Abstracts
Large areas in the Pantanal wetland are covered by monodominant formations, e.g. typical landscapes with local names such as “paratudal”, dominated by T. aurea. Studies on structure of these formations generally include only woody strata, consequently the species richness registered is usually low due to the absence of the ‘ground layer’ of herbaceous and others low species. We recorded 13 species, 12 genera and 11 families for the arboreal stratum. Considering arboreal flora without the dominant (T. aurea) individuals showed great establishment in relation to the flood level between 35 – 45 cm while the individuals of the dominant species of 30 – 45 cm. The diameter distribution revealed that the population of T. aurea did not show the reverse J curve, the usual pattern for species in constant regeneration, also evidenced in inconstant Licourt quotient, indicating an episodic recruitment that could lead to future changes in structure. In the herbaceous strata we recorded 78 species, included in 62 genera and 27 families. Using plots method we sampled 46 species, 40 genera and 22 families, while in line interception we found 65 species distributed in 57 genera and 26 families. The floristic similarity of Sørensen between both methods was 59.4%, with 33 species in common, and the method of line interception was more efficient in detecting richness, with 35% more species found in the same time. According to the methods of plots and line interception applied on the woody stratum, our results gave similar detailed information on the structure of this type of savanna, and in spite of being monodominant it shows high species richness when the herbaceous stratum is taken into account. Plots and line interception methods showed similar results for the woody stratum and high species richness of the herbaceous stratum.
line interception method; plot method; wetland; paratudal; pantanal
Amplas áreas no Pantanal são cobertas por formações monodominantes, tipicamente com nomes locais como “paratudal” dominado por Tabebuia aurea. Estudos na estrutura dessas formações geralmente incluem somente estrato arbóreo, consequentemente, a riqueza de espécies detectada geralmente é baixa devido à ausência do “estrato terrestre” das herbáceas e outras poucas espécies. Nós registramos 13 espécies, 12 gêneros e 11 famílias para o estrato arbóreo. Considerando a flora arbórea sem a espécie dominante (T. aurea) apresentaram um ótimo de estabelecimento em relação ao nível de inundação entre 35 – 45cm, enquanto os indíviduos da espécie dominante de 30 – 45cm. A distribuição diamétrica revelou que a população de T. aurea não apresentou o J reverso, o modelo usual para espécies em constante regeneração, também evidenciado pela não constante no quoeficiente de Licourt. Indicando episódio de recrutamento que poderia levar a futuras mudanças na estrutura. Para o estrato herbáceo geral registramos 78 espécies, incluindo 68 gêneros e 27 famílias. Usando o método de parcelas amostramos 46 espécies, 40 gêneros e 22 famílias, enquanto na interseção na linha nós encontramos 65 espécies distribuídas em 57 gêneros e 26 famílias. A similaridade florísitca de Sørensen entre ambos métodos foi de 59,4%, com 33 espécies em comum, e o método de interseção na linha foi mais eficiente na detecção da riqueza, 35% das espécies foram encontradas no mesmo tempo. De acordo com os métodos de parcelas e interseção na linha aplicado no estrato arbóreo, os nossos resultados deram uma semelhante informação detalhada na estrutura deste tipo de savana, e que apesar de ser monodominante mostrou alta riqueza de espécies quando o estrato herbáceo é levado em conta. Os métodos de parcelas e interseção na linha mostraram resultados similares para o estrato arbóreo e alta riqueza de espécies do estrato herbáceo.
método de interseção na linha; método de parcela; planície inundável; paratudal; pantanal
1. Introduction
A forest is considered monodominant when more than 50% of the
individuals or of its basal area belong to a single species (Connell and Lowman, 1989CONNELL, JH. and LOWMAN, MD., 1989. Low-diversity tropical rain
forests: some possible mechanisms for their existence. American Naturalist, vol.
134, no. 1, p. 88-119.
http://dx.doi.org/10.1086/284967.
http://dx.doi.org/10.1086/284967...
; Hart et al., 1989HART, TB., HART, JA. and MURPHY, PG., 1989. Monodominant and
species-rich forests of the humid tropics: causes for their co-ocurrence.
American Naturalist, vol. 133, no. 5, p. 613-633.
http://dx.doi.org/10.1086/284941.
http://dx.doi.org/10.1086/284941...
). In general, such forests are in sharp
contrast to other tropical forests where species diversity is much greater and they
represent one of the most intriguing enigmas of tropical ecology (Schluter and
Ricklefs, 1993). According to Richards (1952), extensive vegetation
formations dominated by a single or a few species are present in several tropical
regions and can cover hundreds of square kilometres, often occurring adjacent to
other forest types of higher diversity.
Such monodominant formations have been the subject of a few studies, in
which subjects such as floristic composition, structure, soils, population dynamics
were studied. Amongst these, two areas are particularly notable: forest dominated by
Brosimum rubescens Taub. in the state of Mato Grosso
(Marimon et al., 2001aMARIMON, BS., FELFILI, JM. and HARIDASAN, M., 2001a. Studies in
Monodominant Forests in Eastern Mato Grosso, Brazil: I. A Forest of Brosimum
rubescens Taub. Edinburgh Journal of Botany, vol. 58, p.
123-137., bMARIMON, BS., FELFILI, JM. and HARIDASAN, M., 2001b. Studies in
Monodominant Forests in Eastern Mato Grosso, Brazil: II. A Forest in the
Areões Xavante Indian Reserve. Edinburgh Journal of Botany, vol. 58, p.
483-497.) and forest dominated by
Peltogyne gracilipes Ducke on the Island of Maracá, in
Roraima (Milliken and Ratter, 1989, 1998; Nascimento and Proctor, 1994NASCIMENTO, MT. and PROCTOR, J., 1994. Insect Defoliation of a
Monodominant Amazoniam Rainforest. Journal of Tropical Ecology, vol. 10, no. 04,
p. 633-636.
http://dx.doi.org/10.1017/S0266467400008312.
http://dx.doi.org/10.1017/S0266467400008...
; 1997aNASCIMENTO, MT. and PROCTOR, J., 1997a. Soil and Plant Changes
Across a Monodominant Rain Forest Boundary on Maraca Island, Roraima, Brazil.
Global Ecology and Biogeography e Letters, vol. 6, no. 5, p. 387-395.
http://dx.doi.org/10.2307/2997339.
http://dx.doi.org/10.2307/2997339...
, b; Villela and Proctor,
1999VILLELA, DM. and PROCTOR, J., 1999. Litterfall Mass, Chemistry, and
Nutrient Retranslocation in a Monodominant Forest on Maraca Island, Roraima,
Brazil. Biotropica, vol. 31, no. 2, p. 198-211.
http://dx.doi.org/10.1111/j.1744-7429.1999.tb00132.x.
http://dx.doi.org/10.1111/j.1744-7429.19...
).
Wetland monodominant formations are particularly important in the
Brazilian Pantanal and cover large areas forming typical landscapes that receive
local names such as “canjiqueiral” (dominated by
Byrsonima cydioniifolia A. Juss.),
“cambarazal” (of Vochysia divergens Pohl),
“carandazal” (of Copernicia alba Morong ex
Morong and Britton), “paratudal” (of Tabebuia
aurea (Silva Manso) Benth. and Hook.), and others
(Pott and Pott, 1994). Most of these formations are seasonally flooded
savannas. The main studies of monodominant formations in the Pantanal are
(i) of V. divergens forest in the sub-region of
Poconé (Nascimento and Cunha, 1989NASCIMENTO, MT. and CUNHA, CN., 1989. Estrutura e
composição florística de um cambarazal no Pantanal de
Poconé-MT. Acta Botanica Brasilica, vol. 3, no. 1, p.
