreira et al. | 779
single species. The richest genera were Paspalum,
Rhynchospora and Utricularia (8 each), Hyptis and
Cyperus (6), Eleocharis, Ludwigia and Xyris (5),
typical of veredas (Appendix 1).
Comparing the general richness found in the wet
grassland with the transitional areas, we observed
that the first had a slightly lower number of species
(113) than the latter (122). In the transition zone,
we sampled 43 exclusive species, while the wet
grassland showed 33, and 79 were common to both
formations (Appendix 1). The Sørensen Similarity
indexes for habitats and seasons showed a higher
floristic similarity between dry and rainy seasons
in similar environments than in relation to different
environments (Table 1).
In the analysis of the topographic gradient, we
compared wet grassland and transitional area. The
topographic quotas varied from 314 cm, the top
level, where the water table was just close to the
surface, to zero, the lowest ground, where the soil
was waterlogged or flooded since toward the pond
the water rose to the surface (Figures 2 and 3). The
topography was important to determine the pattern
of species distribution in both rainy (p = 0.001) and
dry period (p = 0.001).
In the rainy period, the Principal Coordinates
Analysis (PCoA) indicates two main centroids related
to the topographic quotas of 0-15 cm and 15-30 cm.
The other quotas do not represent a clear pattern of
separation of species, indicating that they are more
Table 1. Sørensen Similarity Index between areas and sampled periods in a vereda, municipality of Terenos,
State of Mato Grosso do Sul, Brazil.
Grassland Rainy
Grassland Dry
Transitions Rainy
Transitions Dry
Grassland/Rainy
Grassland/Dry
.
.
0.81
.
0.65
0.51
0.62
0.55
Transitions/Rainy
Transitions/Dry
.
.
.
.
.
.
0.87
.
Figure 2. Ordination of the herbaceous species by Principal Coordinates Analysis (PCoA) through declivity
of the terrain sampled in a vereda in municipality of Terenos, state of Mato Grosso do Sul, Brazil, in the rainy
period. The numbers mean the topographical quotas in centimeters.
Oecol. Aust. 23(4): 776–798, 2019
780 | Vegetation structure of vereda
Figure 3. Ordination of the herbaceous species by Principal Coordinates Analysis (PCoA) through declivity
of the terrain sampled in a vereda in municipality of Terenos, state of Mato Grosso do Sul, Brazil, in the dry
period. The numbers mean the topographic quotas in centimeters.
similar between them (Figure 2).
The Indicator Species Analysis for the rainy period
revealed 62 species related with distinct topographic
quotas forming two large groups, the first with
22 species and the second with 40 species. The
indicator species of the first group are distributed in
nine families, the richest being Poaceae (7 species),
Cyperaceae (3) and Malvaceae, Lentibulariaceae and
Asteraceae (2). The characteristic species of group I
are: Utricularia gibba (Lamiales, Lentibulariaceae),
Rhytachne rottboellioides (Poales, Poaceae), Byttneria
palustris (Malvales, Malvaceae), Saccharum asperum
(Poales, Poaceae) and Andropogon hypogynus
(Poales, Poaceae). The species of the second group
are represented by 10 families, Poaceae (12 species),
Cyperaceae (11) and Asteraceae (5) having the largest
number. The main indicator species are Eriochrysis
holcoides (Poales, Poaceae), Rhynchospora emaciata
(Poales, Cyperaceae), Andropogon virgatus (Poales,
Poaceae), Paspalum erianthoides (Poales, Poaceae)
and Paspalum flaccidum (Poales, Poaceae).
In the dry period, the Principal Coordinates
Analysis (PCoA) revealed a spaced pattern of the
centroids, indicating that more topographic quotas
Oecol. Aust. 23(4): 776–798, 2019
influenced the pattern of distribution. The quota of
0-15 cm, as well as in the rainy season, indicates the
preference of some species, whilst the quotas 15-30
cm and 30-45 cm are more related between them
than with the others. The highest topographic quotas
demonstrated that the species occur similarly along
the gradient, without forming characteristic groups
(Figure 3).
The Indicator Species Analysis for the dry
period revealed 57 species related to the distinct
topographic quotas. Differently from the rainy
period, when we related the indicator species
with the graph of species distribution, there
is not a clear spatial separation or preference.
However, the analysis indicated some key-species
for the quota of 0-15 cm Schizachyrium gracilipes
(Poales, Poaceae) and Eryngium ebracteatum
(Apiales, Apiaceae) and 15-30 cm, Leersia hexandra
(Poales,
Poaceae),
Caperonia
castaneifolia
(Malpighiales,
Euphorbiaceae),
Eleocharis
plicarhachis (Poales, Cyperaceae), E. acutangula
(Poales,
Cyperaceae),
Mikania
stenophylla
(Asterales, Asteraceae), Phyllanthus stipulatus
(Malpighiales, Phyllanthaceae) and Aeschynomene
Moreira et al. | 781
fluminensis (Fabales, Fabaceae). The quota of 3045 cm presented just a single indicator species,
Lessingianthus rubricaulis (Asterales, Asteraceae).
Wet grassland
In wet grassland, we sampled 113 species, of 71 genera
and 34 families. Poaceae (27 species), Cyperaceae
(20), Asteraceae (11), Lamiaceae (6) and Xyridaceae
and Lentibulariaceae (5) were the richest families,
adding to 65.5% of total richness. The richest genera
were Rhynchospora (8 species), Hyptis, Paspalum,
Utricularia, and Xyris (5 each). Twenty families were
represented by a single species and accumulated
17.8% of the floristic richness (Appendix 1).
