doi:10.21829/myb.2017.

2311527
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Damming water influences the structure,

composition and functions of
adjacent savannahs
Águas represadas influenciam a estrutura, composição e funções de
savanas adjacentes

Vagner Santiago do Vale1*, Ana Paula de Oliveira2, Jamir Afonso do Prado-Junior3, Patrícia Ribeiro Londe4 and
Diego Raymundo Nascimento5

1 Doutor em Ecologia e Conservação de Recursos Na- 2 Doutora em Ecologia e Conservação de Recursos 3 Doutor em Ecologia e Conservação de Recursos Na-
turais, UFU. Naturais, UFU. Instituto de Ciências Biológicas e da turais, UFU. [email protected]
5 Mestrando em Ecologia, UFJF. diegoraynascimen- Saúde, UFV. [email protected] 4 Mestre em Geografia, UFU. [email protected]
[email protected] * Corresponding author. [email protected]

Abstract
Dams are important to human wellbeing; however, they can cause various environmental changes. In the cerrado stricto sensu, waters
from the reservoir modify water supply in the soil and can affect species establishment. It is argued that species adapted to high water
availability can establish close to river banks and water reservoirs and their characteristics are different from those of species from the
cerrado stricto sensu. These characteristics could be more similar to those of species from riparian forests, with higher values of height
and basal area. Thus, we aim to analyze changes in an area of the cerrado stricto sensu, 35 years after the construction of a hydroelectric
dam. We tested different parameters to determine these changes, including richness, diversity, individual height and basal area (using the
square-point method), with individuals <1 m, individuals >1 m, and Bromeliaceae (using the plot method) in three sectors close to the
artificial lakeshore (0 m - 20 m, 40 m - 60 m, and 80 m - 100 m away from the water). Using species diversity, we also compared the
species functional group within these sectors. Sectors closer to the dam acted as an edge, presenting lower basal area and individual’s
height, but species with different traits compared to those of a cerrado stricto sensu area.

Keywords: biomass accumulation, cerrado stricto sensu, tree community, hydroelectric energy, phytosociology, functional groups,
functional traits.

Resumo
Represas são importantes para o bem-estar humano, no entanto, podem causar diversas alterações nos ambientes. No cerrado stricto
sensu, as águas represadas modificam o abastecimento de água no solo e podem alterar o estabelecimento das espécies. Argumenta-se
que as espécies adaptadas à alta disponibilidade de água podem estabelecer-se próximo às margens de rios e reservatórios e possuem
características distintas daquelas de espécies de cerrado stricto sensu. As características seriam mais semelhantes àquelas de espécies de
matas ciliares, com elevada estatura e área basal. Assim, almejamos analisar as mudanças em uma área de cerrado stricto sensu após 35
anos sob a influência de uma usina hidrelétrica. Testamos parâmetros distintos para determinar essas mudanças, tais como a riqueza,
a diversidade, altura dos indivíduos e área basal (utilizando o método do ponto quadrante), com indivíduos < 1 m, indivíduos > 1 m e
Bromeliaceae (utilizando o método de parcelas) em três setores próximo à margem da represa (a 0 m - 20 m, 40 m - 60 m e 80 m - 100
m da represa). Utilizando a diversidade de espécies, também comparamos os grupos funcionais de espécies nesses setores. Os setores
mais próximos à represa possuíam baixa área basal e altura dos indivíduos atuando como uma borda, mas com espécies com diferentes
características em relação ao cerrado stricto sensu.

Palavras chave: acumulação de biomassa, cerrado stricto sensu, comunidade arbórea, energia hidroelétrica, fitossociologia, grupos
funcionais, traços funcionais.

63
 Vale et al. Damming water influences adjacent savannahs

Introduction functional group is a set of species with similar characteris-

Anthropogenic impacts can lead to a structural evolution tics or that present a similar use of resources (Lyon and
from open to closed vegetation, with increased biomass Sagers, 2003; Médail, Roche and Tatoni, 1998). The spe-
reported in various areas of the Cerrado (Moreira, 2000). cies within a functional group play a similar role within the
Variations in water availability due to the presence of community or the ecosystem (Blondel, 2003; Vale et al.,
dams (Nilsson and Bergreen, 2000, Vale et al., 2013a) 2013b). Classification into functional groups allows the
and fires constitute examples of anthropic disturbances gathering of a large number of species into a small group of
that affect plant communities (Hoffman and Solbrig, functional types that present the same responses to distur-
2003). In seasonal environments, such as the Cerrado, bances or similarities in terms of processes of dispersion,
these impacts can determine the predominant physiog- competition and survival (Hubbel, 2005; Skov, 2000).
nomy and the proportion that exists between open (savan- To achieve a good representation of ecological roles,
nahs) and closed (forests) areas (Marimon Júnior and we must choose traits that represent important functions
Haridasan, 2005; Pinheiro and Durigan, 2009; Lopes, relating to the ecosystems (Vale et al., 2011). However,
Vale and Schiavini, 2009); open areas have less water and plants have many attributes that may be used for identi-
experience more fires while closed areas have more water fying functional groups (Pendry et al., 2007) and the
and a lower incidence of fires. In Brazil, such open savan- choice of such attributes must be clearly related to the role
nahs are called “South American Savannahs” or played by the species within the ecosystem. In this study,
“Cerrado” and are very similar structurally to the African we focus on attributes that may represent the impacts cau-
savannahs (Dexter, 2015), with twisted trees with thick sed by the presence of a dam. We therefore consider attri-
bark and a grass stratum that occupies an important por- butes related to local animal-plant interactions, such as
tion of the vegetation, although there is variation in the pollination and dispersion, since these vary according to
floristic patterns. Here, we focus on the impact of a dam the availability of water and the presence of fauna and
on an open cerrado plant community that became a clo- flora within an ecosystem (Nilsson, 1996).
sed forest after 35 years in proximity to the artificially Various animal groups can occupy different positions
water created by the reservoir. in the vertical space formed by vegetation. Birds, mam-
Changes from open to closed areas are particularly mals (Clark, Poulsen and Parker, 2001) and invertebrates
important because it is not only structural aspects (such as (Gossner and Muller, 2011) may prefer to occupy different
number of trees, basal area and individual height) that are vertical layers in a forest, for instance. This preference
modified, but also ecological functions, such as the provi- may be connected to the direct attributes of the tree com-
sion of resources for fauna and carbon assimilation (Vale, munity, such as the abundance of resources, temperature,
Schiavini, Oliveira and Gusson, 2010). In protected areas, leaf density, and luminosity (Campbell and Donato, 2014;
such changes can alter the primary protective purpose O’Hanlon and Holwell, 2011; Walther, Clayton and Gre-
related to the local diversity of fauna and flora, as well as gory, 1999).
influence the various biological roles played by animal Growth responses under high luminosity and morta-
and plant species. lity under shade are particularly important in forests (Pear-
These biological roles are related to the occurrence of son, Burslem, Goeriz and Dalling, 2003), but light is not
species and their attributes (Walker, Kingzig and Lan- restricted in open areas. Furthermore, deciduousness can
gridge, 1999). Many species can thus play similar roles interfere with luminosity at ground level, which can become
within the community and the ecosystem (Hooper and more marked in soils that present a significant hydric deficit
Dukes, 2004; Walker et al., 1999). These species that share during dry periods. Furthermore, species from deciduous
the same attributes may belong to “functional groups”. A forests currently present rapid growth due to high carbon