3-23.;
Arieira and Cunha, 2006ARIEIRA, J. and CUNHA, CN., 2006. Fitossociologia de uma floresta
inundável monodominante de Vochysia divergens Pohl
(Vochysiaceae), no Pantanal Norte, MT, Brasil. Acta Botanica
Brasilica, vol. 20, no. 3, p. 569-580.
http://dx.doi.org/10.1590/S0102-33062006000300007.
http://dx.doi.org/10.1590/S0102-33062006...
);
(ii) of C. alba palm savanna in the sub-regions of
Miranda and Nabileque, Mato Grosso do Sul (Amador et al., 2012AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and
SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba
and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia
Australis, vol. 16, no. 4, p. 846-858.
http://dx.doi.org/10.4257/oeco.2012.1604.09
http://dx.doi.org/10.4257/oeco.2012.1604...
); and (iii) of the arboreal stratum
of T. aurea in the sub-region of Miranda (Ribeiro and Brown, 2002RIBEIRO, SP. and BROWN, VK., 2002. Tree Species Monodominance or
Species-Rich Savannas? The Influence of Abiotic Factors in designing Plant
Communities of The Brazilian Cerrado and Pantanal Matogrossense A Review.
Ecotropica, vol. 8, p. 31-45.; Soares and Oliveira, 2009SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
).
However, studies of the structure of these monodominant formations usually
include only the tree and shrub strata, and use the plot method, e.g. that of
V. divergens by Nascimento and
Cunha (1989)NASCIMENTO, MT. and CUNHA, CN., 1989. Estrutura e
composição florística de um cambarazal no Pantanal de
Poconé-MT. Acta Botanica Brasilica, vol. 3, no. 1, p.
3-23., and Arieira and
Cunha (2006)ARIEIRA, J. and CUNHA, CN., 2006. Fitossociologia de uma floresta
inundável monodominante de Vochysia divergens Pohl
(Vochysiaceae), no Pantanal Norte, MT, Brasil. Acta Botanica
Brasilica, vol. 20, no. 3, p. 569-580.
http://dx.doi.org/10.1590/S0102-33062006000300007.
http://dx.doi.org/10.1590/S0102-33062006...
, and T. aurea savanna by Ribeiro and Brown (2002)RIBEIRO, SP. and BROWN, VK., 2002. Tree Species Monodominance or
Species-Rich Savannas? The Influence of Abiotic Factors in designing Plant
Communities of The Brazilian Cerrado and Pantanal Matogrossense A Review.
Ecotropica, vol. 8, p. 31-45. and Soares and Oliveira (2009)SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
. As a
consequence, the species richness found is usually very low. However, in a study on
C. alba savanna Amador et al.
(2012)AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and
SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba
and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia
Australis, vol. 16, no. 4, p. 846-858.
http://dx.doi.org/10.4257/oeco.2012.1604.09
http://dx.doi.org/10.4257/oeco.2012.1604...
demonstrated that it is species-rich when the herbaceous
stratum is included. As herbs constitute the major part of the floristic diversity
of the Pantanal (Pott and Pott, 1999; Schessl,
1999SCHESSL, M., 1999. Floristic composition and structure of floodplain
vegetation in the Northern Pantanal of Mato Grosso, Brazil. Phyton, vol. 39, p.
303-336.), it is possible that all the monodominant formations of the
Pantanal are in fact species-rich. Hence, for a more complete evaluation of this
vegetation, it is necessary to have more comprehensive methods to describe both
strata.
Two main methods are used to evaluate herbaceous formations in Brazil: the
line interception method (Canfield,
1941CANFIELD, R., 1941. Application of line interception in sampling
range vegetation. Journal of Forestry, vol. 39, p. 388-394., 1950; Brower and Zar,
1977BROWER, JE., and ZAR, JH., 1977. Field and laboratory methods for
general ecology. 2nd. ed. Dubuque: Wm. C. Brown Publishers. 226
p.), used by Munhoz and Felfili
(2006)MUNHOZ, CBR. and FELFILI, JM., 2006. Fitossociologia do estrato
herbáceo-subarbustivo de uma área de campo sujo no Distrito Federal,
Brasil. Acta Botanica Brasilica, vol. 20, no. 3, p. 671-685.
http://dx.doi.org/10.1590/S0102-33062006000300017.
http://dx.doi.org/10.1590/S0102-33062006...
, and the plot method with cover percentages (Brower and Zar, 1977BROWER, JE., and ZAR, JH., 1977. Field and laboratory methods for
general ecology. 2nd. ed. Dubuque: Wm. C. Brown Publishers. 226
p.) used, for example,
by Prado et al. (1994)PRADO, AL., HECKMAN, C. and MARTINS, FR., 1994. The seasonal
succession of biotic communities in wetlands of the tropical wet-and-dry
climatic zone: II. The aquatic macrophyte vegetation in the Pantanal of Mato
Grosso, Brazil. International Revue ges. Hydrobiology, vol. 79, p.
569-589. and
Rebelatto and Cunha (2005). The method of plots with percentage cover can
also be used for study of woody vegetation, supplying a good description where both
strata are important (Damasceno-Junior and Pott, 2011). However,
comparative evaluations of these methods have not yet been used for monodominant
formations of the Pantanal.
Thus, following the precept of study in monodominant formations, the objective of this study is to evaluate the phytosociological structure of the tree and herbaceous stratum monodominant formation of T. aurea (“paratudal”) in the Pantanal wetlands, directing the following questions. (i) Would flood be a factor influencing structure, richness and diversity of species in “paratudal” for both strata? (ii) Can the combination of the methods of the intersection lines and plots show variation in species richness?
2. Material and Methods
2.1. Study area
The Pantanal sub-region of Miranda is located within the following boundaries: to the North, the sub-region of Abobral; to the South, the Chaco woodlands of the municipality of Porto Murtinho; to the East, the sub-region of Aquidauana; and to the West, the Bodoquena highlands and the sub-region of Nabileque (Figure 1). It covers an area of 5,000 km2, of savanna (Cerrado), forest and grassland vegetation including a strong concentration of Copernicia alba palm savanna and especially Tabebuia aurea savanna (Brasil, 1982BRASIL., 1982. Ministério das Minas e Energia. Secretária Geral. Projeto RADAMBRASIL. Folha SD 20 Corumbá. Rio de Janeiro: Ministério das Minas e Energia. Levantamento dos Recursos Naturais, no. 26; Loureiro et al., 1982; Amaral, 1987).
Location of the Pantanal in Brazil (A), and the detail of the Pantanal wetland (B). The circule black indicates the the study area, ranch São Bento, close to the Pantanal research station of UFMS, on the right margin of the river Miranda, at Passo do Lontra, municipality of Corumbá, Mato Grosso do Sul state, Brazil.
Tabebuia aurea forms a park savanna vegetation
(Loureiro et al., 1982), and, in spite of occupying large areas, it is
almost restricted to the Miranda and Nabileque sub-regions of the Pantanal,
associated with slightly alkaline waters and sediments (Pott and Pott,
1994). It shows continuous areas into the Paraguayan and Bolivian wetlands;
the species also occurs sparsely in the sub-regions of the Nhecolândia and
Poconé (Pott and Pott, 1994). The dominant species
(T. aurea) usually occurs on very compact clay
soil, where there are so called mounds or “murundus” (Ribeiro and Brown, 2002RIBEIRO, SP. and BROWN, VK., 2002. Tree Species Monodominance or
Species-Rich Savannas? The Influence of Abiotic Factors in designing Plant
Communities of The Brazilian Cerrado and Pantanal Matogrossense A Review.
Ecotropica, vol. 8, p. 31-45.; Soares and Oliveira, 2009SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
). It has
seasonally flood plain areas with grasses and other herbs covered by water of
variable depth in the flood period.