Eriochrysis holcoides (Poales, Poaceae) showed
the highest cover in the rainy season, followed
by Anthaenantia lanata (Poales, Poaceae), P.
flaccidum, P. erianthoides, Rhynchospora globosa
(Poales, Cyperaceae), A. virgatus, A. hypogynus and
Axonopus uninodis (Poales, Poaceae). In the dry
season, A. uninodis was the species with highest CV,
followed by P. erianthoides, A. lanata, A. virgatus, P.
dedeccae (Poales, Poaceae), P. flaccidum, R. globosa,
A. hypogynus and Rhynchospora marisculus (Poales,
Cyperaceae) (Table 2). The cover of Eryngium
pandanifolium (Apiales, Apiaceae) did not differ
between seasons.
Transition areas
In the transition areas, we sampled 122 species
and 72 genera distributed into 34 families. Poaceae
(28 species), Cyperaceae (20), Asteraceae (11) and
Lentibulariaceae (8) were the richest families,
adding to 55.8% of all recorded species. Another 15
families were represented by a single species or just
17.8% of the floristic richness. The richest genera
were Utricularia (8), Rhynchospora (7) and Xyris,
Eleocharis, Scleria, Hyptis and Ludwigia (4 each)
(Appendix 1).
Saccharum asperum, A. hypogynus and
A. uninodis were the species with the highest
relative cover the rainy season, compared with R.
rottboellioides, A. hypogynus and Imperata tenuis
(Poales, Poaceae) in the dry season (Table 2).
DISCUSSION
Overall floristic data
Poaceae, Cyperaceae and Asteraceae were the
richest families in other studies on wetlands in
Cerrado (Guimarães 2001, Guimarães et al. 2002,
Araújo et al. 2002, Meirelles et al. 2004, Munhoz &
Felfili 2008, Oliveira et al. 2009, Moreira et al. 2011,
Moreira et al. 2015, Bijos et al. 2017), as we also
observed. These families contain species which
Table 2. Species with highest Cover Values in Wet grassland and Transition Area at Vereda, municipality of
Terenos, State of Mato Grosso do Sul, Brazil, in two differents environments and seasons with their respective
Collector Number (COL.). M= Moreira,S.N. Numbers in bold represent the highest cover values.
Family/Species
Col.
Wet grassland
Transition Area
Rainy
Dry
Rainy
Dry
M 143
5.8
6.33
1.04
0.35
M 37
1.11
1.93
6.45
8.58
Andropogon hypogynus Hack.
M 89
7.13
7.16
17.38
16.86
Axonopus uninodis (Hack.) G.A. Black
M 19
6.49
18.6
13.14
11.09
Eriochrysis holcoides (Nees) Kuhlm.
M 101
23.04
3.78
3.65
5.69
Imperata tenuis Hack.
M 208
3.15
2.55
11.7
15.73
Paspalum erianthoides Lindm.
M 303
12.22
14.38
12.55
13.83
Paspalum flaccidum Nees
M 285
12.91
10.15
1.25
1.19
Rhytachne rottboellioides Desv.
M 90
2.07
1.84
5.05
34.32
Saccharum asperum (Nees) Steud.
M 61
1.75
0.25
22.26
3.66
APIACEAE
Eryngium pandanifolium Cham. & Schltdl.
MALVACEAE
Byttneria palustris Cristóbal
POACEAE
Oecol. Aust. 23(4): 776–798, 2019
782 | Vegetation structure of vereda
occur mostly in sunny habitats (Coutinho 1978)
but in wet grassland, the herbaceous dominance
can be attributed to excess of water in the soil, what
hinders the establishment of woody strata (Araújo
et al. 2002). We collected a recently described new
species, Cyperus longiculmis (Poales, Cyperaceae)
(Pereira-Silva et al. 2018).
In spite of wet grassland and transition zone
presenting some exclusive species, the large
number of species (79) in common reflects their
capacity to colonize both habitats with higher
or lower water saturation and a high similarity
between the same habitats, even considering
distinct sampling seasons. Thus, the sampled area
exerts higher influence on the species composition
than does the water regime. A similar result was
already observed by Moreira et al. (2011) in ponds
associated with veredas.
The relationship between topographic quotas
and distribution patterns is attributed to different
moisture conditions along the relief gradient.
Similar results were described in other studies
(see Teixeira & Assis 2005 and Gaya 2014), where
hydromorphic soils have different floristic and
structural peculiarities, besides providing a greater
variety of environments and, consequently, a more
diverse flora. In our study area, the slope is easily
perceived as slow and the drainage is gradual.
The water movement is visible in the veins and
animal tracks, from the highest to the lowest
ground, where the water begins to dam up, as
well as in puddles in small depressions between
grass bunches. The soil was not analyzed, but we
observed that the wet grassland was waterlogged in
spite of the slope because there is a thicker organic
top horizon functioning as a sponge compared
with the transition floodable grassland. Positive
relationships between declivity and species
distribution in forests were described by several
authors (see Gartlan et al. 1986, Oliveira-Filho et al.
1994a, 1994b, Van Den Berg & Oliveira-Filho 1999).
The Principal Coordinates Analysis and the
Indicator Species Analysis in both sampled periods
suggest the formation of groups mainly on the
lowest quotas and that these lack exclusive species
since they occur in the transition zone between
the wet grassland and the associated pond. This
could be explained by transition areas containing
species from either. The transition zones can be
characterized as ecotones. A pioneer study on vereda
Oecol. Aust. 23(4): 776–798, 2019
by Goldsmith (1974) found a type of vegetation
in the transition area and that this change occurs
in a subtle way, as a continuum. In the moist and
flooded gradient of veredas occurs a continuum
of species replacement gradient where the plant
communities have components from neighbouring
zones and also exclusive species (Costa 2007). The
moisture gradient can select different species (Kurtz
et al. 2013). Some occur preferentially in flooded
soils, others in dry soils, plus those occurring in
both environments (Guimarães et al. 2002, Resende
et al. 2013).