64
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

assimilation (Cornelissen, Diez and Hunt, 1996). On the cerrado area after 35 years under the influence of a
other hand, evergreen species tend to produce and keep hydroelectric dam. The main objective was to understand
their leaves throughout the year in milder or moist environ- the impacts of the dam on the savannah communities.
ments, thus hindering the establishment of species that This area was previously far from any water source, but
demand direct light below the canopy of the trees. Another currently lies close to a large body of water. Thus, in 2009,
important feature is tree-bark thickness; this property pro- the entire open cerrado border became forest; however, it
vides savannah species with an increased resistance to fire still preserves some cerrado characteristics, mainly at a
compared to forest species (Hoffman and Solbrig, 2003). distance from the lakeshore.
We cannot disregard this trait, because forest areas often
border the savannahs and fire events can thereby favor the Material and Methods
savannah species (Hoffman and Solbrig, 2003).
Finally, in environments with low water availability, Study area and sampling methods
seed desiccation tolerance can play a key role for species sur- This study was carried out in a forest fragment in the state
vival over time. Seeds that are more tolerant to desiccation of Minas Gerais, southeastern Brazil. The climate of the
are able to survive under conditions of lower humidity zone is characterized by warm temperatures throughout
without losing their viability when hydrated. Thus, the spe- the year with a rainy summer and dry winter, i.e. Aw,
cies that present this feature can constitute a persistent seed according to Köppen (1948). The forest is located at the
bank. In contrast, seeds that are sensitive to desiccation quic- “Pirapitinga Ecological Station” (ESEC – Fig. 1), close to
kly lose their viability, although they can constitute a saplings the Três Marias Dam, in Minas Gerais, Brazil. Damming
bank in environments with high soil moisture (Tweddle, took place in 1962 and ESEC became an island (Gonçalves-
Dickie, Baskin and Baskin, 2003). Alvim and Fernandes, 2001). Flood elevation varies from
It is difficult to understand the consequences of proxi- 559 m to 568 m and the area is around 1090 ha (Azevedo,
mity to the water line on savannah drought-adapted spe- 1987). At present, the forest canopy is rarely >10 m height
cies, and it also remains unknown whether ecosystem and there are many Bromeliaceae on the ground. Species of
functions will be lost or change after many years of dams Bromeliaceae are usually high light-demanding species that
impacts. Furthermore, dams constructions are increasing cover large areas on the ground, and clearly affect the esta-
all around the world, and we must understand the changes blishment of other plant species. Far from the water, there
in vegetation after damming in order to apply more appro- is a transition between the forest and a cerrado vegetation.
priate conservation and management actions in the future. Thus we divided the forest in three sectors: 1) “close”, i.e. 0
We assumed three hypotheses: 1) the sector close to the m - 20 m from the dam; 2) “middle”, i.e. 40 m - 60 m from
dam will have more forest species, with higher basal area the dam, but still about 50 m away from the cerrado stricto
and individual tree heights; 2) due to the presence of more sensu area; and 3) “transition”, 80 m - 100 m from the dam
closed vegetation, the sector close to the dam will have less and closer to the typical cerrado vegetation.
small plants; 3) the influence of the dam will change the We used the square-point method (Cottam and Cur-
ecological roles within this community, with more marked tis, 1956) for sampling trees. This method consists of sam-
effects on sectors in more close proximity to the water. pling the four first trees close to the square point. Only
trees with circumference at breast height (CBH) ≥15 cm
Objectives were included. We sampled 90 points arranged into 6
Considering the structural and functional importance of parallel transects located 100 m apart, for a total of 360
open (in this case, cerrado stricto sensu) and closed (in sampled trees. In each transect, we allocated 15 square
this case, forest) areas, we aim to evaluate changes in a points, 5 close to the forest border (“close”), 5 in the

65
 Vale et al. Damming water influences adjacent savannahs

Figure 1. Location of the Pirapitinga Ecological Station (ESEC), which became an island with damming of the area 35 years ago, State
of Minas Gerais, Brazil. Source: Giácomo, 2009.