The area has loamy-clay and sandy soils, with a strong predominance of the former (Silva et al., 2000SILVA, MP., MAURO, R., MOURÃO, G. and COUTINHO, M., 2000. Distribuição e quantificação de classes de vegetação do Pantanal através de levantamento aéreo. Revista Brasileira de Botânica, vol. 23, no. 2, p. 143-152.). The climate is type Aw of Köppen (1948), with annual rainfall around 1,100 mm. Our study was carried out at São Bento ranch (19°30′S, 57°04′W) close to the Pantanal Research Station of the Federal University of Mato Grosso do Sul (UFMS), on the right bank of the Miranda river, at Passo do Lontra, Corumbá, Mato Grosso do Sul.
2.2 Methods
We used two methods for the phytosociological analyses (as following by Damasceno-Junior and Pott (2011)POTT, A., OLIVEIRA, AKM., DAMASCENO-JUNIOR, GA. and SILVA, JSV., 2011. Plant diversity of the Pantanal wetland. Brazilian Journal of Biology, vol. 71, no. 1, sup. Suppl 1, p. 265-273. PMid:21537599.) (i) the plot method, where large plots were used for arboreal and smaller nested plots for herbaceous strata, and (ii) in addition the line interception for the latter.
The trees were sampled using ten plots of 0.15 ha (50×30 m), totalling 1.5 ha. These were randomly distributed in each of the T. aurea stands. The diameter of all trees with dbh ≥ 3.18 cm was measured as was that of their larger branches. Height was also estimated using a 2 m stick. The height of the water mark of the last flood on tree-trunks was also measured. In each plot 20 nested subplots (totalling 50 m2) were placed in random directions approximately 10m apart. In each plot the percentage cover per species was estimated visually; the same was done for bare ground and with that covered by dead material.
The second method was that of line interception (Canfield, 1941CANFIELD, R., 1941. Application of line interception in sampling range vegetation. Journal of Forestry, vol. 39, p. 388-394., 1950), for determination of the composition and linear cover of the ground species. Eight lines of 50 m were placed in random directions, inside and outside the plots of trees, with each line subdivided into sampling units of 1m, giving a total of 400 m. The method consists of recording the position and the cover area of each species, that is, the length which each inventoried species occupies per sample along the lines and is then divided by the total length of the species under the line, thus estimating the proportion of the area covered by that species. The sum of the horizontal projection of each species in all sampling units corresponds to the value of absolute lineal cover of the species in the area. The relative cover (RC) was determined by dividing the AC of each species by the sum of the AC of all multiplied by 100. The record of occurrence of each species in sampling units was used to calculate frequency. In both methods a concept of importance value (IV) for herbaceous vegetation was used; the concept was adapted where the relative frequency and RC were added to form the IV for which the maximum is 200 (Damasceno-Junior and Pott, 2011). We recorded bare ground and/or dead matter together with the vegetation analysis.
Individuals of T. aurea were distributed in diameter classes with optimal interval class (IC=6.0) calculated by the formulae of Spiegel (1976). Where, IC = A/NC; NC = 1+3,3 logN; A = range diameters; NC = number of classes e N = number of individuals.
The Licourt quotient “q”, dividing each diameter class
by the previous one (Meyer,
1952MEYER, HA., 1952. Structure, growth, and drain in balanced
uneven-aged forests. Journal of Forestry, vol. 50, no. 2, p.
85-92.), was calculated allowing inferences about recruitment and
mortality to be made. A constant relation between classes indicates that
recruitment and mortality rates are balanced (Nascimento et al., 2004NASCIMENTO, ART., FELFILI, JM. and MEIRELLES, EM., 2004.
Florística e estrutura da comunidade arbórea de um remanescente de
Floresta Estacional Decidual de encosta, Monte Alegre, GO, Brasil. Acta Botanica
Brasilica, vol. 18, no. 3, p. 659-669.
http://dx.doi.org/10.1590/S0102-33062004000300023.
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).
Species were identified in the field, using the nomenclature of APG III (2009), and when this was not possible, herbarium specimens were collected and later identified by specialists at the Universidade Federal de Mato Grosso do Sul (UFMS), Campo Grande campus.
The tree stratum was analysed using the program Mata Nativa 2
(Cientec, 2009CIENTEC, 2009. Mata Nativa 2: sistema para análise
fitossociológica e elaboração de planos de manejo de florestas
nativas. Viçosa: Cientec-Consultoria e Desenvolvimento de
Sistemas.), to
calculate the phytosociological parameters. In both methods, Shannon's
diversity (H′) and equability index (E) for the
herbaceous vegetation were calculated as recommended by Munhoz and Felfili (2006)MUNHOZ, CBR. and FELFILI, JM., 2006. Fitossociologia do estrato
herbáceo-subarbustivo de uma área de campo sujo no Distrito Federal,
Brasil. Acta Botanica Brasilica, vol. 20, no. 3, p. 671-685.
http://dx.doi.org/10.1590/S0102-33062006000300017.
http://dx.doi.org/10.1590/S0102-33062006...
using the cover per
species and not the number of individuals.
3. Results
3.1. Arboreal and herbaceous strata
A total of 88 species belonging to 72 genera and 30 families were recorded (Table 1). The most represented families were Poaceae with 18 species (20.2% of the total species), Fabaceae with 12 (13.5%), Malvaceae with 8 (9.0%), Rubiaceae with 5 (5.6%), Cyperaceae with 4 (4.5%), and Alismataceae, Asteraceae, Euphorbiaceae, Onagraceae and Verbenaceae each with 3 species (3.4%) (Table 1).
List of species recorded in a tropical floodable Tabebuia aurea monodominant savanna, sub-region of Miranda, Pantanal, MS, Brazil. And sampling method: A - tree plot method; B - herbaceous plot method; C - line interception method for herbaceous species.
In the 1.5 ha sampled for tree stratum of T. aurea savanna, we recorded 448 individuals of 13 species from 12 genera and 11 families. The most represented families were Bignoniaceae and Fabaceae with two species each and the other families occurred only once. T. aurea had the highest IV, followed by B. cydoniifolia (Table 2). The diversity index (H′) was 0.84, with an evenness (E) of 0.33.
Phytosociological parameters of species observed in a tropical floodable Tabebuia aurea monodominant savanna, sub-region of Miranda, Pantanal, MS, Brazil. N = number of individuals; BA = basal area; AD = absolute density; RD = relative density; AF = absolute frequency, RF = relative frequency; ADo = absolute dominance; RDo = relative dominance; CV = coverage value, CV% = coverage value percentage, IV = importance value, IV% = importance value percentage.
Judging by the watermarks on the trunks, 30-45 cm of flooding was the commonest level for T. aurea, the dominant species, while other species showed 35-45 cm (Figure 2). The diameter distribution did not show the reverse “J” pattern as expected for a population in constant regeneration as in the case of the dominant species, T. aurea (Figure 3).
(a) Representation of the relationship of population density of tree species without Tabebuia aurea with the height of the water mark (cm) (p = 0.002), and (b) relationship of population density of Tabebuia aurea with the height of the water mark (cm) (p < 0.001), in a tropical floodable Tabebuia aurea monodominant savanna, sub-region of Miranda, Pantanal, Mato Grosso do Sul state, Brazil.
Frequency distribution of diameters (DBH) of individuals of Tabebuia aurea (Bignoniaceae) and Licourt quotient “q” sampled in a tropical floodable Tabebuia aurea monodominant savanna, sub-region of Miranda, Pantanal, Mato Grosso do Sul state, Brazil.