The indicator species of the rainy period reflect
basically the growth habit of the individuals.
Many cespitous tall grasses have vigorous growth
in wetlands, such as A. hypogynus, A. virgatus,
E. holcoides, P. erianthoides, P. flaccidum, R.
rottboellioides and S. asperum, as well as Cyperaceae,
e.g. R. emaciata. The submerged or palustrine herb
Utricularia gibba (Lamiales, Lentibulariaceae) has
abundant ramified stolons (Baleeiro et al. 2017),
what contributes to its higher frequency and cover,
preferentially in wetter habitats, along animal
tracks throughout the area.
The indicator species in the dry period
correspond majorly to other botanical families. P.
stipulatus, A. fluminensis and L. rubricaulis do not
present stoloniferous propagation, such as Poaceae
and Cyperaceae, however, occur in specific zones,
i.e. areas with less flooding.
Wet grassland
Similar to several reports the richest families in
wet grassland were Poaceae, Cyperaceae and
Asteraceae (Guimarães et al. 2002, Araújo et al.
2002, Meirelles et al. 2004, Munhoz & Felfili 2007,
Munhoz & Felfili 2008, Oliveira et al. 2009, Moreira
et al. 2011, Moreira et al. 2015, Bijos et al. 2017), plus
the main genera Rhynchospora and Xyris (Araújo
et al. 2002, Munhoz & Felfili 2007, Resende et al.
2013). Araújo et al. (2002) suggest that these richest
families have species well adapted to the conditions
of soil and waterlogging, commonly found in
veredas, reflected by the high representativity of the
herbaceous stratum.
The highest cover values (CV) represent species
with dense tussocks as much in the dry as in the
rainy season, such as A.lanata and P. erianthoides
(Poaceae) and R. globosa (Cyperaceae). The
dense tussocks of filiform species can hinder the
Moreira et al. | 783
establishment of other species in wetter zones of
the vereda (Araújo et al. 2002), such as A. lanata,
A. hypogynus, A. uninodis and R. rottboellioides.
E. pandanifolium (Apiaceae) showed high CV in
both seasons. Differently from grasses, this species
grows in groups and shows high competitiveness
for space with graminoids, mainly for having
rosette-like large strong leaves (Cardozo 2017),
pushing others aside. The species which form
dense tussocks, such as grasses, tend to show
higher cover than small and slender herbs in high
abundance (Munhoz & Felfili 2008). We observed
that some species of Lamiaceae (Hyptis spp.),
Ochnaceae (Sauvagesia racemosa), Apocynaceae
(Rhabdadenia
ragonesei),
Orobanchaceae
(Escobedia grandiflora), Gesneriaceae (Sinningia
elatior)
and
Lentibulariaceae
(Utricularia
praelonga) elongate stems over the filiform tussocks
to expose the flowers, as also reported by Araújo et
al. (2002). The dense grass bunch expands to its
periphery and tends to die in the middle, where
the decayed matter becomes colonized by ferns
and shrubs (e. g. Miconia chamissois, Myrtales,
Melastomataceae).
There is a higher probability to find fertile plants
during the rainy season. We found the highest
species richness in the rainy season (Munhoz
& Felfili 2006), what does not correspond to the
establishment of true aquatic plants but to the
appearance of species between seasons or just not
sampled before.
deficit. This tussock grass is only grazed after burn
or during critical dry or flood periods (Pott 1982),
though in the study area it occurs in the transition
temporary pond/wet grassland, where cattle can
have access but prefers softer grasses, what could
explain its high coverage. We observed that I.
tenuis formed dense dominant populations, what
is due to the multiple aggregate culms (Welker &
Longhi-Wagner 2012) from the strong rhizome net.
Frequent in the studied vereda, R. rottboellioides is
not yet cited for the Central-West region in the list
of the Brazilian Flora (Flora do Brasil 2018).
We concluded that the studied vereda presents
a continuum of environmental conditions related
to soil moisture and waterlogging of the distinct
topographic quotas. Thereby, some species tend to
occur preferentially in certain microhabitats whilst
others show a wider plasticity.
Transition areas
Araújo, G. M., Barbosa, A. A. A., Arantes, A. A., &
Amaral, A. F. 2002. Composição florística de
Veredas no município de Uberlândia, MG.
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Baccaro, C. A. D. 1994. As unidades
geomorfológicas e a erosão nos chapadões
do município de Uberlândia. Sociedade &
Natureza, Uberlândia, 6(11/12), 19-33
Baleeiro, P. C., Moreira, A. D. R., Silva, N. G.,
& Bove, C. P. 2017. Flora do Rio de Janeiro:
Lentibulariaceae. Rodriguésia, 68(1), 59-71.
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Bijos, N. R., Eugênio, C. U. O., Mello, T. R. B., Souza,
G. F., & Munhoz, C. B. R. 2017. Plant species
composition, richness, and diversity in the
palm swamps (veredas) of Central Brazil. Flora,
236-237, 94-99. DOI: 10.1016/j.flora.2017.10.002
Brasil. 2012. Lei No. 12.651, de 25 de maio de 2012.
Poaceae, Cyperaceae and Asteraceae correspond
to the richest families in most reports on wetlands,
such as veredas (Araújo et al. 2002, Guimarães et
al. 2002, Meirelles et al. 2002, Tannus & Assis 2004,
Munhoz & Felfili 2006, Munhoz & Felfili, 2007,
Oliveira et al. 2009, Moreira et al. 2011, Resende et
al. 2013, Moreira et al. 2015, Silva et al. 2017, Bijos
et al. 2017, Moreira et al. 2017). Coutinho (1998)
and Araújo et al. (2002) mention that these three
families present a great number of genera and
heliophilous species.