middle forest (“middle”), and 5 in a transition site with Tree community functional analysis
cerrado stricto sensu (“transition”). Each set of 5 points in This analysis was performed for all tree species found
the same sector was regarded as 1 sample. within the three sectors. For each species, three physiolo-
We measured all of the young trees (CBH <15 cm), gical traits were evaluated: 1) shade tolerance, 2) deci-
seedlings and saplings in 75 plots (2 m x 2 m); these indi- duousness and 3) seed desiccation tolerance. In addition,
viduals were regarded as a regeneration layer. Regenera- two structural traits were considered: 4) vertical strata;
ting individuals were classified into 2 categories: 1) and 5) bark thickness. Finally, there were 2 animal-plant
seedlings, saplings, and young trees <1 m height; and 2) interaction types: 6) dispersal syndrome; and 7) pollina-
young trees ≥1 m height. The number of species of Brome- tion syndrome (Table 1). We obtained information rela-
liaceae was determined in these plots. To compare these ting to the ecological attributes from scientific papers and
structural parameters, we used Kruskall-Wallis and post specialized books, as well as through field observation
hoc Median tests, both non-parametric tests, since the and consulting specialists.
data did not comply with the assumptions of normality. A matrix with the tree species and 25 traits was thus
We calculated the richness and density of the arbo- adopted for analysis. We converted data to a presence/abs-
real and regenerating strata within the “close”, “middle”, ence matrix with species and their ecological attributes.
and “transition” sectors. We compared the basal area and An ecological distance matrix was then determined (Pet-
individual tree heights between these sectors using analy- chey and Gaston, 2002). The functional groups were
sis of variance (ANOVA), followed by an a posteriori revealed by two analysis types, using the distance matrix
Tukey test with a 5% significance level (Wilkinson, 2002). and a “cluster analysis”, as explained below.

66
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Table 1. Traits and their respective ecological functions used for obtaining the distance matrix.

Trait Classification Categories Ecological function

Dispersal syndrome1 Diaspores dispersed by wind Anemochory Long distance dispersion, little affected by
fragmentation
Free fall or ballistic mechanisms Autochory Short distance dispersion, highly affected by
fragmentation
Birds Ornitochory Medium distance dispersion, affected by frag-
mentation and food resource for the birds
Mammals Mammaliochory Short distance dispersion, highly affected
by fragmentation and food resource for the
mammals
Deciduousness Species in which all leaves fall Deciduous High litter release and canopy openness to
within the dry season other plants
Remain throughout the year Evergreen Shade and refuge for animals and provides
other plants with increased shading
Shade tolerance Light environment required for
establishment
Tolerates shade under other trees Shade tolerant Low growing species and climax species;
related to forest maturity
Needs direct sunlight to develop Light demanding Fast growing species; generally represents
early regeneration stages
Vertical strata Based on the stature commonly
reached by adult individuals
Always reach the canopy Typical canopy Greater biomass accumulation when compa-
red to smaller species from the shade below
the canopy
Sometimes reach the canopy Intermediate Integrate canopy and understory; it may repre-
sent various strata
Usually do not reach the upper Typical understory Low basal area, but high density species; this
forest layers is a different habitat for animals
Seed desiccation Related to seeds remaining in situ
tolerance until next season
Seeds able to survive until next Orthodox Forms seed banks in the forest
season
Seeds unable to survive until next Non-orthodox Forms a “seedling bank”, but not a seed bank
season
Pollination Species were classified by animal
syndrome 2
pollinator
Body size < 5 mm Very small insects Promotes genetic variability, and provides very
small insects with food resource
Small bees, wasps, and flies (body Small insects Promotes genetic variability, and provides
size 5-12 mm) bees, wasps, and flies with food resource

67
 Vale et al. Damming water influences adjacent savannahs

Table 1. Traits and their respective ecological functions used for obtaining the distance matrix. Continuation...

Trait Classification Categories Ecological function

Bees with large size body (> 12 mm) Big bees Promotes genetic variability, and provides big
and isolated bees with food resource
Beetles Beetles Promotes genetic variability. and provides
beetles with food resource
Butterflies and moths Lepidoptera Promotes genetic variability, and provides
moths and butterflies with food resource
Bats Bats Promotes genetic variability, and provides
bats with food resource
Bark thickness Species were classified by stem
protection against fire
Stem with high bark thickness Highly protected Prevents against fire damage. Individuals can
stem persist after a fire event
Rough stem Moderately protec- Species can undergo fire damage and face
ted stem negative effects in reproductive terms
Stem without bark protection Little protected Cannot prevent fire damage and the trees may
stem die following a fire event
1 = Pijl (1982); 2 = Faegri and Pijl (1979).

Cluster analysis test. The same procedure was adopted to address the
The groups were defined by Euclidian distance and a abundance of species traits at all three sectors.
dendrogram generated using the group average. Multi-
variate clustering methods were used to determine Results
which plant species groups could emerge. These We found 57 species, 39 close to the dam, 38 in the middle
methods calculate similarity or measurements of asso- sector and 29 in the transition site in the cerrado area
ciation, based on the extent to which species share (Table 2); however, only 18 species occurred in all three
attributes (Leishman and Westoby, 1992). Cophenetic sectors. We observed 10 exclusive species in the sector
correlation was calculated for the generated dendro- close to the dam, in the middle sector there were 9 exclu-
gram; since this is equivalent to Pearson’s coefficient, sive species but only 2 species were exclusive to the transi-
we proposed it as a concordance measure between the tion sector. Species abundance also varied between the
obtained group and the original distance matrix (Bus- sectors. Many species were more abundant in one sector.
sab and Morettin, 1990). All these analyses were per- Alibertia edulis and Qualea parvifolia were much more
formed using the software Fitopac-Shell, version 1.6.4 abundant close to the dam. Siparuna guianensis, a typical
(Shepherd, 2004). forest understory species, was the most abundant species
Once the functional groups were found, we perfor- in the middle sector, presenting few individuals in other
med analysis of variance to test differences with regard to sectors. Vatairea macrocarpa and Qualea grandiflora,
the abundance of functional groups within the three sec- two typical cerrado species, had more individuals in the
tors (close, middle and transition), followed by a Tukey transition site in the cerrado (Table 2).