The families with the largest number of species in the herbaceous stratum were Poaceae (18), Fabaceae (10), Malvaceae (8), Cyperaceae and Rubiaceae with 4 species each, totalling 53.7% of the recorded species. In the plots we sampled 46 species, 40 genera and 22 families. Line interception gave 65 species distributed in 57 genera and 26 families. Index of species Shannon diversity (H′) and equability (E) were respectively 2.67 nats/individuals and 0.68 for the method of plots, and 2.25 nats/individuals and 0.53 for the method of the line interception.
The phytosociological survey of the herbaceous stratum showed 78 species, included in 62 genera and 27 families (Table 3). Sebastiania hispida and Andropogon hypogynus showed the highest importance values (IV) in both methods employed (Figure 4). The class bare area (bare ground and/or dead matter) had the largest IV in both methods (29.0% for plots and 48.4% for line interception), the other categories were species showing differences in cover and IV (Table 3).
Importance Value (IV) of the ten species most representative by method of plots (a) and line interseption method (b) in a tropical floodable Tabebuia aurea monodominant savanna, sub-region of Miranda, Pantanal, Mato Grosso do Sul state, Brazil. IV (Importance Value), RC (Relative Cover) and RF (Relative Frequency).
Phytosociological parameters of herbaceous species found in a tropical floodable Tabebuia aurea monodominant savanna. sub-region of Miranda. Pantanal. MS. Brazil. Line interception and Plots methods. RC = Relative Cover; RF = relative frequency; VIh = importance value for herbaceous plants.
3.2. Comparison of methods
The Sørensen floristic similarity comparing the results of both methods was 59.4%, with 33 species in common (Table 3). Despite this, both methods showed lower values of Shannon diversity index (H′) (2.25 line and 2.67 plot) for species and equability (E) (0.68 line and 0.53 plot), the diversity detected by line interception was more efficient, with 35% more species recorded than for the plots (Table 3).
4. Discussion
4.1. Arboreal stratum
Our study recorded more species in the arboreal stratum
(13) than in the surveys by Soares and
Oliveira (2009)SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
in three T. aurea areas in
the Pantanal sub-region of Miranda, where six recorded tree species occurred,
all of them also found in our study (T. aurea, B. cydoniifolia,
Erythroxylon anguifugum, Inga vera, Handroanthus heptaphyllus, Zanthoxylum
rigidum) (Table
2). Such low diversity of woody species is reported for other
monodominant formations (Nascimento and
Cunha 1989NASCIMENTO, MT. and CUNHA, CN., 1989. Estrutura e
composição florística de um cambarazal no Pantanal de
Poconé-MT. Acta Botanica Brasilica, vol. 3, no. 1, p.
3-23.; Arieira and Cunha
2006ARIEIRA, J. and CUNHA, CN., 2006. Fitossociologia de uma floresta
inundável monodominante de Vochysia divergens Pohl
(Vochysiaceae), no Pantanal Norte, MT, Brasil. Acta Botanica
Brasilica, vol. 20, no. 3, p. 569-580.
http://dx.doi.org/10.1590/S0102-33062006000300007.
http://dx.doi.org/10.1590/S0102-33062006...
; Amador et al.,
2012AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and
SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba
and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia
Australis, vol. 16, no. 4, p. 846-858.
http://dx.doi.org/10.4257/oeco.2012.1604.09
http://dx.doi.org/10.4257/oeco.2012.1604...
), demonstrating that the higher the water level and the longer
its duration, the smaller the number of tolerant species. Damasceno-Junior et al. (2005)DAMASCENO-JUNIOR, GA., SEMIR, J., ANTONIO MAËS DOS SANTOS, F.
and DE FREITAS LEITÃO-FILHO, H., 2005. Structure, distribution of species
and inundation in a riparian forest of Rio Paraguai, Pantanal, Brazil. Flora,
vol. 200, no. 2, p. 119-135.
http://dx.doi.org/10.1016/j.flora.2004.09.002.
http://dx.doi.org/10.1016/j.flora.2004.0...
, Amorim and Batalha (2006)AMORIM, PK. and BATALHA, MA., 2006. Soil characteristics of a
hyperseasonal cerrado compared to a seasonal cerrado and a floodplain grassland:
implications for plant community structure. Brazilian Journal of Biology, vol.
66, no. 2B, p. 661-670.
http://dx.doi.org/10.1590/S1519-69842006000400010.
PMid:16906298
http://dx.doi.org/10.1590/S1519-69842006...
, Carvalho et al. (2006)CARVALHO, FA., NASCIMENTO, MT., BRAGA, JMA. and RODRIGUES, PJFP.,
2006. Estrutura da comunidade arbórea da floresta atlântica de baixada
periodicamente inundada na Rebio de Poço das Antas, RJ. Rodriguesia, vol.
5, p. 503-518., Silva and Batalha (2008)SILVA, IA. and BATALHA, MA., 2008. Species convergence into
life-forms in a hyperseasonal cerrado in central Brazil. Brazilian Journal of
Biology, vol. 68, no. 2, p. 329-339.
http://dx.doi.org/10.1590/S1519-69842008000200014.
PMid:18660961
http://dx.doi.org/10.1590/S1519-69842008...
and
Cianciaruso and Batalha
(2009)CIANCIARUSO, MV. and BATALHA, MA., 2009. Short-term community
dynamics in seasonal and hyperseasonal cerrados. Brazilian Journal of Biology,
vol. 69, no. 2, p. 231-240.
http://dx.doi.org/10.1590/S1519-69842009000200002.
PMid:19675923
http://dx.doi.org/10.1590/S1519-69842009...
discuss water saturation of the soil as a selective
force for tree species, limiting diversity by acting as an environmental filter
against intolerant species. However, tree species conditioned to flooding and
showing tolerance to such stress were recorded (Pott and Pott 1994; Damasceno-Junior et al., 2005DAMASCENO-JUNIOR, GA., SEMIR, J., ANTONIO MAËS DOS SANTOS, F.
and DE FREITAS LEITÃO-FILHO, H., 2005. Structure, distribution of species
and inundation in a riparian forest of Rio Paraguai, Pantanal, Brazil. Flora,
vol. 200, no. 2, p. 119-135.
http://dx.doi.org/10.1016/j.flora.2004.09.002.
http://dx.doi.org/10.1016/j.flora.2004.0...
; Pott et al., 2011POTT, A., OLIVEIRA, AKM., DAMASCENO-JUNIOR, GA. and SILVA, JSV.,
2011. Plant diversity of the Pantanal wetland. Brazilian Journal of Biology,
vol. 71, no. 1, sup. Suppl 1, p. 265-273. PMid:21537599.).
Of the case species T. aurea dominates compared to
the other species recorded in the Paratudal, showing an advantage because of the
flood pulse. T. aurea is the species most tolerant of such
stress and thus dominates in wetter conditions (Pott and Pott, 1994; Damasceno-Junior et al., 2005DAMASCENO-JUNIOR, GA., SEMIR, J., ANTONIO MAËS DOS SANTOS, F.
and DE FREITAS LEITÃO-FILHO, H., 2005. Structure, distribution of species
and inundation in a riparian forest of Rio Paraguai, Pantanal, Brazil. Flora,
vol. 200, no. 2, p. 119-135.
http://dx.doi.org/10.1016/j.flora.2004.09.002.
http://dx.doi.org/10.1016/j.flora.2004.0...
; Pott and
Pott, 1999). As described by Soares and
Oliveira (2009)SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
, the growth of T. aurea on
the mounds (murundus) is probably a factor that allows seedling
survival in times of flood, when these mounds protect the young individuals from
submersion.