Saccharum asperum can occur either in drier
areas of the vereda edge (Oliveira et al. 2009)
or water courses and marshes, with high cover
(Meirelles et al. 2002). Andropogon hypogynus
was representative in both seasons, showing to
be adapted to either flooding or relative water
ACKNOWLEDGEMENTS
We thank the Universidade Federal de Mato
Grosso do Sul (UFMS) for sponsoring the study. To
Conselho Nacional de Desenvolvimento Científico
e Tecnológico (CNPq) for scholarship to Suzana
Neves Moreira and grants to Arnildo Pott and
Geraldo Alves Damasceno Junior. To Embrapa
Gado de Corte that gave us access to the area and
kept firebreaks.
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Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. Species sampled in Wet grassland and Transition Areas at Vereda, municipality of Terenos, State of Mato Grosso do Sul, Brazil, with their respective
Collector Number (Col.), Relative Frequency (RF), Relative Cover (RC) and Cover Values (CV). The value was described for two different seasons (Rainy and
Dry) and two different environments (Wet Grassland and Transition Area). M= Suzana Neves Moreira and P= Vali Joana Pott. The numbers in bold represent
the highest coverage values.
Wet grassland
Family/Species
Col.
Rainy
Transition Area
Dry
Rainy
RC
RF
CV
RC
RF
CV
M 084
0.06
0.12
0.18
0.06
0.13
0.19
P 10.918
0.08
0.37
0.45
0.4
0.27
0.67
Dry
RC
RF
CV
RC
RF
CV
0.53
1.17
1.7
1.7
3.85
5.55
0
0.16
0.16
3.96
2.02
5.98
0.31
0.09
0.4
ALISMATACEAE
Echinodorus longipetalus Micheli
Helanthium tenellum (Mart.) Britton
Sagittaria rhombifolia Cham.
M 87
APIACEAE
Eryngium ebracteatum Lam.
M 178
0.08
0.12
0.2
Eryngium floribundum Cham. & Schltdl.
M 31
0.74
0.61
1.35
0.94
0.66
1.6
0.25
0.16
0.4
0.83
0.75
1.58
M 143
3.72
2.08
5.8
3.94
2.39
6.33
0.57
0.47
1.04
0.07
0.28
0.35
Mandevilla widgrenii C. Ezcurra
M 284
0.06
0.37
0.43
0.3
0.62
0.92
2.99
5.91
8.91
Rhabdadenia madida (Vell.) Miers
M 26
0.06
0.37
0.43
0.03
0.23
0.27
0.03
0.19
0.22
Secondatia densiflora A. DC.
M 58
0.1
0.19
0.29
Widgrenia corymbosa Malme
M 54
Eryngium
Schltdl.
pandanifolium
Cham.
&
APOCYNACEAE
0.13
0.14
0.25
0.4
0.65
0.32
1.63
1.95
0.61
2.07
2.68
0.03
0.16
0.18
0.08
0.09
0.17
ARECACEAE
Mauritia flexuosa L.f.
M 366
ASTERACEAE
Acilepidopsis echitifolia (Mart. ex DC.)
H.Rob.
M 192
Achyrocline alata (Kunth) DC.
M 156
0.59
0.86
1.45
0.06
0.13
0.19
Appendix 1. Continued on next page...
Moreira et al. | 787
Oecol. Aust. 23(4): 776–798, 2019
0.01
Wet grassland
Family/Species
Campuloclinium
(Less.) DC.
Col.
macrocephalum
Rainy
Transition Area
Dry
Rainy
Dry
RC
RF
CV
RC
RF
CV
RC
RF
CV
RC
RF
CV
M 358
Chromolaena laevigata (Lam.) R.M.
King & H. Rob.
M 199
0.52
0.49
1.01
0.56
0.27
0.83
0.04
0.16
0.2
Chromolaena palmaris (Sch. Bip. ex
Baker) R.M. King. & H. Rob.
M 359
1.29
1.84
3.13
0.06
0.13
0.19
0.06
0.23
0.29
Conyza bonariensis (L.) Cronquist
M 135
0.02
0.16
0.17
0.05
0.19
0.24
Leptostelma tweediei (Hook. & Arn.)
D.J.N.Hind & G.L.Nesom
Lessingianthus bardanoides (Less.) H.
Rob.
Lessingianthus rubricaulis (Humb. &
Bonpl.) H.Rob.
M 107
0.06
0.25
0.31
0.14
0.27
0.41
M 100
0.36
0.25
0.61
0.07
0.27
0.34
0.1
0.39
0.49
0.04
0.19
0.23
M 176
0.7
1.1
1.8
0.02
0.13
0.15
0.32
1.17
1.49
0.45
1.88
2.33
Mikania cordifolia (L. f.) Willd.
M 201
0
0.12
0.12
0.14
0.23
0.37
Mikania stenophylla Holmes
M 95
0.12
0.39
0.51
0.03
0.23
0.26
Pichrosia longifolia D. Don
Trichogonia crenulata (Gardner) D.J.N.
Hind
Vernonanthura chamaedrys (Less.)
H.Rob.
BEGONIACEAE
Begonia cucullata Willd.
P 8914
0.19
0.86
1.05
M 125
0.67
1.1
1.77
1.63
1.33
2.96
0.22
0.47
0.69
M 40
0.11
0.37
0.48
0.31
0.27
0.58
0.49
0.7
1.19
0.23
0.66
0.89
M 108
0.01
0.12
0.13
0.01
0.13
0.14
0.02
0.16
0.17
0.01
0.09
0.1
0.02
0.16
0.17
0.65
0.23
0.89
0.15
0.09
0.25
CAMPANULACEAE
Lobelia aquatica Cham.