68
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Table 2. Species list and their abundance in a cerrado physiognomy at the “Pirapitinga Ecological Station”, Minas Gerais, Brazil.

Species Close to the dam Middle Transition Total Group

Alibertia edulis (Rich.) Kuntze 8 2 2 12 G5
Annona crassiflora Mart. - 1 1 2 G1
Aspidosperma macrocarpon Mart. 1 - 3 4 G2
Aspidosperma tomentosum Mart. 6 2 6 14 G2
Astronium fraxinifolium Schottex Spreng. 4 7 5 16 G3
Bowdichia virgilioides Kunth 2 - 1 3 G3
Byrsonima coccolobifolia Kunth 1 2 3 6 G1
Byrsonima crassa Nied. 1 1 3 5 G1
Brosimum gaudichaudi Trécul - 1 - 1 G1
Cecropia pachystachya Trécul* 1 - - 1 G4
Copaifera langsdorffii (Desf.) Kuntze 1 2 - 3 G3
Dimorphandra mollis Benth. - 1 - 1 G5
Diospyros sericea A. DC. 5 4 4 13 G4
Dipteryx alata Vogel - 1 - 1 G4
Emmotum nitens (Benth.) Miers - 1 - 1 G4
Eriotheca pubescens (Mart. & Zucc.) Schott & Endl. 12 13 5 30 G3
Stenocalyx dysentericus (DC.) O. Berg 6 1 4 11 G1
Hancornia speciosa Gomes 1 - - 1 G1
Heteropterys byrsonimifolia A. Juss. 1 6 3 10 G2
Hirtella gracilipes (Hook. F.) Prance 1 - - 1 G4
Hymenaea stigonocarpa Mart. Ex Hayne 1 - - 1 G1
Inga fagifolia (L.) Willd. ex Benth. 1 - - 1 G5
Inga edulis Mart. 2 1 - 3 G5
Machaerium opacum Vogel 1 1 1 3 G3
Magonia pubescens A. St.-Hil 11 8 20 39 G2
Myrcia tomentosa (Aubl.) 2 - - 2 G5
Ouratea hexasperma (A. St.-Hil.) Baill. - - 1 1 G1
Pera glabrata (Schott) Poepp. ex Baill.* 1 1 - 2 G4
Plathymenia reticulata Benth. 1 - 3 4 G3
Plenckia populnea Reissek - - 1 1 G1
Pouteria ramiflora (Mart.) Radlk. 1 3 3 7 G1
Protium heptaphyllum (Aubl.) Marchand 2 3 1 6 G4
Pseudobombax longifolium - 2 1 3 G3
Psidium pohlianum (O. Berg) Landrum 1 - 1 2 G1
Pterodon pubescens Vogel 1 4 - 5 G3

69
 Vale et al. Damming water influences adjacent savannahs

Table 2. Species list and their abundance in a cerrado physiognomy at the “Pirapitinga Ecological Station”, Minas Gerais, Brazil. Continuation...

Species Close to the dam Middle Transition Total Group

Qualea grandiflora Mart. 2 5 10 17 G2
Qualea multiflora Mart. 3 2 4 9 G3
Qualea parviflora Mart. 6 1 - 7 G2
Roupala Montana Aubl. 1 2 1 4 G2
Rudgea viburnoides (Cham.) Benth. - 1 - 1 G5
Tachigali vulgaria Harms - 3 1 4 G3
Siparuna guianensis Aubl.  7 16 9 32 G4
Siphoneugena densiflora (Krug & Urb.) Proença* 1 - - 1 G4
Styrax ferrugineus Ness & Mart. - 1 - 1 G1
Strychnos pseudoquina A.St.-Hil. - 2 - 2 G1
Stryphnodendron adstringens (Mart.) Coville 1 1 - 2 G1
Tabebuia aurea (Silva Manso) Benth. & Hook. f. ex S. Moore 1 - 1 2 G3
Handroanthus ochraceus (Cham.) Mattos - 1 - 1 G3
Tapirira guianensis Aubl.* 1 - - 1 G4
Terminalia argêntea Mart. 3 - - 3 G3
Vatairea macrocarpa (Benth.) Ducke - 1 9 10 G3
Virola sebifera Aubl. - 1 - 1 G4
Vochysia cinnamomea Pohl. 1 - - 1 G2
Vochysia rufa Mart. - 1 - 1 G2
Xylopia aromatica (Lam.) Mart. 17 14 13 44 G4

* Species usually found in forest with abundant water resources.