The advantage afforded is reflected in the importance value as well as dominance of T. aurea, a factor referred to by several authors, who mention it as a species associated with mounds, i.e. small elevations on the plain (miniature islands), where T. aurea avoid floods. However, in our area T. aurea occur not only on mounds, but also more rarely on the lower ground in between them, although there is an indication that T. aurea cannot survive below the floodline (Pott and Pott, 1994; Ribeiro and Brown, 2002RIBEIRO, SP. and BROWN, VK., 2002. Tree Species Monodominance or Species-Rich Savannas? The Influence of Abiotic Factors in designing Plant Communities of The Brazilian Cerrado and Pantanal Matogrossense A Review. Ecotropica, vol. 8, p. 31-45.). The mound/T. aurea association tends to be more frequent where duration and level of flood are higher, such as in the case of the Paratudal under influence of the Paraguay river flood, providing the trees higher probability of survival to grow almost exclusively on the mounds. Silva Júnior and Felfili (1996) add that these islands or mounds have drainage conditions and determine good soil aeration favouring the propagation of tree species.
In the Licourt quotient, where “q” is not constant, as in the case of the values of our study, there is a discrepancy between rates of mortality and recruitment that characterise changes in forest structure (Felfili et al., 1998FELFILI, JM., SILVA JUNIOR, MC. and NOGUEIRA, PE., 1998. Levantamento da vegetação arbórea na região de Nova Xavantina, MT. Boletim do Herbário Ezechias Paulo Heringer, vol. 3, p. 63-81.). However, it is probable that, according to Felfili (1997)FELFILI, JM., 1997. Diameter and height distributions in a gallery forest community and some of its main species in central Brazil over a six-year period (1985-1991). Revista Brasileira de Botânica, vol. 20, no. 2, p. 155-162., species require a broad spatial and temporal scale to achieve balance between these events. Although establishment of individuals under varying conditions may cause differences in the distribution pattern of tree diameters. Such differences could be in variation in flood conditions on anthropic activity such as burning and grazing.
4.2. Herbaceous stratum
The floristic richness of Poaceae and their dominance cover in of many habitats in the Pantanal are well known, and they are very important as natural forage. The Poaceae, Malvaceae, Rubiaceae and Cyperaceae are also abundant in the Pantanal, the last particularly in more humid areas. More severe flooding limits the numbers of herbaceous species but under such conditions semi-aquatic and aquatic perennials appear such as Echinodorus grandiflorus, E. paniculatus, E. teretoscapus, Ludwigia lagunae, L. octovalvis, L. tomentosa, Pontederia cordata, P. subovata (Pott and Pott, 2000). Whereas Axonopus leptostachyus, Paspalum laxum, Reimarochloa acuta, Setaria parvilfora are indicators of dry periods (Pott and Pott, 1994; Schessl, 1999SCHESSL, M., 1999. Floristic composition and structure of floodplain vegetation in the Northern Pantanal of Mato Grosso, Brazil. Phyton, vol. 39, p. 303-336.).
In the area analysed, as in other areas subject to flooding, the
effect is to limit the number of both woody and herbaceous species in the
environment (Rebellato and Cunha,
2005REBELLATO, L. and CUNHA, CN., 2005. Efeito do “fluxo sazonal
mínimo da inundação” sobre a composição e
estrutura de um campo inundável no Pantanal de Poconé, MT, Brasil.
Acta Botanica Brasilica, vol. 19, no. 4, p. 789-799.
http://dx.doi.org/10.1590/S0102-33062005000400015.
http://dx.doi.org/10.1590/S0102-33062005...
); clearly this varies with the duration severity and
frequency of flooding (Scremin-Dias et al.
2011SCREMIN-DIAS, E., LORENZ-LEMKE, AP. and OLIVEIRA, AKM., 2011. The
floristic heterogeneity of the Pantanal and the occurrence of species with
different adaptive strategies to water stress. Brazilian Journal of Biology,
vol. 71, no. 1, sup. Suppl 1, p. 275-282.
http://dx.doi.org/10.1590/S1519-69842011000200006.
PMid:21537600
http://dx.doi.org/10.1590/S1519-69842011...
).
This conditioning for herbaceous diversity values were below those
recorded for the Pantanal flood areas of Poconé for both periods
(flood 4.01-3.29 drought) (Rebellato and Cunha, 2005REBELLATO, L. and CUNHA, CN., 2005. Efeito do “fluxo sazonal
mínimo da inundação” sobre a composição e
estrutura de um campo inundável no Pantanal de Poconé, MT, Brasil.
Acta Botanica Brasilica, vol. 19, no. 4, p. 789-799.
http://dx.doi.org/10.1590/S0102-33062005000400015.
http://dx.doi.org/10.1590/S0102-33062005...
). And there was also a lower value
from Carandazal of the sub-region of Miranda (1.15) and a higher one
for Carandazal of the sub-region of Nabileque (0.33) (Amador et al., 2012AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and
SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba
and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia
Australis, vol. 16, no. 4, p. 846-858.
http://dx.doi.org/10.4257/oeco.2012.1604.09
http://dx.doi.org/10.4257/oeco.2012.1604...
). These
differences in the compared diversity values, are associated to environmental
conditions, such as the lower value recorded for Carandazal from Nabileque. The
reason is that areas under inundation of the Paraguay River are subject to more
strict conditions (Amador et al.,
2012AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and
SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba
and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia
Australis, vol. 16, no. 4, p. 846-858.
http://dx.doi.org/10.4257/oeco.2012.1604.09
http://dx.doi.org/10.4257/oeco.2012.1604...
). Schessl
(1999)SCHESSL, M., 1999. Floristic composition and structure of floodplain
vegetation in the Northern Pantanal of Mato Grosso, Brazil. Phyton, vol. 39, p.
303-336. observed that in the more flooded Pantanal
environment of Poconé, there is a strong variation in the coverage and
composition of species promoted by hydric seasonality. Therefore, depending on
the intensity and duration of flooding, different responses of ground layer
characterise communities in relation to richness and diversity, as observed in
the Paratudal.
Within the herbaceous species that best featured the structure
phytosociology for both methodologies were Sebastiania hispida
which is characterised as weedy and an indicator of disturbance and fire
(Pott and Pott, 1994), and Andropogon hypogynus cited
as a dominant grass species in the Miranda sub-region (Pott et al., 2011POTT, A., OLIVEIRA, AKM., DAMASCENO-JUNIOR, GA. and SILVA, JSV.,
2011. Plant diversity of the Pantanal wetland. Brazilian Journal of Biology,
vol. 71, no. 1, sup. Suppl 1, p. 265-273. PMid:21537599.). According to Soares and Oliveira (2009)SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de
Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347.
http://dx.doi.org/10.1590/S0100-67622009000200015.
http://dx.doi.org/10.1590/S0100-67622009...
A. hypogynus was second in density and frequency in two of
three T. aurea savanna areas analysed. The success of
A. hypogynus may be attributed to its occurrence in
different microhabitats, while its maintenance throughout the year is due to the
high physiological and morphological plasticity which makes it tolerant to
either flooding or drought (Junk et al.,
2006JUNK, WJ., CUNHA, CN., WANTZEN, KM., PETERMANN, P., STRÜSSMANN,
C., MARQUES, MI. and ADIS, J., 2006. Biodiversity and its conservation in the
Pantanal of Mato Grosso, Brazil. Aquatic Sciences, vol. 68, no. 3, p. 278-309.
http://dx.doi.org/10.1007/s00027-006-0851-4.
http://dx.doi.org/10.1007/s00027-006-085...
; Rebellato et al.,
2012REBELLATO, L., NUNES DA CUNHA, C. and FIGUEIRA, JEC., 2012.
Respostas da Comunidade Herbácea ao Pulso de Inundação no
Pantanal de Poconé, Mato Grosso. Oecologia Australis, vol. 16, no. 4, p.
797-818.
http://dx.doi.org/10.4257/oeco.2012.1604.06
http://dx.doi.org/10.4257/oeco.2012.1604...