M 152
Lobelia camporum Pohl
M 113
Pratia hederacea Hook. & Arn.
M 45
0.06
0.12
0.18
0.05
0.13
0.18
Appendix 1. Continued on next page...
788 | Vegetation structure of vereda
Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued
Appendix 1. ...Continued
Wet grassland
Family/Species
Col.
Rainy
RC
CHARACEAE
Nitella furcata (Roxb. ex Bruzelius)
Agardh
CHLORANTHACEAE
M 307
Hedyosmum brasiliense Miq.
M 43
RF
Transition Area
Dry
CV
RC
RF
Rainy
CV
Dry
RC
RF
CV
0.09
0.39
0.48
0.11
0.54
0.65
0.01
0.16
0.16
RC
RF
CV
0.17
0.56
0.73
0.03
0.19
0.22
CUCURBITACEAE
Melothria fluminensis Gardner
M 28
0.05
0.25
0.3
Ascolepis brasiliensis (Kunth) Benth. ex
C.B. Clarke
M 11
0.3
1.47
1.77
Cyperus brevifolius (Rottb.) Endl. ex
Hassk.
23
0.04
0.12
0.16
Cyperus haspan L.
M 67
0.04
0.12
0.16
Cyperus longiculmis Pereira-Silva, Hefler
& R. Trevis.
M 71
Cyperus rigens var. impolitus (Kunth)
Hefler & Longhi-Wagner
P 10.841
0.37
1.23
1.6
0.71
1.86
2.57
0.03
0.23
0.27
M 77
0.2
0.86
1.06
0.25
0.27
0.52
0.03
0.16
0.19
0.09
0.93
1.02
0.2
0.94
1.14
0.03
0.39
0.42
0.34
0.66
1
0.24
0.62
0.86
0.75
2.72
3.47
CYPERACEAE
Cyperus valie Pereira-Silva. Hefler & R.
Trevis.
Eleocharis acutangula (Roxb.) Schult.
0.53
0.65
P 10.274
M 310
Eleocharis capillacea Kunth
P 10.848
0.13
0.86
0.99
0.12
0.53
0.65
Eleocharis minima Kunth
P 10.985
0.01
0.49
0.5
0.01
0.13
0.14
M7
0
0.12
0.12
Eleocharis nudipes Palla
Appendix 1. Continued on next page...
Moreira et al. | 789
Oecol. Aust. 23(4): 776–798, 2019
Cyperus unioloides R. Br.
0.12
Wet grassland
Family/Species
Col.
Rainy
RC
Eleocharis
Svenson
plicarhachis
(Griseb.)
RF
Transition Area
Dry
CV
RC
RF
Rainy
CV
M 309
RC
RF
CV
RC
RF
CV
0.58
2.56
3.14
0.48
1.6
2.07
Lipocarpha humboldtiana Nees
M 12
0.17
0.74
0.91
Rhynchospora conferta (Nees) Boeckeler
M 10
0.04
0.25
0.29
M
0.2
0.98
1.18
M 116
0.26
0.49
0.75
Rhynchospora loefgrenii Boeckeler
M 14
3.33
4.9
8.23
4.41
5.05
9.46
Rhynchospora marisculus Lindl. ex Nees
M 80
2.7
2.45
5.15
3.97
3.06
7.03
Rhynchospora rugosa (Vahl) Gale
M 325
0.62
0.86
1.48
1.06
1.73
2.79
Rhynchospora trispicata (Nees) Schrad.
ex Steud.
M 204
0.41
0.49
0.9
0.95
1.33
2.28
0.68
1.09
M 294
1.69
1.84
3.53
0.5
1.33
1.83
0.3
Rhynchospora corymbosa (L.) Britton
Rhynchospora
Boeckeler
Rhynchospora
Boeckeler
(Nees)
emaciata
velutina
(Kunth)
Scleria hirtella Sw.
Dry
1.31
3.32
4.63
0.2
0.93
1.13
0.02
0.09
0.11
0.05
0.4
0.45
0.18
0.23
0.41
0.35
0.19
0.53
1.8
3.03
4.83
0.99
2.82
3.81
0.12
0.19
0.3
0.08
0.09
0.17
0.03
0.19
0.22
1.76
0.19
0.38
0.57
0.93
1.24
0.35
0.94
1.29
0.06
0.62
0.68
M8
0.08
0.16
0.24
Scleria leptostachya Kunth
M 148
0.46
1.1
1.56
0.39
1.73
2.12
0.14
0.62
0.76
0.03
0.19
0.22
Scleria lithosperma (L.) Sw.
M 162
0.29
1.35
1.64
0.64
2.66
3.3
0.01
0.47
0.48
0.31
0.38
0.68
Scleria microcarpa Nees ex Kunth
M 75
0
0.16
0.16
0.12
0.09
0.21
ERIOCAULACEAE
Eriocaulon sellowianum Kunth
M 69
0.2
0.74
0.94
0.27
0.53
0.8
0
0.16
0.16
0.35
0.38
0.73
Syngonanthus caulescens (Poir.) Ruhland
M3
0.52
1.1
1.62
0.65
1.33
1.98
0.02
0.16
0.17
0.04
0.28
0.32
0.31
1.63
1.95
0.54
1.88
2.41
EUPHORBIACEAE
Caperonia castaneifolia (L.) A. St.-Hil.
M 165
Appendix 1. Continued on next page...