A pattern was found in terms of basal area (F = 4.32, formation. The 5 groups clearly have distinct functions
p = 0.015) and individual heights (F = 9.82, p < 0.001): (Table 2). Two groups present typical savannah species
these structural parameters were higher in the middle sec- traits (G1 and G2): they are short (up to 6 m in height),
tor (Fig. 2A and 2B). In the regeneration layer, the density deciduous species (a feature typical of seasonal environ-
of plants >1 m height did not show a difference (Fig. 3B; H ments), present prior pollination by small insects, moths
= 7.02, p = 0.30), however, for individuals <1 m height (H and large bees (well-known pollinators in cerrado areas).
= 26.96, p = 0.002) and bromeliad species (H = 14.37, p Most species from both groups also have stems that are
= 0.008) the lowest values occurred in the sector close to protected against fire. These groups basically differ in
the dam (Fig. 3A and 3C). terms of dispersion: Group G1 provides mammals and
birds with food resources while G2 is dispersed by wind.
Tree functional groups Few species from G1 are non-orthodox, meaning that this
The cluster forms 5 groups (Figure 4) and the pollination group can form both seed and seedling banks within the
syndrome was the only species trait not useful for group rainy season.

70
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Figure 2. Analysis of variance of basal area (A) and height (B) in three sectors of a cerrado area at the “Pirapitinga Ecological Station”,
Minas Gerais, Brazil. Lower case letters denote the results of a post hoc Tukey test; different letters (a and b) represent significant
differences (p <0,05).

Figure 3. Kruskall-Wallis test to determine the number of individuals above and below 1 m in height (A and B, respectively), as well
as the number of Bromeliaceae (C) in the three sectors of a cerrado area at the “Pirapitinga Ecological Station”, Minas Gerais, Brazil.
Lower case letters (a, b and c) represent the results of median test. Different letters represent significant differences (p < 0,01).

The other three groups are formed by traits more cies group with marked deciduousness and dispersion by
typical of forest species; however, they are clearly distinct. wind. All species have desiccation tolerance and they can
G3 and G4 are formed by taller species (usually > 10 m), form a permanent seed bank. Most species from G3 have
but G5 has strictly small trees. G3 is the only forest spe- high protection against fire, provided by thick trunks and

71
 Vale et al. Damming water influences adjacent savannahs

barks. This group may be regarded as the most protected

in case of a fire event, since, in addition to the thickness of
the trunks and bark, its individuals are tall.
G4 is also a tall group; however, its individuals do
not have a marked protection against fire. This may be
regarded as the group most different from savannah in the
forest, since it consists of many shade-tolerant species.
Some species from this group, such as Protium heptaphy-
llum, Siparuna guianensis, Ocotea corymbosa, Pera gra-
blata, and Tapirira guiannesis, are also observed in moist
forest physiognomies. In this way, the group clearly repre-
sents the influence of the dam by considering the former
cerrado vegetation. This group consists of evergreen spe-
cies and provides food resources to the fauna, especially to
bird species. This is the only group that presents pollina-
tion by very small insects; however, most pollination is
provided by small insects, just like the other groups. This
group is also non-orthodox and can form a seedling bank Figure 4. Functional groups based on ecological distance
within the rainy season. formed by clustering in a cerrado forest in southeastern Brazil.
G5 also has non-orthodox species; however, these The cluster was defined by Euclidian distance and group average.
differ from those of G4, since they provide mammals and Groups characteristics are shown in Table 2.
birds with food resources. This group may be highly
affected by fire, because it consists of small species with
no significant protection against fire. It is also the smallest as cerrado stricto sensu (Giácomo, 2009), presented forest
group, with only 7 species - most of which are shade-tole- characteristics due to the establishment of the many forest
rant species from the moist forest understory. species that currently share the space with the cerrado
Comparing the groups formed close to the dam and species. Floristic composition is still quite similar to that
in the middle sector, there is a lower number of short trees of cerrado vegetation, and includes species from cerrado
(G1 and G2 - groups of savannah species), a high number stricto sensu, such as Qualea parviflora, Eriotheca pubes-
of tall forest trees (G3 and G4) and the highest number of cens, Magonia pubescens, and Xylopia aromatica (Rat-
short forest trees from the forest understory (G5) close to ter, Bridgewater and Ribeiro, 2003; Lopes, Vale, Oliveira
the dam (Fig. 5). The opposite occurs in the transition and Schiavini, 2011). However, species from moist soils
sector, which has a higher number of cerrado species, and were also observed, such as Alibertia edulis and Protium
the lower number of forest species, from groups 3 and 4. heptaphyllum. Even small changes in the water regime
In the middle sector, the predominant groups were G4 can induce changes in vegetation structure (Gusson, Vale,
and G3 (tall forest species) with few G1 and G2 trees Araújo and Schiavini, 2012; Vale et al., 2013a) and the
(savannah species). dam clearly favors water-associated species, such as those
observed in other dams (Vale et al., 2013a). Some recent
Discussion studies have shown that richness can increase over time,
The influence of the dam on the cerrado vegetation after due to the establishment of moisture-associated associa-
35 years is clear; the tree community, previously regarded ted species in sectors near the lakeshore created by the