).
The high importance values for bare area is caused by a dry period
and much of the dead matter comes from species established there, and were
undergoing a succession stage after flooding. Still, according to Junk et al. (1989)JUNK, WJ., BAYLEY, PB. and SPARKS, RS., 1989. The flood pulse
concept in river-floodplain systems. Canadian Journal of Fishers and Aquatic,
vol. 106, p. 110-127., the
maintenance of an initial state of succession in the floodplain, as a result of
constant renovation through the flood pulse, promotes the high productivity. The
presence of bare ground and/or dead matter promotes soil moisture, and thus
allows survival for several weeks of some aquatic species, as previously
described. These aquatic species considered, together with annual species, form
the temporary species and their occurrence and importance depends on time of
observation and collection. Such a process is also related to the seed bank,
indicating an extremely important role in the resilience of these transitional
communities (Brock et al., 2003BROCK, MA., NIELSEN, DL., SHIEL, RJ., GREEN, JD. and LANGLEY, JD.,
2003. Drought and aquatic community resilience: the role of eggs and seeds in
sediments of temporary wetlands. Freshwater Biology, vol. 48, no. 7, p.
1207-1218.
http://dx.doi.org/10.1046/j.1365-2427.2003.01083.x.
http://dx.doi.org/10.1046/j.1365-2427.20...
;
Rebellato and Cunha 2005REBELLATO, L. and CUNHA, CN., 2005. Efeito do “fluxo sazonal
mínimo da inundação” sobre a composição e
estrutura de um campo inundável no Pantanal de Poconé, MT, Brasil.
Acta Botanica Brasilica, vol. 19, no. 4, p. 789-799.
http://dx.doi.org/10.1590/S0102-33062005000400015.
http://dx.doi.org/10.1590/S0102-33062005...
; Junk et al., 2006JUNK, WJ., CUNHA, CN., WANTZEN, KM., PETERMANN, P., STRÜSSMANN,
C., MARQUES, MI. and ADIS, J., 2006. Biodiversity and its conservation in the
Pantanal of Mato Grosso, Brazil. Aquatic Sciences, vol. 68, no. 3, p. 278-309.
http://dx.doi.org/10.1007/s00027-006-0851-4.
http://dx.doi.org/10.1007/s00027-006-085...
; Alves Pagotto et al., 2011ALVES PAGOTTO, M., DE MORAES LIMA SILVEIRA, R., NUNES DA CUNHA, C.
and FANTIN-CRUZ, I., 2011. Distribution of Herbaceous Species in the soil seed
bank of a flood seasonality area, northern Pantanal, Brazil. International
Review of Hydrobiology, vol. 96, no. 2, p. 149-163.
http://dx.doi.org/10.1002/iroh.201111315.
http://dx.doi.org/10.1002/iroh.201111315...
).
Although there are no data regarding the intensity of human intervention in the area, especially with regard to the effects of grazing and burning, the presence of species considered ruderal such as Melochia arenosa, M. parvifolia, M. simplex, Sida cerradoensis, could contribute to elevate the species richness.
4.3. Comparison of methods
We recorded in both methods dominance of the same species of the
herbaceous stratum, Sebastiania hispida and Andropogon
hypogynus, which influence the richness and diversity values
recorded. There was 59.8% of Sorensen similarity between methods,
indicating strong floristic relationship. According to Felfili et al.
(2001), Sørensen indices over 50% indicate elevated
similarity between two areas. Munhoz and Felfili
(2007)MUNHOZ, CB. and FELFILI, JM., 2007. Florística do estrato
herbáceo-subarbustivo de um campo limpo úmido em Brasília,
Brasil. Biota Neotropica, vol. 7, no. 3, p. 205-215.
http://dx.doi.org/10.1590/S1676-06032007000300022.
http://dx.doi.org/10.1590/S1676-06032007...
reported similar similarity values when analysing
the floodplain grassland by the method of linear interception, correlating their
result with edaphic homogeneity.
The line interception method was sensitive in the detection of the
diversity. That is probably because it allows the equivalent of larger areas to
be evaluated linearly. This method is equivalent to using a long narrow plot. As
already discussed in the literature (see Condit et al., 1996CONDIT, R., HUBBELL, S., LAFRANKIE, JV., SUKUMAR, R., MANOKARAN, N.,
FOSTER, RB. and ASHTON, PS., 1996. Species-area and species-individual
relationships for tropical trees: a comparison of three 50-ha plots. Journal of
Ecology, vol. 84, no. 4, p. 549-562.
http://dx.doi.org/10.2307/2261477.
http://dx.doi.org/10.2307/2261477...
), there is a higher probability of sampling
more species along the line, because it is possible to sample a higher number of
microhabitats. On the other hand, plots are better for trees, giving a more
precise idea of community structure as a whole. It is, however, possible to
survey trees by the line intersection method; however, this was not used in our
study because of the low arboreal density.
5. Conclusion
In spite of being a monodominant formation of Tabebuia aurea, the vegetation has significant richness with a total of 88 species recorded, when herbaceous stratum is taken into account. Of the employed methods, plots have the advantage of integrating the data on tree and on ground vegetation, thus allowing a more detailed analysis, however, the method of line interception is better in detecting species richness.
Flooding is a selective factor in occurrence species in both tree and ground strata, and thus influences richness and diversity. However, there is a need for continuous monitoring of the area so that one can understand successional changes of this vegetation type relative the to flood pulse, as well as anthropic processes such as grazing and fire.
Acknowledgements – We are grateful to the Postgraduate Programme in Plant Biology and to all students of the course of “Phytosociological Sampling Methods Adapted to the Vegetation Formations of the Pantanal”. We are grateful to the Brazilian CNPq and CAPES organizations for research and Senior Professor grants to Dr. A. Pott. And we would also like to thank the reviewers.
References
- ALVES PAGOTTO, M., DE MORAES LIMA SILVEIRA, R., NUNES DA CUNHA, C. and FANTIN-CRUZ, I., 2011. Distribution of Herbaceous Species in the soil seed bank of a flood seasonality area, northern Pantanal, Brazil. International Review of Hydrobiology, vol. 96, no. 2, p. 149-163. http://dx.doi.org/10.1002/iroh.201111315.
» http://dx.doi.org/10.1002/iroh.201111315 - AMADOR, GA., DAMASCENO-JÚNIOR, GA., CASAGRANDE, JC. and SARTORI, ALB., 2012. Structure of two communities dominated by Copernicia Alba and associations with soil and inundation in Pantanal Wetland, Brazil. Oecologia Australis, vol. 16, no. 4, p. 846-858. http://dx.doi.org/10.4257/oeco.2012.1604.09
» http://dx.doi.org/10.4257/oeco.2012.1604.09 - AMARAL, JAM.,1987. Solos do Pantanal. In Recursos Forrageiros Nativos do Pantanal Mato-Grossense. ALLEM A. C. and VALLS J. F. M. Brasília: Embrapa-Cenargen. p. 19-22. Documentos, no. 8
- AMORIM, PK. and BATALHA, MA., 2006. Soil characteristics of a hyperseasonal cerrado compared to a seasonal cerrado and a floodplain grassland: implications for plant community structure. Brazilian Journal of Biology, vol. 66, no. 2B, p. 661-670. http://dx.doi.org/10.1590/S1519-69842006000400010. PMid:16906298
» http://dx.doi.org/10.1590/S1519-69842006000400010 - ANGIOSPERM PHYLOGENY GROUP – APG III, 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society, vol. 161, no. 2, p. 105-121. http://dx.doi.org/10.1111/j.1095-8339.2009.00996.x.