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Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued
Appendix 1. ...Continued
Wet grassland
Family/Species
Sapium hasslerianum Huber
Col.
M 144
Rainy
Transition Area
Dry
RC
RF
CV
0.02
0.12
0.14
RC
RF
Rainy
CV
Dry
RC
RF
CV
RC
RF
CV
FABACEAE
Aeschynomene fluminensis Vell.
M 50
0.11
0.23
0.35
0.26
0.38
0.64
Aeschynomene sensitiva Sw.
M 175
0.1
0.31
0.41
0.08
0.09
0.17
0.02
0.16
0.17
0.01
0.09
0.1
0.11
0.31
0.42
0.04
0.09
0.13
0.02
0.19
0.2
0.02
0.09
0.11
0.01
0.09
0.1
GENTIANACEAE
Chelonanthus alatus (Aubl.) Pulle
M 117
0.07
0.37
0.44
Schultesia aptera Cham.
P 10.870
0.3
1.47
1.77
Schultesia gracilis Mart.
M 114
0.97
2.7
3.67
Schultesia heterophylla Miq.
P 10.849
GESNERIACEAE
Sinningia elatior (Kunth) Chautems
M 59
0.32
0.93
1.25
0.02
0.13
0.15
0.81
1.06
1.87
0.35
1.46
1.81
HYPNACEAE
Isopterygium tenerifolium Mitt.
P 10.969
IRIDACEAE
M 98
0.17
0.86
1.03
Cantinoa althaeifolia (Pohl ex Benth.)
Harley & J.F.B.Pastore
M 301
0.82
0.98
1.8
Hyptis crenata Pohl ex Benth.
M 944
0.01
0.12
0.13
Hyptis lavandulacea Pohl ex Benth.
M 188
0.18
0.37
0.55
Hyptis microphylla Pohl ex Benth.
M 360
Hyptis pachyarthra Briq.
M 109
LAMIACEAE
0.26
0.61
0.87
0.09
0.53
0.62
0.08
0.02
0.31
0.16
0.39
0.17
Appendix 1. Continued on next page...
Moreira et al. | 791
Oecol. Aust. 23(4): 776–798, 2019
Cypella laxa Ravenna
Wet grassland
Family/Species
Col.
Rainy
RC
Dry
CV
RC
RF
CV
M 336
0.12
0.16
0.28
0.07
0.09
0.16
Utricularia foliosa L.
M 184
0.51
1.17
1.67
Utricularia gibba L.
M 129
0
0.12
0.12
2.11
4.66
6.77
0.01
0.09
0.1
Utricularia hydrocarpa Vahl
M 180
0.01
0.12
0.13
0.61
2.72
3.33
Utricularia myriocista A. St.-Hil. &
Girard
M 134
0.21
0.62
0.84
Utricularia nervosa Weber ex Benj.
M 30
0.25
0.37
0.62
0.05
0.27
0.32
0.01
0.09
0.1
Utricularia praelonga A. St.-Hil. & Girard
M1
0
0.12
0.12
0.05
0.4
0.45
0.01
0.09
0.1
Utricularia trichophylla Spruce ex Oliv.
M 76
0.03
0.37
0.4
Utricularia tricolor A. St.-Hil.
M 130
0.02
0.09
0.11
Hyptis recurvata Poit.
M 82
0.48
1.96
2.44
RC
RF
CV
0.05
0.4
0.45
0.24
0.66
0.9
Dry
RF
M 170
CV
Rainy
RC
Hyptis pulchella Briq.
RF
Transition Area
LAURACEAE
Nectandra gardneri Meisn.
LENTIBULARIACEAE
0.25
0.23
0.48
MALPIGHIACEAE
Heteropterys procoriacea Nied.
M 38
Heteropterys eglandulosa A. Juss.
M 345
Heteropterys orinocensis (Kunth) A.Juss.
M 371
0.28
0.37
0.65
0.31
0.4
0.71
0.37
0.74
1.11
0.6
1.33
1.93
0.14
0.31
0.45
0.14
0.19
0.33
2.18
4.27
6.45
2.19
6.38
8.58
MALVACEAE
Byttneria palustris Cristóbal
M 37
Melochia simplex A. St.-Hil.
M 362
0.57
1.55
2.13
0.04
0.09
0.13
Melochia villosa (Mill.) Fawc. & Rendle
M 282
0.13
0.31
0.44
0.25
0.75
1
Appendix 1. Continued on next page...
792 | Vegetation structure of vereda
Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued
Appendix 1. ...Continued
Wet grassland
Family/Species
Col.
Rainy
RC
Sida rhombifolia L.
RF
Transition Area
Dry
CV
RC
RF
Rainy
CV
-
Dry
RC
RF
CV
RC
RF
CV
0.01
0.16
0.16
0.87
2.41
3.28
0.02
0.19
0.2
0.36
1.17
1.53
0.94
2.25
3.19
MAYACACEAE
Mayaca sellowiana Kunth
M 83
0.06
0.25
0.31
0.1
0.4
0.5
MELASTOMATACEAE
Acisanthera alsinaefolia Triana
-
Acisanthera divaricata Cogn.
M 314
Acisanthera variabilis (DC.) Triana
M 339
Miconia chamissois Naudin
M 34
Tibouchina gracilis (Bonpl.) Cogn.