72
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

monly found in seasonal semideciduous forest (Lopes,

Schiavini, Oliveira and Vale, 2012), which is often asso-
ciated with streams, confirming that the influence of the
dam can reach a distance of 40-60 m from the water´s
edge.
Despite the high basal area, many small trees colo-
nize the middle sector and the large trees are distant from
each other, creating open spaces. The same occurs in the
transition sector, but the trees are smaller. These open
patches provide sunlight conditions and facilitate the esta-
blishment of small light-demanding plants (many <1 m
Figure 5. Number of trees found in three sectors at a forest in height). The occurrence of many Bromeliaceae on the
Três Marias Dam, Minas Gerais, Brazil. ground in the middle and transition sectors confirms the
Groups 1(white), and 2 (small squares) consist of cerrado species, G3 (light gray), high light incidence. Savannahs are considered open areas
G4 (dark gray), and G5 (black) consist of forest species.
and this feature means that these areas are still influenced
by their savannah history prior to construction of the
dam. Changes after damming are therefore focused on the
dam (Vale et. al., 2015). However, for the following three sector close to the dam, 0 to 20 m far from the river, at
reasons, this increase in species does not compensate for least for some vegetation structures.
those lost due to the dam construction: 1) species richness In the transition sector, the predominance of savan-
in dammed riparian forests is lower than that of non- nah species is clear; there are many individuals of species
dammed riparian forest (Liu, Liu, Liu, Song and Zhang, such as Qualea grandiflora and Magonia pubescens, and
2013); 2) the new species are different to those that exis- many other small trees, as well as large trees such as
ted in the area prior to construction of the dam (here the Vatairea macrocarpa. Small trees are related to forest
cerrado species are clearly fewer in sectors near the water edges (Lima-Ribeiro, 2008; Laurance et al., 2006) and
and 3) the flooded area destroys the past systems, and this both the sector close to the dam and the transition sector
loss is difficult to replace. constitute forest edges. However, one is influenced by the
Many of these species have thin trunks, are shade- dam and the other by the savannah vegetation. The
tolerant and from the forest understory. They are there- middle sector is thus protected to a greater extent from
fore usually short trees, such as A. edulis. Other studies border effects and consequently presents the biggest trees.
show that the bark thickness of cerrado species is three The larger number of small trees (< 1 m height) and
times greater than that of forest species (Hoffman and Bromeliaceae (light-demanding species on the ground) in
Solbrig, 2003), and the basal area is even lower. the transition sector implies an environment with more
The sector close to the dam may be regarded as an sunlight, in contrast to the sector close to the dam, which
edge where the wind can topple big trees, decreasing the is a shaded environment. Savannah plant communities are
total basal area (Laurance et al., 2006). The middle sector open areas without light limitation unlike the forest,
contained more typical canopy species, such as Astronium where a large amount of light is intercepted by the species
fraxinifolium, Pterodon pubescens, and Copaifera lans- that form the canopy (Poorter, Bongers and Bongers,
dorffii, compared to the other sectors. This sector there- 2006). Moreover, high soil moisture can kill seedlings and
fore presented the highest basal area and height of trees. young trees due to hypoxia or anoxia (Nilsson and Ber-
These canopy tree species in the middle sector are com- green, 2000), hindering plant establishment.

73
 Vale et al. Damming water influences adjacent savannahs

Perhaps the change from a cerrado physiognomy to a species, differing strongly in this respect from a savannah.
forest physiognomy is not only the result of the dam. The The damming influence was found at a distance of at least
ESEC is currently a protected area and fire, normally a 40-60 m from the artificial lakeshore and this was eviden-
common disturbance in cerrado areas, is infrequent. ced by the establishment of big trees and a well developed
When protected from fire events, some cerrado communi- understory, in contrast to the savannah which does not pre-
ties can turn into more closed environments and, over sent such a structure. Functionally, the tree species that
time, can become forests (Durigan and Ratter, 2006). colonized these sectors (up to 60 m in distance from the
These areas, for instance, had not undergone a fire event water) had thinner bark and thus less protection against
for 38 years (Durigan and Ratter, 2006), facilitating the fire, were taller and did not lose their leaves in the dry sea-
establishment of forest species, which would normally son, all of which are common traits for moist forest. Clearly,
have been destroyed by fire events in the absence of any many years of the dams influence has changed the commu-
form of timber protection. nity and converted an open cerrado phytophysiognomy
This change in functional group species (based on into a closed forest community, at least within a distance of
traits) not only suggests structural changes, but also chan- 60 m from the water´s edge and management of these areas
ges in ecological roles. Forests contain a large amount of must take this fact into full account.
animals that forage throughout the year for fruits and
pollen (Oliveira and Paula, 2001; Pinheiro and Ribeiro, Acknowledgments
2001; Vale et al., 2013b), whereas the production of fruits We thank the two anonymous reviewers for their helpful
and flowers in savannahs is concentrated in the late dry comments, which improved the manuscript and the Pro-
period (Oliveira and Paula, 2001) and resources for fauna grama de Concessão de Bolsa de Incentivo ao Pesquisador
may be insufficient during other periods. PROBIP (UEG) for financial support.
Due to the absence of fire and/or the presence of the
dam, this community clearly assumes the characteristics References
of a seasonal forest, since many seeds do not have desicca- Azevedo, L.G. (1987). Ensaio metodológico de identificação e
tion tolerance, there are evergreen plants, those without avaliação de unidades ambientais: a Estação Ecológica
bark protection and those dispersed by animals, as well as de Pirapitinga, MG. Brasília, DF: Embrapa.
shade-tolerant trees (groups 4 and 5 are the most repre- Blondel, J. (2003). Guilds or functional groups: does it matter?.
sentative of these traits). However, some cerrado characte- Oikos, 100(2), 223-231.
ristics still remain, such as deciduous small trees with Bussab, W. O. and Morettin, P. A. (1990). Estatística Básica.
large bark thickness, and this may be due to their savan- São Paulo: Atual.
nah history. With no fire events in the area, more forest Campbell, J. L., Donato, D. C. (2014). Trait-based approaches
species can establish (especially species from groups 4 and to linking vegetation and food webs in early-seral forest of
5), assisting the conversion into forest; however, a monito- the Pacific Northwest. Forest Ecology and Management,
ring survey would be required in order to provide further 324, 173-178.
data pertaining to this issue. Clark, C. J., Poulsen, J. R. and Parker, V. T. (2001). The role of
arboreal seed dispersal groups on the seed rain of a
Conclusion lowland tropical forest. Biotrópica, 33(4), 606-620.
After 35 years of influence of water from the dam, the Cornelissen, J. H. C., Diez, P. C. and Hunt, R. (1996). Seedling
cerrado stricto sensu has altered its structure, floral compo- growth, allocation and leaf attributes in a wide range of
sition and functional traits. Patches close to the dam water woody plant species and types. Journal of Ecology, 84(5),
now act as a forest edge and contain many water-associated 755-765.