» http://dx.doi.org/10.1111/j.1095-8339.2009.00996.x - ARIEIRA, J. and CUNHA, CN., 2006. Fitossociologia de uma floresta inundável monodominante de Vochysia divergens Pohl (Vochysiaceae), no Pantanal Norte, MT, Brasil. Acta Botanica Brasilica, vol. 20, no. 3, p. 569-580. http://dx.doi.org/10.1590/S0102-33062006000300007.
» http://dx.doi.org/10.1590/S0102-33062006000300007 - BRASIL., 1982. Ministério das Minas e Energia. Secretária Geral. Projeto RADAMBRASIL. Folha SD 20 Corumbá. Rio de Janeiro: Ministério das Minas e Energia. Levantamento dos Recursos Naturais, no. 26
- BROCK, MA., NIELSEN, DL., SHIEL, RJ., GREEN, JD. and LANGLEY, JD., 2003. Drought and aquatic community resilience: the role of eggs and seeds in sediments of temporary wetlands. Freshwater Biology, vol. 48, no. 7, p. 1207-1218. http://dx.doi.org/10.1046/j.1365-2427.2003.01083.x.
» http://dx.doi.org/10.1046/j.1365-2427.2003.01083.x - BROWER, JE., and ZAR, JH., 1977. Field and laboratory methods for general ecology. 2nd. ed. Dubuque: Wm. C. Brown Publishers. 226 p.
- CANFIELD, R., 1941. Application of line interception in sampling range vegetation. Journal of Forestry, vol. 39, p. 388-394.
- CANFIELD, R.,1950. Sampling range by the line interception method. Southwestern Forest and Range Experiment Station.
- CARVALHO, FA., NASCIMENTO, MT., BRAGA, JMA. and RODRIGUES, PJFP., 2006. Estrutura da comunidade arbórea da floresta atlântica de baixada periodicamente inundada na Rebio de Poço das Antas, RJ. Rodriguesia, vol. 5, p. 503-518.
- CIANCIARUSO, MV. and BATALHA, MA., 2009. Short-term community dynamics in seasonal and hyperseasonal cerrados. Brazilian Journal of Biology, vol. 69, no. 2, p. 231-240. http://dx.doi.org/10.1590/S1519-69842009000200002. PMid:19675923
» http://dx.doi.org/10.1590/S1519-69842009000200002 - CIENTEC, 2009. Mata Nativa 2: sistema para análise fitossociológica e elaboração de planos de manejo de florestas nativas. Viçosa: Cientec-Consultoria e Desenvolvimento de Sistemas.
- CONDIT, R., HUBBELL, S., LAFRANKIE, JV., SUKUMAR, R., MANOKARAN, N., FOSTER, RB. and ASHTON, PS., 1996. Species-area and species-individual relationships for tropical trees: a comparison of three 50-ha plots. Journal of Ecology, vol. 84, no. 4, p. 549-562. http://dx.doi.org/10.2307/2261477.
» http://dx.doi.org/10.2307/2261477 - CONNELL, JH. and LOWMAN, MD., 1989. Low-diversity tropical rain forests: some possible mechanisms for their existence. American Naturalist, vol. 134, no. 1, p. 88-119. http://dx.doi.org/10.1086/284967.
» http://dx.doi.org/10.1086/284967 - DAMASCENO-JUNIOR, GA. and POTT, A.,2011. Métodos de Amostragem em estudos fitossociológicos sugeridos para o Pantanal. In FELFILI, J.M., EISENLOHR, P.V., MELO, M.M.R.F., ANDRADE, L.A. and MEIRA NETO, J.A.A. (Eds.) Fitossociologia no Brasil: Métodos e Estudos de Casos. Viçosa: Editora UFV.
- DAMASCENO-JUNIOR, GA., SEMIR, J., ANTONIO MAËS DOS SANTOS, F. and DE FREITAS LEITÃO-FILHO, H., 2005. Structure, distribution of species and inundation in a riparian forest of Rio Paraguai, Pantanal, Brazil. Flora, vol. 200, no. 2, p. 119-135. http://dx.doi.org/10.1016/j.flora.2004.09.002.
» http://dx.doi.org/10.1016/j.flora.2004.09.002 - FELFILI, JM., 1997. Diameter and height distributions in a gallery forest community and some of its main species in central Brazil over a six-year period (1985-1991). Revista Brasileira de Botânica, vol. 20, no. 2, p. 155-162.
- FELFILI, JM., SILVA JUNIOR, MC. and NOGUEIRA, PE., 1998. Levantamento da vegetação arbórea na região de Nova Xavantina, MT. Boletim do Herbário Ezechias Paulo Heringer, vol. 3, p. 63-81.
- FELFILI, JM., SILVA JUNIOR, MCF., REZENDE, AV., HARIDASAN, M., FILGUEIRAS, TS., MENDONÇA, RC., WALTER, BMT. and NOGUEIRA, PE.,2001. O projeto Biogeografia do Bioma Cerrado: hipóteses e padronização da metodologia. In GARAY, I. and DIAS, B. (Orgs.). Conservação da biodiversidade em ecossistemas tropicais. Petrópolis: Vozes, p. 157-163.
- HART, TB., HART, JA. and MURPHY, PG., 1989. Monodominant and species-rich forests of the humid tropics: causes for their co-ocurrence. American Naturalist, vol. 133, no. 5, p. 613-633. http://dx.doi.org/10.1086/284941.
» http://dx.doi.org/10.1086/284941 - JUNK, WJ., BAYLEY, PB. and SPARKS, RS., 1989. The flood pulse concept in river-floodplain systems. Canadian Journal of Fishers and Aquatic, vol. 106, p. 110-127.
- JUNK, WJ., CUNHA, CN., WANTZEN, KM., PETERMANN, P., STRÜSSMANN, C., MARQUES, MI. and ADIS, J., 2006. Biodiversity and its conservation in the Pantanal of Mato Grosso, Brazil. Aquatic Sciences, vol. 68, no. 3, p. 278-309. http://dx.doi.org/10.1007/s00027-006-0851-4.
» http://dx.doi.org/10.1007/s00027-006-0851-4 - KÖPPEN, WP.,1948. Climatologia: con un estudio de los climas de la tierra. México: Fondo de Cultura Económica.
- LOUREIRO, RL., SOUZA LIMA, JP. and FONZAR, BC.,1982. Vegetação: As regiões fitoecológicas, sua natureza e seus recursos econômicos. In: Brasil. Ministério das Minas e Energia. Projeto Radam Brasil. Folha SE 21. Corumbá e parte da Folha SE 20. Rio de Janeiro: Ministério das Minas e Energia. Levantamento de Recursos Naturais, no. 27.
- MARIMON, BS., FELFILI, JM. and HARIDASAN, M., 2001a. Studies in Monodominant Forests in Eastern Mato Grosso, Brazil: I. A Forest of Brosimum rubescens Taub. Edinburgh Journal of Botany, vol. 58, p. 123-137.
- MARIMON, BS., FELFILI, JM. and HARIDASAN, M., 2001b. Studies in Monodominant Forests in Eastern Mato Grosso, Brazil: II. A Forest in the Areões Xavante Indian Reserve. Edinburgh Journal of Botany, vol. 58, p. 483-497.
- MEYER, HA., 1952. Structure, growth, and drain in balanced uneven-aged forests. Journal of Forestry, vol. 50, no. 2, p. 85-92.
- MILLIKEN, W. and RATTER, JA., (Eds.). 1998. Maracá - The Biodiversity and Environment of an Amazonian Rainforest. Chichester: John Wiley and Sons.
- MILLIKEN, W. and RATTER, JA.,1989. The vegetation of the Ilha de Maracá. Edinburgh: Royal Botanic Garden.