M 85
0.21
0.66
0.87
0.34
0.7
1.04
0.32
0.66
0.98
0.47
1.71
2.18
0.34
1.06
1.4
0.17
0.62
0.79
0.02
0.19
0.21
0.12
0.12
0.24
0.34
1.59
1.93
0.04
0.39
0.43
0.15
0.47
0.62
M 174
0.2
0.54
0.74
0.22
0.66
0.88
Ludwigia irwinii Ramamoorthy
M 86
0.37
1.09
1.46
0.41
1.41
1.82
Ludwigia nervosa (Poir.) H. Hara
M 74
0.52
1.59
2.11
0.99
1.99
2.98
0.36
1.09
1.45
0.48
1.5
1.98
Ludwigia sericea (Cambess.) H. Hara
M 39
0.07
0.12
0.19
0.06
0.13
0.19
0.08
0.16
0.24
0.2
0.47
0.67
M 123
0.06
0.12
0.18
0.06
0.13
0.19
0.06
0.16
0.21
0.04
0.09
0.13
MYRSINACEAE
Rapanea umbellata (Mart.) Mez
M 96
OCHNACEAE
Sauvagesia racemosa A. St.-Hil.
M 32
ONAGRACEAE
Ludwigia bullata (Hassl.) H. Hara
(Micheli)
ORCHIDACEAE
Cyrtopodium paludicola Hoehne
Appendix 1. Continued on next page...
Moreira et al. | 793
Oecol. Aust. 23(4): 776–798, 2019
Ludwigia
filiformis
Ramamoorthy
M 166
Wet grassland
Family/Species
Col.
Rainy
Transition Area
Dry
RC
RF
CV
RC
RF
Rainy
CV
Habenaria nuda Lindl.
M 324
0.02
0.12
0.14
Habenaria pungens Cogn.
M 323
0.02
0.12
0.14
M 94
0.14
0.37
0.51
0.06
0.13
0.19
M 363
0.07
0.12
0.19
0.19
0.13
0.32
0.02
0.13
0.15
Dry
RC
RF
CV
0.01
0.16
0.16
0.11
0.31
0.43
RC
RF
CV
0.14
0.28
0.42
0.15
0.09
0.25
11.79
5.07
16.86
OROBANCHACEAE
Escobedia grandiflora (L. f.) Kuntze
PASSIFLORACEAE
Passiflora pottiae Cervi & Imig
PHYLLANTHACEAE
Hieronyma alchorneoides Allemão
Phyllanthus
Webster
stipulatus
(Raf.)
M 364
G.L.
M 203
PIPERACEAE
Piper fuligineum Kunth
Piper macedoi Yunck.
M 36
-
PLANTAGINACEAE
Bacopa monnierioides (Cham.) B.L. Rob.
M 41
0.03
0.31
0.34
Bacopa reflexa (Benth.) Edwall
M 213
0
0.16
0.16
Bacopa scabra Descole & Borsini
M 205
0.05
0.25
0.3
Andropogon bicornis L.
M 137
0.06
0.12
0.18
Andropogon glaziovii Hack.
M 221
Andropogon hypogynus Hack.
M 89
3.58
3.55
7.13
3.57
3.59
7.16
Andropogon macrothrix Trin.
M 104
0.24
0.12
0.36
0.39
0.4
0.79
0.02
0.13
0.15
POACEAE
12.96
4.42
17.38
Appendix 1. Continued on next page...
794 | Vegetation structure of vereda
Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued
Appendix 1. ...Continued
Wet grassland
Family/Species
Col.
Rainy
Transition Area
Dry
Rainy
Dry
RC
RF
CV
RC
RF
CV
RC
RF
CV
RC
RF
CV
P 10058
3.87
4.78
8.65
4.91
6.24
11.15
0.65
1.32
1.97
0.74
1.88
2.61
Anthaenantia lanata (Kunth) Benth.
M 334
9.08
4.41
13.49
6.88
4.65
11.53
Anthaenantiopsis trachystachya (Nees)
Mez ex Pilg.
M 56
0.78
0.37
1.15
0.75
0.4
1.15
0.12
0.16
0.28
Arundinella hispida (Humb. & Bonpl. ex
Willd.) Kuntze
M 24
0.04
0.13
0.17
Axonopus cf. comans (Trin. ex Döll)
Kuhlm.
P 10.777
0.96
0.66
1.62
7.33
3.76
11.09
Andropogon virgatus Desv.
M 19
5.02
1.47
Axonopus purpusii (Mez) Chase
M 20
Axonopus siccus (Nees) Kuhlm.
M 20
1.74
0.86
2.6
Coelorhachis aurita (Steud.) A. Camus
M 60
0.29
0.61
0.9
0.27
0.66
Eriochrysis cayennensis P. Beauv.
M 17
0.47
1.35
1.82
1.22
Eriochrysis holcoides (Nees) Kuhlm.
M 101
14.83
8.21
23.04
Hyparrhenia bracteata (Humb. & Bonpl.
ex Willd.) Stapf
M 189
0.58
0.98
1.56
Hyparrhenia rufa (Nees) Stapf
M 361
Ichnanthus procurrens (Nees ex Trin.)
Swallen
M 22
0.18
0.49
0.67
0.15
0.4
0.55
Imperata tenuis Hack.
M 208
1.92
1.23
3.15
1.62
0.93
2.55
Leersia hexandra Sw.
M 259
Paspalum dedeccae Quarin
M 343
Paspalum erianthoides Lindm.
M 303
9.65
2.57
Paspalum erianthum Nees ex Trin.
M 304
0.02
0.12
6.49
12.36
6.24
9.88
3.26
13.14
0.14
0.16
0.29
0.93
0.43
0.7
1.12
0.31
0.75
1.06
1.99
3.21
0.44
0.7
1.14
1.38
1.41
2.79
2.05
1.73
3.78
1.86
1.79
3.65
3.53
2.16
5.69
0.37
0.66
1.03
0.61
0.78
1.39
0.41
0.75
1.16
0.08
0.08
0.09
0.1
8.05
3.65
11.7
10.47
5.26
15.73
2.49
2.87
5.36
0.69
3
3.69
9.68
2.87
12.55
10.26
3.57
13.83
18.6
6.22
4.38
10.6
12.22
11.32
3.06
14.38
0.14
2.27
2.13
4.4
Appendix 1. Continued on next page...