74
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Cottam, G. and Curtis, J. T. Curtis. (1956). The use of distance Laurance, W. F., Nascimento, H. E. M. Laurance, S. G.,
measures in phytosociological sampling. Ecology, 37(3), Andrade, A., Ribeiro, J. E. L. S., Giraldo, J. P., Lovejoy, T.
451-460. E., Condit, R., Chave, J., Harms, K. E. and D’Angelo, S.
Dexter, K. G. (2015). Floristics and biogeography of vegetation (2006). Rapid decay of tree-community composition in
in seasonally dry tropical regions. International Forestry Amazonian forest fragments. Proceedings of the National
Review, 17(S2),10-31. Academy of Sciences of the United States of America,
Durigan, G. and Ratter, J. A. (2006). Successional changes in 103(50), 19010-19014.
cerrado and cerrado/forest ecotonal vegetation in western Leishman, M. R. and Westoby, M. (1992). Classifying plants
São Paulo state, Brazil, 1962-2000. Journal of Botany, into groups on the basis of associations of individual traits
63(1), 119-130. evidence from Australian semi-arid woodlands. Journal
Faegri, K. and Pijl, L. (1979). The principles of pollination eco- of Ecology, 80(3), 417-424.
logy (3rd ed.). Oxford: Pergamon Press. Liu, W. Liu, G., Liu, H., Song, Y. and Zhang, Q. (2013). Subtro-
Giácomo, R. G. (2009). Fitossociologia, Aporte de Serapil- pical reservoir shorelines have reduced plant species and
heira, Estoques de Carbono e Nitrogênio em Diferentes functional richness compared with adjacent riparian wet-
Formações Vegetais na Estação Ecológica de Pirapitinga lands. Environment Restoration Letters, 8,1-10.
– MG. Unpublished master’s thesis. Ciências Ambientais Lima-Ribeiro, M.D. (2008). Edge effects on vegetation and
e Florestais. Universidade Federal Rural do Rio de population structure in Cerradao fragments of
Janeiro. Southwest Goias, Brazil. Acta Botanica Brasilica, 22(2),
Gonçalves-Alvim, S. J. and Fernandes, G. W. (2001). Comuni- 535-545.
dades de insetos galhadores (Insecta) em diferentes fisio- Lopes, S. F., Vale, V. S. and Schiavini, I. (2009). Efeito de quei-
nomias do cerrado em Minas Gerais, Brasil. Revista madas sobre a estrutura e composição da comunidade
Brasileira de Zoologia, 18, 289-385. vegetal lenhosa do cerrado sentido restrito em Caldas
Gossner, M. M. and Muller, M. J. (2011). The influence of spe- Novas, GO. Revista Árvore, 33(3), 695-704.
cies traits and q-metrics on scale-specific beta-diversity Lopes, S. F., Schiavini, I., Oliveira, A. P. and Vale, V. S. (2012).
components of arthropod communities of temperate An Ecological Comparison of Floristic Composition in
forests. Landscape Ecology, 26, 411-424. Seasonal Semideciduous Forest in Southeast Brazil: Impli-
Gusson, A. E., Vale, V. S, Araújo, G. M. and Schiavini, I. (2012). cations for Conservation. International Journal of Fores-
Variações temporais na densidade de espécies lenhosas try Research, 2012. doi:10.1155/2012/537269
regenerativas em áreas impactadas por Usinas Hidrelétri- Lopes, S. F., Vale, V. S., Oliveira, A. P. and Schiavini, I. (2011).
cas. Revista de Biologia e Ciências da Terra, 12(1), 83-88. Análise comparativa da estrutura e composição florística
Hoffmann, W. A. and Solbrig, O. T. (2003). The role of topkill de Cerrado no Brasil Central. Interciencia, 36(1), 8-15,
in the differential response of savannah woody species to Lyon, S. and Sagers, C. L. (2003). Correspondence analysis of
fire. Forest Ecology and Management, 180, 273-286. functional groups in a riparian landscape. Plant Ecology,
Hooper, D. U. and Dukes, J. S. (2004). Overyielding among 164(2), 171-183.
plant functional groups in a long-term experiment. Eco- Marimon Junior, B.H. and Haridasan. M. (2005). Comparação
logy Letters, 7, 95-105. da vegetação arbórea e características edáficas de um
Hubbel, S.P. (2005). Neutral theory in community ecology and cerradão e um cerrado stricto sensu em áreas adjacentes
the hypothesis of functional equivalence. Functional Eco- sobre solo distrófico no leste de Mato Grosso, Brasil. Acta
logy, 19, 166-172. Botanica Brasilica, 19(4), 913-926.
Köppen, W. (1948). Climatología: con un estudio de los climas Médail, F., Roche, P. and Tatoni, T. (1998). Functional groups
de la Tierra. México: Fondo de Cultura Económica. in phytoecology: an application to the study of isolated