- MUNHOZ, CB. and FELFILI, JM., 2007. Florística do estrato herbáceo-subarbustivo de um campo limpo úmido em Brasília, Brasil. Biota Neotropica, vol. 7, no. 3, p. 205-215. http://dx.doi.org/10.1590/S1676-06032007000300022.
» http://dx.doi.org/10.1590/S1676-06032007000300022 - MUNHOZ, CBR. and FELFILI, JM., 2006. Fitossociologia do estrato herbáceo-subarbustivo de uma área de campo sujo no Distrito Federal, Brasil. Acta Botanica Brasilica, vol. 20, no. 3, p. 671-685. http://dx.doi.org/10.1590/S0102-33062006000300017.
» http://dx.doi.org/10.1590/S0102-33062006000300017 - NASCIMENTO, ART., FELFILI, JM. and MEIRELLES, EM., 2004. Florística e estrutura da comunidade arbórea de um remanescente de Floresta Estacional Decidual de encosta, Monte Alegre, GO, Brasil. Acta Botanica Brasilica, vol. 18, no. 3, p. 659-669. http://dx.doi.org/10.1590/S0102-33062004000300023.
» http://dx.doi.org/10.1590/S0102-33062004000300023 - NASCIMENTO, MT. and CUNHA, CN., 1989. Estrutura e composição florística de um cambarazal no Pantanal de Poconé-MT. Acta Botanica Brasilica, vol. 3, no. 1, p. 3-23.
- NASCIMENTO, MT. and PROCTOR, J., 1994. Insect Defoliation of a Monodominant Amazoniam Rainforest. Journal of Tropical Ecology, vol. 10, no. 04, p. 633-636. http://dx.doi.org/10.1017/S0266467400008312.
» http://dx.doi.org/10.1017/S0266467400008312 - NASCIMENTO, MT. and PROCTOR, J., 1997a. Soil and Plant Changes Across a Monodominant Rain Forest Boundary on Maraca Island, Roraima, Brazil. Global Ecology and Biogeography e Letters, vol. 6, no. 5, p. 387-395. http://dx.doi.org/10.2307/2997339.
» http://dx.doi.org/10.2307/2997339 - NASCIMENTO, MT. and PROCTOR, J., 1997b. Population Dynamics of Five Tree Species in a Monodominant Peltogyne Forest and Two Other Forest Types on Maraca Island, Roraima, Brazil. Forest Ecology and Management, vol. 94, no. 1-3, p. 115-128. http://dx.doi.org/10.1016/S0378-1127(96)03968-0.
» http://dx.doi.org/10.1016/S0378-1127(96)03968-0 - POTT, A. and POTT, VJ.,1994. Plantas do Pantanal. Corumbá. Brasília: EMBRAPA. 150p.
- POTT, A. and POTT, VJ.,1999. Flora do Pantanal, listagem atual de Fanerógamas. In Anais II Simpósio sobre Recursos Naturais e Socio-econômicos do Pantanal. Embrapa Pantanal, Corumbá, Brasil. p. 297-325.
- POTT, A., OLIVEIRA, AKM., DAMASCENO-JUNIOR, GA. and SILVA, JSV., 2011. Plant diversity of the Pantanal wetland. Brazilian Journal of Biology, vol. 71, no. 1, sup. Suppl 1, p. 265-273. PMid:21537599.
- POTT, VJ. and POTT, A.,2000. Plantas Aquáticas do Pantanal. Brasília: EMBRAPA. 404p.
- PRADO, AL., HECKMAN, C. and MARTINS, FR., 1994. The seasonal succession of biotic communities in wetlands of the tropical wet-and-dry climatic zone: II. The aquatic macrophyte vegetation in the Pantanal of Mato Grosso, Brazil. International Revue ges. Hydrobiology, vol. 79, p. 569-589.
- REBELLATO, L. and CUNHA, CN., 2005. Efeito do “fluxo sazonal mínimo da inundação” sobre a composição e estrutura de um campo inundável no Pantanal de Poconé, MT, Brasil. Acta Botanica Brasilica, vol. 19, no. 4, p. 789-799. http://dx.doi.org/10.1590/S0102-33062005000400015.
» http://dx.doi.org/10.1590/S0102-33062005000400015 - REBELLATO, L., NUNES DA CUNHA, C. and FIGUEIRA, JEC., 2012. Respostas da Comunidade Herbácea ao Pulso de Inundação no Pantanal de Poconé, Mato Grosso. Oecologia Australis, vol. 16, no. 4, p. 797-818. http://dx.doi.org/10.4257/oeco.2012.1604.06
» http://dx.doi.org/10.4257/oeco.2012.1604.06 - RIBEIRO, SP. and BROWN, VK., 2002. Tree Species Monodominance or Species-Rich Savannas? The Influence of Abiotic Factors in designing Plant Communities of The Brazilian Cerrado and Pantanal Matogrossense A Review. Ecotropica, vol. 8, p. 31-45.
- RICHARDS, PW.,1952. The Tropical Rain Forest. Cambridge University Press.
- SCHESSL, M., 1999. Floristic composition and structure of floodplain vegetation in the Northern Pantanal of Mato Grosso, Brazil. Phyton, vol. 39, p. 303-336.
- SCHLUTER, D. and RICKLEFS, RE.,1993. Species diversity: an introduction to the problem. In RICKLEFS, R. E. and SCHLUTER, D. Species diversity in ecological communities: historical and geographical perspectives. Chicago: University of Chicago Press , p. 1-10.
- SCREMIN-DIAS, E., LORENZ-LEMKE, AP. and OLIVEIRA, AKM., 2011. The floristic heterogeneity of the Pantanal and the occurrence of species with different adaptive strategies to water stress. Brazilian Journal of Biology, vol. 71, no. 1, sup. Suppl 1, p. 275-282. http://dx.doi.org/10.1590/S1519-69842011000200006. PMid:21537600
» http://dx.doi.org/10.1590/S1519-69842011000200006 - SILVA, IA. and BATALHA, MA., 2008. Species convergence into life-forms in a hyperseasonal cerrado in central Brazil. Brazilian Journal of Biology, vol. 68, no. 2, p. 329-339. http://dx.doi.org/10.1590/S1519-69842008000200014. PMid:18660961
» http://dx.doi.org/10.1590/S1519-69842008000200014 - SILVA JÚNIOR, MC. and FELFILI, JM.,1996. A vegetação da Estação Ecológica de Águas Emendadas. Brasília, DF: Instituto de Ecologia e Meio Ambiente do Distrito Federal. 43 p.
- SILVA, MP., MAURO, R., MOURÃO, G. and COUTINHO, M., 2000. Distribuição e quantificação de classes de vegetação do Pantanal através de levantamento aéreo. Revista Brasileira de Botânica, vol. 23, no. 2, p. 143-152.
- SOARES, JJ. and OLIVEIRA, AKM., 2009. O Paratudal do Pantanal de Miranda, Corumbá-MS, Brasil. Revista Arvore, vol. 33, no. 2, p. 339-347. http://dx.doi.org/10.1590/S0100-67622009000200015.
» http://dx.doi.org/10.1590/S0100-67622009000200015 - SPIEGEL, MP.,1976. Estatística. São Paulo: McGraw-Hill do Brasil. 580 p.
- VILLELA, DM. and PROCTOR, J., 1999. Litterfall Mass, Chemistry, and Nutrient Retranslocation in a Monodominant Forest on Maraca Island, Roraima, Brazil. Biotropica, vol. 31, no. 2, p. 198-211. http://dx.doi.org/10.1111/j.1744-7429.1999.tb00132.x.
» http://dx.doi.org/10.1111/j.1744-7429.1999.tb00132.x
Publication Dates
-
Publication in this collection
May 2014
History
-
Received
9 Aug 2012 -
Accepted
4 Feb 2013