Moreira et al. | 795
Oecol. Aust. 23(4): 776–798, 2019
Axonopus uninodis (Hack.) G.A. Black
Wet grassland
Family/Species
Col.
Rainy
Transition Area
Dry
Rainy
Dry
RC
RF
CV
RC
RF
CV
RC
RF
CV
RC
RF
CV
6.7
3.45
10.15
0.63
0.62
1.25
0.53
0.66
1.19
0.08
0.16
0.24
0.04
0.09
0.13
Paspalum flaccidum Nees
M 285
8.99
3.92
12.91
Paspalum glaucescens Hack.
M 365
1.29
1.23
2.52
Paspalum lenticulare Kunth
M 327
Paspalum maculosum Trin.
M 299
Paspalum wrightii Hitchc. & Chase
M 355
Rheochloa scabriflora Filg.. P.M. Peterson
& Y. Herrera
M 330
0.06
0.49
0.55
0.19
0.66
0.85
0.01
0.16
0.16
0.09
0.28
0.37
Rhytachne rottboellioides Desv.
M 90
0.6
1.47
2.07
1.31
0.53
1.84
3.11
1.94
5.05
24.37
9.95
34.32
Saccharum asperum (Nees) Steud.
M 61
1.38
0.37
1.75
0.12
0.13
0.25
16.36
5.9
22.26
1.41
2.25
3.66
Saccharum villosum Steud.
M 73
0.28
0.25
0.53
2.97
2.52
5.49
5.22
3.42
8.63
3.71
3.85
7.55
Sacciolepis vilvoides (Trin.) Chase
M 140
3.87
3.92
7.79
0.11
0.4
0.51
Schizachyrium condensatum (Kunth)
Nees
P 10.787
0.01
0.12
0.13
0.4
0.93
1.33
0.1
0.47
0.56
0.05
0.28
0.33
Schizachyrium gracilipes (Hack.) A.
Camus
M 340
0.21
1.17
1.38
0.06
0.66
0.72
Setaria parviflora (Poir.) Kerguélen
M 1278
0.1
0.16
0.25
0.02
0.09
0.12
0.05
0.19
0.24
0.01
0.09
0.1
Steinchisma decipiens (Nees ex Trin.)
W.V. Br.
M 93
Steinchisma laxum (Sw.) Zuloaga
M 18
Trichanthecium
caaguazuense
(Henrard) Zuloaga & Morrone
M 157
Trichanthecium parvifolium
Zuloaga & Morrone
(Lam.)
M 128
0.11
0.49
0.6
0.07
0.39
0.46
0.05
0.39
0.44
0.02
0.7
0.72
Appendix 1. Continued on next page...
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Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued
Appendix 1. ...Continued
Wet grassland
Family/Species
Col.
Rainy
RC
RF
Transition Area
Dry
CV
RC
RF
Rainy
CV
Dry
RC
RF
CV
RC
RF
CV
0.06
0.16
0.21
0.04
0.09
0.13
0.29
0.54
0.84
0.4
0.47
0.87
0.14
0.78
0.92
0.02
0.19
0.21
0.02
0.16
0.17
1.01
1.55
2.56
0.08
0.56
0.64
PONTEDERIACEAE
Pontederia parviflora Alexander
M 368
PRIMULACEAE
Anagalis minima (L.) E.H.L. Krause
M 42
0.07
0.27
0.34
2.17
2.13
4.3
PTERIDACEAE
Pityrogramma calomelanos (L.) Link
M 224
1.72
2.7
4.42
M 168
0.04
0.12
0.16
RUBIACEAE
Borreria
pulchristipula
Bacigalupo & E.L. Cabral
(Bremek.)
Hexasepalum radula (Willd.) Delprete &
J.H. Kirkbr.
M 92
SAPINDACEAE
Matayba elaeagnoides Radlk.
M 338
SCROPHULARIACEAE
M6
0.02
0.13
0.15
SOLANACEAE
Schwenckia juncoides Chodat
Solanum subinerme Jacq.
M 55
P 10.552
STYRACACEAE
Styrax camporum Pohl
M 65
Appendix 1. Continued on next page...
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Melasma stricta (Benth.) Hassl.
Wet grassland
Family/Species
Col.
Rainy
Transition Area
Dry
Rainy
Dry
RC
RF
CV
RC
RF
CV
RC
RF
CV
RC
RF
CV
M 88
0.14
0.25
0.39
0.06
0.13
0.19
0.04
0.16
0.2
0.06
0.09
0.16
Xyris jupicai Rich.
M 348
0.06
0.12
0.18
1.27
2.39
3.66
0.21
0.47
0.68
0.13
0.38
0.51
Xyris laxiflora F. Muell.
M 126
0.47
0.74
1.21
0.84
1.06
1.9
0.62
1.71
2.33
0.41
1.03
1.44
Xyris savanensis Miq.
M 351
1.74
1.96
3.7
0.11
0.53
0.64
Xyris schizachne Mart.
M 353
0.04
0.12
0.16
0.08
0.39
0.47
0.05
0.28
0.34
Xyris tortula Mart.
M 350
0.01
0.12
0.13
0.12
0.23
0.36
0.08
0.19
0.26
URTICACEAE
Cecropia pachystachya Trécul
XYRIDACEAE
798 | Vegetation structure of vereda
Oecol. Aust. 23(4): 776–798, 2019
Appendix 1. ...Continued