75
 Vale et al. Damming water influences adjacent savannahs

plant communities in Mediterranean France. Acta Oeco- Ribeiro, C.A.L. Fonseca and J.C Sousa-Silva. Cerrado:
logica, 19(3), 263-274. caracterização e recuperação de Matas de Galeria (pp.
Moreira, A. G. (2000). Effects of fire protection on savannah 335-378). Cerrados, Planaltina: Embrapa.
structure in Central Brazil. Journal of Biogeography, Poorter, L., Bongers, L. and Bongers, F. (2006). Architecture of
27(4), 1021-1029. 54 moist-forest tree species: Traits, trade-offs, and
Nilsson, C. (1996). Remediating river margin vegetation along functional groups. Ecology, 87(5), 1289-1301.
fragmented and regulated rivers in the North: What is Ratter, J.A., Bridgewater, S. and Ribeiro, J. F. (2003). Analysis
possible?. Regulated Rivers-Research & Management, of the floristic composition of the Brazilian cerrado vege-
12, 415-431. tation III: comparison of the woody vegetation of 376
Nilsson, C. and Berggren, K. (2000). Alterations of riparian areas. Edinburgh: Journal of Botany, 60(1), 57–109.
ecosystems caused by river regulation. Bio Science, 50(9): Shepherd, G. J. (2004). FITOPAC-SHELL (Preliminar version)
783-792. [Software]. Campinas: Universidade Estadual de Campinas.
O’Hanlon, J. C. and Holwell G. I. (2011). The influence of Skov, F. (2000). Distribution of plant functional attributes in a
abrupt forest edges on praying mantid populations. Insect managed forest in relation to neighbourhood structure.
Conservation and Diversity, 4(2), 107-114. Plant Ecology, 146(2), 121-130.
Oliveira, P. E. A. M. and Paula, F. R. (2001). Fenologia e biolo- Tweddle, J. C., Dickie, J. B., Baskin, C. C. and Baskin, J. M.
gia reprodutiva de plantas de Matas de Galeria. In J. F. (2003). Ecological aspects of seed desiccation sensitivity.
Ribeiro, C. A. L. Fonseca, J. C. Sousa-Silva. Cerrado: Journal of Ecology, 91, 294–304.
caracterização e recuperação de Matas de Galeria (pp. Vale, V. S., Schiavini, I., Oliveira, A. P. and Gusson, A. E.
303-328). Planatina: Embrapa Cerrados. (2010). When ecological functions are more important
Pearson, T. R. H., Burslem, D. F. R. P., Goeriz, R. E. and than richness: A conservation approach. Journal of Eco-
Dalling, J. W. (2003). Regeneration niche partitioning in logy and the Natural Environment, 2(12), 270-280.
neotropical pioneers: effects of gap size, seasonaldrought Vale, V. S., Schiavini, I., Prado-Junior, J. A., Oliveira, A. P. and
and herbivory on growth and survival. Oecologia, 137(3), Gusson, A. E. (2015). Rapid changes in tree composition
456-465. and biodiversity: consequences of dams on dry seasonal
Pendry, C. A., Dick, J., Pullan, M. R., Knees, S. G., Miller, A. forests. Revista Chilena de Historia Natural, 88, 1-13.
G., Neale, S. and Watson, M. F. (2007). In search of a Vale, V.S., Dorneles, M.C., Schiavini, I., Mendonça, E. T.,
functional flora-towards a greater integration of ecology Almeida, C. G., Silva P. A. and Crespilho, R. F. (2011).
and taxonomy. Plant Ecologic, 192(2), 161–167. Grupos funcionais e sua importância ecológica na vege-
Petchey, O. L. and Gaston, K. J. (2002). Functional diversity tação arbórea em um remanescente florestal urbano,
(FD), species richness and community composition. Eco- Uberlândia, MG. Natureza on line, 9(2): 67-75.
logy Letters, 5(3), 402 – 411. Vale, V. S., Schiavini, I., Araújo, G. M., Gusson, A. E., Lopes,
Pijl, V.D. (1982). Principles of dispersal in higher plants (2nd S. F., Oliveira, A. P., Prado-Junior, J. A., Arantes, C. S.
ed.). New York: Springer Verlag. and Dias-Neto, O. C. (2013a). Fast changes in seasonal
Pinheiro, E. S. and Durigan, G. (2009). Geotecnologias aplicadas forest communities due to soil moisture increase after
à análise da dinâmica do cerrado na Estação Ecológica de damming. International Journal of Tropical Biology,
Assis, SP. In J. C. N. Epiphanio and L. S. Galvão. Anais do 61(4), 1901-1917.
XIV Simpósio brasileiro de sensoriamento remoto (pp. Vale, V. S., Schiavini, I., Lopes, S. F., Oliveira, A. P., Dias-Neto,
2905-2912). Natal, Río Grande del Norte, Brasil. O. C. and Gusson, A. E. (2013b). Functional groups in a
Pinheiro, F. and Ribeiro, J. F. (2001). Síndromes de dispersão de semi deciduous seasonal forest in Southeastern Brazil.
sementes em Matas de Galeria do Distrito Federal. In J. F. Biotemas, 26(2), 45-58.

76
Madera y Bosques vol. 23, núm. 1: 63-77 Primavera 2017

Walker, B. H. (1992). Biodiversity and Ecological Redundancy. Wilkinson, L. (2002). Systat (version 10.2) [Software]. Chicago.
Conservation Biology, 6(1), 18-23.
Walker, B. H., Kinzig, A. and Langridge, J. (1999). Plant attri-
bute diversity, resilience, and ecosystem function: The
Manuscript received on July 23th 2014.
nature and significance of dominant and minor species. Accepted on December 14th 2016.
Ecosystems, 2, 95-113.
This paper must be cited as:
Walther, B. A., Clayton D. H. and Gregor, R. D. (1999). Showi- Vale, V. S., Oliveira, A. P., Prado-Junior, J. A., Ribeiro Londe, P. and
Nascimento, D. R. (2017). Damming water influences the structure,
ness of neotropical birds in relation to ectoparasite abun-
composition and functions of adjacent savannahs. Madera y
dance and foraging stratum. Oikos, 87, 157-165. Bosques 23(1), 63-77. doi:10.21829/myb.2017.2311527

77